Conservation Biology

Fred Van Dyke · Rachel L.
Lamb
Conservation
Biology
Foundations, Concepts, Applications
Third Edition
Conservation Biology
Fred Van Dyke • Rachel L. Lamb
Foundations, Concepts, Applications
Third Edition
Fred Van Dyke Rachel L. Lamb
Au Sable Institute University of Maryland
Mancelona, MI, USA College Park, MD, USA
ISBN 978-3-030-39532-2 ISBN 978-3-030-39534-6 (eBook)

https://doi.org/10.1007/978-3-030-39534-6
# Springer Nature Switzerland AG 2008, 2020

This work is subject to copyright. All rights are reserved by the Publisher, whether the whole or part of the material is
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Photo courtesy and copyright # of Rudi van Aarde. Context derived from: Mole, M.A., Rodrigues DÁraujo, S., van
Aarde, R.J., Mitchell, D., and Fuller, A., 2018. Savanna elephants maintain homeothermy under African heat. J Comp
Physiol B 188, 889–897. https://doi.org/10.1007/s00360-018-1170-5
This Springer imprint is published by the registered company Springer Nature Switzerland AG.
The registered company address is: Gewerbestrasse 11, 6330 Cham, Switzerland
Cover Photo Description
African elephants are among the world’s most charismatic megafauna and beloved by many in
the world of conservation. As the world’s largest land mammal, they transform their surround-
ings, altering ecosystems, creating habitats like water holes, and influencing plant species
composition. Once dominating the African savannah with populations close to 10 million,
elephants have dwindled to less than 500,000 individuals across the entire continent. Poaching
for ivory, habitat loss, and human-wildlife conflict have intensified, leaving conservationists
with a growing conservation problem.
This photo is drawn from the efforts of conservation scientists studying elephants in
Southern Africa. Researchers in the Okavango Delta of Botswana have been working to better
understand elephant biology and the capacity of these animals to cope with the ongoing impacts
of climate change, a new threat multiplier on the landscape. Although many elephants in this
research park, such as the one pictured in the cover photo, are accustomed to humans, these
semi-free-ranging savanna elephants offer conservationists clues to the species ability to
thermoregulate in the midst of a warming world. As the region continues to grow hotter and
drier over the next several decades, conservation solutions will depend as much on climate-
informed strategies as on traditional efforts to curb direct human disturbance.
What this photo displays arises from real science. What this photo symbolizes is the need for
real connection between people and nonhuman species. One subject is a human hand, which for
too long has been an instrument of violence in an abusive relationship toward nature globally.
The other is an elephant’s forehead. We cannot know what an elephant thinks, but we can learn
what an elephant needs. And to meet that need, it is not enough to be informed by science.
Conservation will not succeed unless we also touch the Earth’s creatures with compassion. Just
as conservationists must be instructed by science, they must also be directed by benevolence.
We have written this book for readers who desire both. Without such connection, which this
photo visually displays and conceptually symbolizes, there can be no outcome of lasting benefit
to biodiversity.
If there is hope for success in conservation biology, it requires effort aided by a new
generation of conservation scientists, managers, policy-makers, ethicists, and activists who
continue the work of the discipline’s founders while, at the same time, renovating contemporary
conservation practice with new transdisciplinary strategies that fully engage the growing
challenges of the Anthropocene. We offer this book as an invitation to the reader, whatever
your background and wherever in the world you may be, to learn about this effort and to join in
it. The work of conservation remains important not only for the African elephant but for the tens
of thousands of global species threatened by extinction. But humans are not helpless. There is
still much that we can do and time to do it.
v
Photo courtesy and copyright # of Rudi van Aarde. Context derived from: Mole, M.A., Rodrigues DÁraujo, S.,
van Aarde, R.J., Mitchell, D., and Fuller, A. 2018. Savanna elephants maintain homeothermy under African
heat. Journal of Comparative Physiology B 188, 889–897. https://doi.org/10.1007/s00360-018-1170-5
vii
Foreword
The first and second editions of this book have been great tools for conservation, but this new
edition is even more important and vital today to the living world. In the 11 year interval since
the last edition of this book was published, many things have changed for the better and for the
worse. Conservation has new methods and equipment, but the environmental situation has
unfortunately gotten much worse. On the one hand, molecular biology has developed as a vital
part of conservation because of the greater genetic and evolutionary understanding that it brings
to planning what species to save. This makes the updated chapter here on conservation genetics
particularly important. On the other hand, climate change has accelerated unbelievably and is
now a serious threat to biodiversity. Together with habitat destruction, climate change has
become a major cause of species extinction. As I write this in 2019, the Amazon forest is
burning furiously, and this makes me more aware of the need for conservation action and for the
recruitment of young conservation scientists who will learn from all the useful material in this
book. The first edition of this book was very US-oriented, but this edition is truly international
and important for conservationists wherever in the world they are operating.
Much of the deforestation and burning is for short-term profit with little thought for the
future or care about the effects on local and indigenous peoples. Profit rather than preservation
is the trend today in many places. This makes this treatment of conservation extremely
important because it addresses both practical action in the field and the moral, philosophical,
religious, legal, and ethical aspects of the situation. If the political, legal, and ethical issues are
not addressed, we will not win the battle to make the considerable changes needed to conserve a
major proportion of the world’s biodiversity or to have a sustainable future for humankind.
A new concept introduced to this edition is the Anthropocene epoch and all the implications
of it for conservation and sustainable management. Conservation today must not be confined to
virgin, untouched ecosystems, but if we want to maintain the ecosystem services upon which
the whole world and humanity itself depend, we must manage and conserve the novel
ecosystems of anthropogenic origin and even the urban environment that is discussed here in
some detail.
I like this book because of its comprehensive nature, covering so many topics of relevance to
conservation. Each chapter has been reviewed by experts in the area who have ensured that it is
presenting accurate and up-to-date information. I am impressed by the way that so much new
and recent information of importance to conservation from the last decade has been added to
this edition. The abundant illustrations and tables continue to make the information easily
understandable and accessible. It is a book that challenges one to think through the many points
of engagement and questions that are asked along the way. I know that this volume will help
young conservationists to prepare themselves for a career that is vital for the future of our
planet, but it will continue to be a useful tool for all who are working with species conservation.
Ghillean Prance, F. R. S.
ix
Preface
Personal Reflection: Origins of the Third Edition – I (Fred Van Dyke) can still recall my
surprise when I received a phone call several years ago by an editor at Springer Science and
Business Media. She said something like, “Your book [Conservation Biology: Foundations,
Concepts, Applications, 2nd Edition] has done well [for a “small market” offering]. Would you
consider writing a third edition?” The “considering” took some time, but the project was
launched several months later in the fall of 2016.
Realizing that the science and practice of conservation had changed a great deal since the
second edition was published in 2008, especially in the growth in importance of the social
sciences in conservation, I knew I would need help, particularly in matters related to things like
biogeography, climate change, environmental law and policy, and economics. I enlisted the aid
of a younger but exceptionally able colleague in Rachel Lamb, PhD Candidate and Flagship
Fellow of the University of Maryland, College Park. I knew that Rachel had already packed a
great deal of study and experience into the decade she had been active in conservation, serving
and working as, among other things, a Greater Research Opportunities Fellow for the US
Environmental Protection Agency, working in cases of environmental compliance for the Texas
(USA) Commission on Environmental Quality, developing human-integrated conservation
projects alongside A Rocha Peru, and interfacing directly with and between conservationists
and policy-makers through her work at the University of Maryland. And if she didn’t know all
that she was getting into when I asked her to help me, that was just as well, or she might not
have agreed to be a coauthor.
“I” will now become “we” for the remainder of this Preface, as the rest of what you read
here, and the things in the third edition it describes, are the product of our combined efforts and
insights. As in the previous effort, we see conservation as human action, usually in the form of
deliberate intervention, to prevent biodiversity loss. Conservation is informed by science (with
“science” here having a broad universe) to remedy deficiencies in a species’ environment,
almost entirely caused by humans, which now threaten its persistence. But although conserva-
tion must be informed by science, it can succeed only through changes in human behavior.
Therefore, conservation must address those things which most influence such behavior, includ-
ing the effects of ethics and values, environmental law and policy, and socioeconomic
constraints. Such an understanding of conservation, which will be manifested in this third
edition as it was in the second, has important implications, not only regarding what subjects to
include but how to present the subjects that are included.
A Comprehensive View of Conservation: How This Book Is Organized – Conservation is a
human activity that begins with an end in mind, and its desired outcomes are not morally neutral
objectives. Conservation sees biodiversity as “good” and therefore aims to create conditions for
biodiversity to not only be preserved but to flourish. It treats such ends not merely as scientifi-
cally predictable outcomes but as desirable moral goals. Because conservation has these ends, it
depends on science, especially biology, to understand the needs of nonhuman species and the
conditions under which they can attain stable, self-sustaining populations in equally stable and
sustainable environments. But knowing that the virtues of conservation, beginning with the
preservation of biodiversity, cannot be practiced without skill and that the skills needed to
xi
xii Preface
preserve biodiversity must be understood and applied at all levels of life, this third edition, like
its predecessors, makes the scientific aspects of conservation of genetic diversity, populations,
habitats, and ecosystems its core content (Chaps. 5, 6, 7, 8, and 9). And, as any good physical
trainer will tell you in the gym, you’re only as strong as your core.
But core strength alone is not enough, just as science is not enough, in itself, to accomplish
conservation goals. Therefore, this third edition “surrounds” the core biology of conservation
with those things necessary to complete its intentions. The first four chapters provide an
essential context for conservation to be rightly understood and effectively practiced. First,
conservation must be understood for what it is and how it came to be. Such understanding
requires a grasp of historical context and a knowledge of the “story” of conservation, and that is
where we begin in Chap. 1. From that context arises conservation’s most important subject and
the focus of its mission – the study and preservation of biodiversity. Thus, biodiversity follows
as the subject of its own chapter (Chap. 2) to establish what biodiversity is, how it is measured
and valued, its current status on Earth, and the strategies being employed to preserve it.
Following this examination of biodiversity is the one and only entirely new chapter of this
third edition – The Anthropocene: Conservation in a Human-Dominated Nature. Since the
publication of the second edition in 2008, the world human population has increased from 6.7
billion to nearly 8 billion. It has also become more than 50% urban. The majority of the world’s
ice-free lands are now strongly influenced, if not completely dominated, by intensive human
activity. Humans have become much more than an “ecological disturbance” which conserva-
tionists are obliged to “mitigate.” They are today a geophysical, biological, and climatological
force, with effects far greater than “natural” processes. Humans are creators of new ecosystems,
worldwide conveyors of other species, and harvesters of 40% of the Earth’s primary produc-
tion. Such influences, with the increasing recognition of their effects, profoundly impact
conservation actions and strategies now and far into the future. Chap. 3 examines the
implications of a human-dominated Earth for continued conservation practice. Chapter 4
follows with an examination of what we, the authors, consider the most powerful and pervasive
of the current array of anthropogenic events – global climate change – as we did in the second
edition but now with new studies and applications. Climate planning must now be a founda-
tional component of conservation planning.
Following examination of the core sciences of conservation biology (Chaps. 5, 6, 7, 8, and
9), the next three chapters take up the social, economic, and legal aspects of conservation
through examination of conservation ethics (Chap. 10), economics and development
(Chap. 11), and law and policy (Chap. 12). Not everyone can or will make a living as a
conservationist, but we believe everyone can and should live a life of conservation. However,
without the full integration of ethics, law, policy, and economics into conservation efforts,
particularly in ongoing conversations around “sustainable development,” this is not possible.
And without these disciplines of ethics and social science to complete its work, biological
science does not, on its own, achieve conservation objectives. Only when conservation fully
includes ethical and moral perception, recognized legal statutes and policy directives, and
careful consideration of economic development and sustainability can there be any hope that
desired outcomes of conservation programs will become real attainments that preserve
biodiversity.
The final chapter (Chap. 13) takes the work of conservation to a personal level – conserva-
tion as vocation. As in the second edition, we explore the ways and means through which one
can enter a career in conservation (through both traditional and increasingly diverse nontradi-
tional paths), how to understand and prepare for work and career in conservation, how to
function effectively as a contributor in a conservation effort, and, over time, how to develop the
traits and skills needed to lead such efforts.
Some Things Old and Some Things New – Like the first and second editions that preceded it,
this third edition reflects our conviction that conservation biology should be taught as a unity of
Preface xiii
thought and practice expressed through a coherent foundation of concepts, theories, facts, and
values, not as a loose assemblage of compartmentalized disciplinary expertise. Thus, we do not
attempt to present every subject that comprises conservation science today (which has already
become an impossible task for any single text), but instead define the context and relationships
of controlling ideas, problems, and applications that we, through our own study and experience
as well as that of others, actually make conservation work. We also remain convinced that
effective learners – those who display high, long-term retention rates of what they have
studied – learn highly organized information. Thus, we have continued to give careful attention
to the organization of ideas in every chapter, and among chapters, so that the organization of
ideas is itself part of the way that readers will understand the concepts presented. It is the
connections of these ideas to one another that matter as much as the ideas themselves.
We have continued our effort, begun in the second edition, to present examples and case
histories from throughout the world to define and display the thought and practice of today’s
global conservation effort and in every chapter highlight some of these case histories in new
Information Boxes. We also have worked to present and engage different intellectual paradigms
and perspectives, not shying away from controversy and disagreement when they are part of the
current development of conservation in the present age. To stimulate the discussions necessary
to process contrasting ideas, we also include, in each chapter, “Points of Engagement”
questions on subjects that students and other readers will become conversant with in particular
chapters. We continue to offer the well-received “Synthesis” sections at the end of every
chapter. These are not intended to provide the “final word” on any subject but to show some
paths of resolution and future orientation to the difficulties and challenges inherent in all aspects
of conservation study and practice.
In the second edition, every illustration was black and white. We are delighted, in this third
edition, to present many of our illustrations (most photos and some figures) in glorious color!
The biodiversity of the Earth is not only compellingly interesting, it is vividly beautiful.
Pictures are only imperfect representations of it, but those representations should be as accurate
and as attractive as possible. The biodiversity we study as conservationists deserves no less.
Practical Considerations – For those teaching a one-semester course under a typical 15–16-
week schedule, we would suggest an average of 3–4 class sessions for each of the book’s
13 chapters (39–42 of 45–48 class periods in a semester), recognizing that there will always be
sessions consumed by things like class orientation, exams, course evaluations, guest speakers,
scheduled breaks and days off, and other events that need their own place in the class schedule
and course syllabus. But if addressing each chapter in three or four sessions, we suggest a
logical order of, first, emphasis on the foundational ideas of the chapter’s subject related to the
problems unique to that subject; second, explanation of the core scientific concepts and theories
that bring meaning to the great diversity of information and data that surround the subject; and,
finally, effective applications of these concepts and theories and the outcomes they have
produced.
Every chapter is informed by many sources (typically over a hundred are cited in each one),
and we suggest that those papers or other sources the instructor finds most interesting,
illuminating, or constructively controversial be assigned for reading and discussion as the
chapter is covered. Learning how to effectively read and use the literature of conservation is
part of the work of conservation. We encourage you to equip your students for this work and
hope that our work in the literature review of each chapter will help you do it.
We hope you will find this book a useful tool, whether you are engaged in conservation
management, research, or teaching, a student in a conservation course, or simply a reader who
wants to learn more about some of the things that conservation scientists think and do. And we
welcome your feedback on how to make our book better suited to these purposes. Despite the
many setbacks, disappointments, and, sometimes, tragedies that mark conservation effort
today, we should not only acknowledge but be encouraged that, worldwide, the conservation
of biodiversity is today not only a matter of scientific study but a social and moral goal of entire
xiv Preface
communities, societies, and nations. This is a new event in human development and engages
humanity in a great and noble task. This book is one of our attempts to contribute to it. We
invite all readers and users of this book to join us in the adventure of this effort.
Mancelona, MI, USA Fred Van Dyke

College Park, MD, USA Rachel L. Lamb
About the Authors
Fred Van Dyke is Executive Director of the Au Sable Institute, an environmental and
conservation education and research organization located in northern Michigan, USA. Prior
to coming to Au Sable, Fred served as Professor and Chair of the Biology Department at
Wheaton College, Wheaton, IL, USA, and Director of Wheaton’s Environmental Studies
Program. He received his PhD in Environmental and Forest Biology from the College of
Environmental Science and Forestry, State University of New York--Syracuse. He has served
as a Wildlife Biologist for the Montana Department of Fish, Wildlife and Parks, a Scientific and
Management Consultant to the US National Park Service, and an Ex Officio Member of
numerous interdisciplinary management teams of the US Forest Service. His studies of wildlife
ecology, plant ecology, restoration ecology, fire ecology, and plant and animal response to
environmental disturbance have been published in numerous international scientific journals
and books. He is the author of the widely and internationally used textbook Conservation
Biology: Foundations, Concepts, Applications, second edition (Springer 2008).
Contact email: [email protected]
Rachel L. Lamb is a University Flagship Fellow, Harvey Fellow, and PhD Candidate in
Geographical Sciences at the University of Maryland, College Park (UMD). She also holds a
Master of Public Policy and Master of Science in Sustainable Development and Conservation
Biology from UMD. She has worked for numerous agencies and organizations, including the
Texas Commission on Environmental Quality, A Rocha Peru, National Socio-Environmental
Synthesis Center (SESYNC), Society for Conservation Biology, and the US Environmental
Protection Agency. Her current work focuses on the socioeconomic applications of NASA
Carbon Monitoring System products to advance climate-smart land use with benefits for
biodiversity. During the summers, she serves as a member of the faculty of the Au Sable
Institute (Michigan, USA) as an Assistant Professor, teaching courses in environmental law and
policy as well as land use and land resources policy. In 2015, she was named a White House
Champion of Change by the Obama Administration for her efforts in protecting the environ-
ment and human communities from effects of climate change.
Contact email: [email protected]
xv
Acknowledgments
Writing acknowledgments is a necessary pleasure for authors, an opportunity to say the

obvious – “we couldn’t have done it without you!” And there are a great many “yous” with
whom we could not have done without. We begin with Springer Associate Editor, Marlene
Moore, our faithful and perseverant Springer Science and Business Media Editor. Earlier in the
project, we were also immensely helped by Springer Associate Editor, Takeesha Moerland-
Torpey. Publishing Editor Izabella Witkowska started the project with us. At the end, our hard
work in writing was supported by the even harder work of obtaining permissions, contacting
reviewers, and reworking tables and figures by our splendid Editorial and Research Assistant,
Katie DeVoss. Thanks to Katie’s diligence, professionalism, and dogged determination to track
down every photo, figure, and table, we were able to devote ourselves to writing without
distraction, at least not editorial distractions. We are grateful to all of our reviewers who gave us
their time, knowledge, experience, and expertise to enhance our understanding of all of the
topics covered in this text and to point out our errors of fact and interpretation. We are equally
grateful for their insights about teaching, letting us know, from their experience, what works in
a classroom and what does not. They made our work infinitely better, and any errors or
misjudgments that remain are entirely our fault. We are also indebted to the many scholars,
scientists, and conservation managers who provided us with so many fine photos, illustrations,
and insights that are now also a part of this text. Finally, we are grateful to our families who
encouraged us, persevered with us, and bore with our absences (in both presence and spirit)
during the many hours spent, not only at work but at home, that were needed to complete this
effort.
Fred Van Dyke
Au Sable Institute
1 October 2019
Rachel Lamb
University of Maryland, College Park
1 October 2019
xvii
List of External Reviewers
Third Edition Reviewers
Katie DeVoss, Editorial and Research Assistant
Charles Acosta, Northern Kentucky University

Steven Bouma-Prediger, Hope College
Diane Debinski, Montana State University
John Dernbach, Widener University
David Foster, Messiah College
Richard Fredrickson, Independent Consulting Biologist
Heath Garris, Covenant College
Joanna Goger, University of Maryland, College Park
Eric Gustafson, US Forest Service
George Hurtt, University of Maryland, College Park
Thomas Lovejoy, United Nations Foundation and George Mason University
Benjamin Lowe, University of Florida
Robert O’Hagan, Borneo Orangutan Survival Foundation
L. Kristen Page, Wheaton College (Illinois)
Vern Peters, The King’s University (Alberta)
Sir Ghillean Prance, Royal Botanic Gardens, Kew (Retired)
Holmes Rolston III, Colorado State University
Nadine Rorem, Wheaton College (Illinois)
Rodney Scott, Wheaton College (Illinois)
Simon Stuart, Synchronicity Earth
Steven Thomas, US National Park Service
Timothy Van Deelen, University of Wisconsin, Madison
Benjamin Van Ee, University of Puerto Rico, Mayagüez
Katie Weakland, Bethel University (Indiana)
Second Edition Reviewers
Michael J Bigelow, Illustrator and Permissions Assistant

Lauren Anderson, Illustrator and Permissions Assistant
Charles Acosta, Northern Kentucky University

Dorothy Boorse, Gordon College
Steven Bouma-Prediger, Hope College
Gregory Chandler, University of North Carolina at Wilmington
Jiquan Chen, University of Toledo
Eric Gustafson, US Forest Service
Paula Kleintjes Neff, University of Wisconsin at Eau Claire
xix
xx List of External Reviewers
Colleen Lynch, University of South Dakota

Kathi Malueg, University of Colorado at Colorado Springs
Krista Peppers, University of Central Arkansas
Bradley Reynolds, University of Tennessee at Chattanooga
Leslie Rissler, University of Alabama
Anne Russon, York University
Carolyn Hull Sieg, US Forest Service
Steven Thomas, US National Park Service
Noah Toly, Wheaton College (Illinois)
Matt White, Ohio University
Robert C. Whitmore, West Virginia University
John Wood, The King’s University (Alberta)
First Edition Reviewers
Krista Peppers, Editorial Consultant
Vickie L. Backus, Middlebury College

Lawrence S. Barden, University of North Carolina at Charlotte
David Ehrenfeld, Cook College, Rutgers University
Laura Jackson, The University of Northern Iowa
Frances C. James, Florida State University
Susan S. Kephart, Willamette University
Mark E. Knauss, Shorter College
Jonathan A. Newman, University of Oxford
Richard A. Niesenbaum, Muhlenberg College
Kristian S. Omland, Union College
Andrew G. Peterson, Columbia University
William F. Porter, State University of New York, College of Environmental Science and
Forestry
Eleanor J. Sterling, American Museum of Natural History, Center for Biodiversity and
Conservation
Stephen C. Trombulak, Middlebury College
Robert W. Yost, Indiana University – Purdue University at Indianapolis
Contents
1 The History and Distinctions of Conservation Biology . . . . . . . . . . . . . . . . . . 1

1.1 Perspectives for an Inquiry into Conservation Biology . . . . . . . . . . . . . . . 1
1.1.1 A Remarkable Meeting . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1
1.1.2 The Emergence of Conservation Biology as a Professional and
Scientific Discipline . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
1.2 The Origins of Conservation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
1.2.1 What Is “Conservation”? . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
1.2.2 Ancient Traditions of Conservation . . . . . . . . . . . . . . . . . . . . . . . 4
1.2.3 Conservation as Expression of Privilege . . . . . . . . . . . . . . . . . . . 6
1.2.4 Conservation as Right Relationship with Nature – The Arcadian
Vision . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
1.2.5 Conservation as Knowledge – The Invitation to Study and
Appreciate Nature . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
1.2.6 Conservation as Preservation of Landscape – The Washburn
Expedition Goes to Yellowstone . . . . . . . . . . . . . . . . . . . . . . . . . 9
1.3 Foundations and History of Conservation in the United States . . . . . . . . . . 11
1.3.1 Conservation as Moral Mission – John Muir and Theodore
Roosevelt . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
1.3.2 Conservation as Utilitarian Purpose – Gifford Pinchot and
Sustainable Yield . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
1.4 Aldo Leopold and the Formation of the “Wilderness Ideal” . . . . . . . . . . . . 16
1.5 The Emergence of Global Conservation . . . . . . . . . . . . . . . . . . . . . . . . . . 17
1.5.1 Multilateral Treaties – The Beginnings of International
Conservation Efforts . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17
1.5.2 Forums for International Conservation – The UN and
the IUCN . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19
1.5.3 New Expressions of Resource Management, National Parks and
Nature Preserves . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21
1.5.4 Conservation as Preservation of Culture and Livelihood – The
Extractive Reserve . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22
1.5.5 Indigenous People, Integrated Development, and Conservation
Concern . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24
1.6 Return to Start: What Is the Place of Conservation Biology in the World
Conservation Effort? . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25
1.6.1 The Emergence of Conservation Biology from the Applied
Sciences . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25
1.6.2 The Intellectual Inception of Conservation Biology . . . . . . . . . . . 28
1.6.3 A Time of Transition: Protecting Nature from People to Protecting
Nature for People . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30
Literature Cited . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33
xxi
xxii Contents
2 Biodiversity: Concept, Measurement, and Management . . . . . . . . . . . . . . . . . 35

2.1 Biodiversity and Conservation Biology . . . . . . . . . . . . . . . . . . . . . . . . . . 35
2.2 Biodiversity and Ecosystem Function . . . . . . . . . . . . . . . . . . . . . . . . . . . . 36
2.3 Is Conservation Effort Saving Biodiversity? . . . . . . . . . . . . . . . . . . . . . . . 39
2.3.1 Conservation Governance – The IUCN and Global Biodiversity
Conservation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39
2.3.2 The Current Status of Species Biodiversity . . . . . . . . . . . . . . . . . 40
2.3.3 What Causes Biodiversity Loss? . . . . . . . . . . . . . . . . . . . . . . . . . 42
2.4 The Problem of Concept: What Is Biodiversity? . . . . . . . . . . . . . . . . . . . . 44
2.4.1 A Conceptual Definition of Biodiversity . . . . . . . . . . . . . . . . . . . 44
2.4.2 Biodiversity and the Definition of Species . . . . . . . . . . . . . . . . . . 45
2.4.3 The Species Concept in Conservation . . . . . . . . . . . . . . . . . . . . . 47
2.5 How Do We Measure the Earth’s Biodiversity? . . . . . . . . . . . . . . . . . . . . 48
2.5.1 What Biodiversity Measurements Tell Us . . . . . . . . . . . . . . . . . . 48
2.5.2 Interrelationships of Alpha, Beta, and Gamma Diversity . . . . . . . . 51
2.6 Rarity and Diversity . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 51
2.6.1 It is Common to be Rare . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 51
2.6.2 Habitat Generalists Versus Habitat Specialists . . . . . . . . . . . . . . . 52
2.6.3 Large Populations Versus Small Populations . . . . . . . . . . . . . . . . 53
2.6.4 Widespread Distribution Versus Restricted Distribution . . . . . . . . 54
2.6.5 Conserving Endemic Species . . . . . . . . . . . . . . . . . . . . . . . . . . . 57
2.7 The Problem of Distribution: Where Is Biodiversity Found? . . . . . . . . . . . 57
2.7.1 Global Patterns of Biodiversity Distribution . . . . . . . . . . . . . . . . . 57
2.7.2 Identifying Key Biodiversity Areas – Conservation with
Incomplete Data . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 59
2.8 Preserving and Managing Biodiversity . . . . . . . . . . . . . . . . . . . . . . . . . . . 60
2.8.1 Past Approaches to Conservation Management: Conservation
Legislation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 60
2.8.2 Protected Areas . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 61
2.8.3 Biodiversity Conservation, Landscape Conservation, and Human
Development . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 61
2.8.4 Urban Biodiversity Conservation . . . . . . . . . . . . . . . . . . . . . . . . 64
2.8.5 Biodiversity Technology: Finding Areas of Conservation Value
Using Remotely Sensed Data . . . . . . . . . . . . . . . . . . . . . . . . . . . 65
2.8.6 Should Management of Biodiversity be Species-Based or
Ecosystem-Based? . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 68
2.9 Better Indicators for Biodiversity Conservation . . . . . . . . . . . . . . . . . . . . . 68
2.9.1 The Value of Taxon-specific Surrogates . . . . . . . . . . . . . . . . . . . 68
2.9.2 Can Taxon Surrogates Analyze Global Patterns
of Biodiversity? . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 69
2.10 How Do We Prioritize Areas for Biodiversity? . . . . . . . . . . . . . . . . . . . . . 70
2.10.1 Current Global Prioritization Strategies . . . . . . . . . . . . . . . . . . . . 70
2.10.2 Developing More Advanced Integrated Global Conservation
Strategies . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 73
2.10.3 Management Approaches to Biodiversity at Landscape Levels . . . 74
2.10.4 Regional Biodiversity – Defining Functional Conservation
Areas . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 75
3 The Anthropocene: Conservation in a Human-Dominated Nature . . . . . . . . . 81
3.1 Dawn of the Anthropocene: Human Impacts Define a Geologic Epoch . . . . 81
3.1.1 Scientists Cast a Vote . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 81
3.1.2 The Broader Debate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 82
3.1.3 The Anthropogenic Biome . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 84
Contents xxiii
3.2 Understanding and Managing the Novel Ecosystem . . . . . . . . . . . . . . . . . 88

3.2.1 Origins of Novel Ecological Associations . . . . . . . . . . . . . . . . . . 88
3.2.2 Are Novel Ecosystems Good or Bad? . . . . . . . . . . . . . . . . . . . . . 92
3.2.3 Managing Anthropogenic Biomes and Novel Ecosystems . . . . . . . 93
3.3 The Ecology of Non-native and Invasive Species . . . . . . . . . . . . . . . . . . . 95
3.3.1 How Do Invasive Species Affect Existing Ecosystems
and Create New Ones? . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 95
3.3.2 How Humans Move Invasive Species . . . . . . . . . . . . . . . . . . . . . 97
3.3.3 Patterns and Characteristics of Successful Invasions
and Invaders . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 98
3.3.4 Understanding Invasive Processes . . . . . . . . . . . . . . . . . . . . . . . . 99
3.3.5 Forming a “Theory of Invasion Biology” – Past Efforts . . . . . . . . 102
3.3.6 State of the Art: Current Theories and Management Paradigms
for Invasive Species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 104
3.4 Biodiversity Conservation in Urban Landscapes . . . . . . . . . . . . . . . . . . . . 106
3.4.1 Growth of the Urban Landscape . . . . . . . . . . . . . . . . . . . . . . . . . 106
3.4.2 Understanding Cities as Ecological Systems . . . . . . . . . . . . . . . . 107
3.4.3 Explaining Urban Ecological Responses: Traditional and
Contemporary Approaches . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 108
3.4.4 Biodiversity in Urban Landscapes . . . . . . . . . . . . . . . . . . . . . . . . 109
3.4.5 Can Urban Areas Be Managed for Biodiversity Conservation? . . . 112
3.4.6 Changing Liability to Asset: Incorporating People into Urban
Conservation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 113
3.4.7 How to Do It – Six Strategies for Conservation Practitioners in
Urban Areas . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 115
3.5 Whither the Anthropocene? What Strategy Creates a Place for Nature? . . . 117
3.5.1 The Emergence of “Neoprotectionism” . . . . . . . . . . . . . . . . . . . . 117
3.5.2 Conservation as Human and Economic Development . . . . . . . . . . 119
3.5.3 “Convivial” Conservation – Local Autonomy for Local
Benefit . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 119
3.5.4 Can Different Approaches Find Reconciliation or Resolution? . . . 121
4 Biodiversity Conservation and Climate Change . . . . . . . . . . . . . . . . . . . . . . . 125
4.1 Climate and Climate Change . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 125
4.1.1 Why Does Climate Change Threaten Biodiversity? . . . . . . . . . . . 125
4.1.2 What Is “Climate” and What Is “Climate Change”? . . . . . . . . . . . 127
4.1.3 How Is Contemporary Change Different from Past Climate
Change? . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 129
4.1.4 The Implications of Rapidly Rising CO2 . . . . . . . . . . . . . . . . . . . 130
4.2 The Global Fingerprint of Climate Change on Biodiversity . . . . . . . . . . . . 134
4.2.1 Common Ecological Responses to Current Climate Change . . . . . 134
4.2.2 Phenological Changes and Mismatched Interactions Across
Trophic Levels . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 136
4.2.3 Observed Ranges Shifts and Changes to Local Abundance . . . . . . 139
4.2.4 Increased Threat of Extinction . . . . . . . . . . . . . . . . . . . . . . . . . . 142
4.3 Foundational Tools for Assessing Future Climate Impacts . . . . . . . . . . . . . 149
4.3.1 Integrated Vulnerability Assessments for Biodiversity . . . . . . . . . 149
4.3.2 The Bioclimate Envelope: The Correlative Approach to Modeling
Climate Effects on Individual Species . . . . . . . . . . . . . . . . . . . . . 151
xxiv Contents
4.3.3 Dynamic Global Vegetation Models: Process-Based Approaches to

Modeling Species Response to Climate . . . . . . . . . . . . . . . . . . . . 154
4.3.4 Tracking Climate Velocity: Calculating the Pace of Climate
Change . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 155
4.3.5 Decision-Making with Imperfect Estimates of the Future . . . . . . . 158
4.4 Conservation Strategies in a Time of Climate Change . . . . . . . . . . . . . . . . 158
4.4.1 Beyond Traditional Conservation Approaches . . . . . . . . . . . . . . . 158
4.4.2 Fine- and Coarse-Filter Strategies . . . . . . . . . . . . . . . . . . . . . . . . 159
4.5 Policy Initiatives for Climate Change and Conservation . . . . . . . . . . . . . . . 164
5 Conservation Genetics . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 171
5.1 Conservation Genetics and Conservation Biology . . . . . . . . . . . . . . . . . . . 171
5.2 Bottlenecks, Inbreeding, and Population Decline . . . . . . . . . . . . . . . . . . . . 172
5.2.1 The Theoretical Foundation . . . . . . . . . . . . . . . . . . . . . . . . . . . . 172
5.2.2 Of Bottlenecks and Bison . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 173
5.3 Measuring Genetic Diversity in Populations . . . . . . . . . . . . . . . . . . . . . . . 174
5.3.1 Foundational Measures of Genetic Diversity . . . . . . . . . . . . . . . . 174
5.3.2 Loss of Genetic Diversity over Time: Bottlenecks and Genetic
Drift . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 175
5.3.3 Genetic Drift and Effective Population Size . . . . . . . . . . . . . . . . . 177
5.3.4 Bottlenecks, Small Populations and Rare Alleles . . . . . . . . . . . . . 178
5.3.5 Measuring Genetic Change with Genetic Technology . . . . . . . . . . 179
5.4 Solving the Problem of Inbreeding . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 182
5.4.1 What Do We Mean by “Inbreeding” and How Would
We Measure It? . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 182
5.4.2 The Problem of Inbreeding Depression . . . . . . . . . . . . . . . . . . . . 184
5.4.3 Measuring the Inbreeding Coefficient . . . . . . . . . . . . . . . . . . . . . 185
5.5 Can Inbreeding Cause Extinction? . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 186
5.5.1 Experiments on Inbreeding . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 186
5.5.2 Inbreeding in Wild Populations . . . . . . . . . . . . . . . . . . . . . . . . . . 186
5.5.3 Outbreeding Depression . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 188
5.6 Landscape Genetics and Habitat Fragmentation . . . . . . . . . . . . . . . . . . . . 188
5.6.1 Habitat Fragmentation: A Genetic Threat to Large and Small
Populations . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 188
5.6.2 Landscape-Induced Genetic Differentiation . . . . . . . . . . . . . . . . . 189
5.7 Managing Genetic Diversity in Wild Populations . . . . . . . . . . . . . . . . . . . 190
5.7.1 Importing Genetic Diversity: Genetic Restoration of Inbred
Populations . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 190
5.7.2 Hybridization and Introgression: The Case of the Red Wolf . . . . . 193
5.8 Managing Genetic Diversity in Captive Populations . . . . . . . . . . . . . . . . . 196
5.8.1 Measuring the Cost of Adaptation to Captivity . . . . . . . . . . . . . . . 196
5.8.2 Managed Breeding: Mitigating Effects of Inbreeding
in Captivity . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 197
5.8.3 The Okapi: Analyzing Parameters of Captive Breeding
Management . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 197
5.8.4 Managing Captive Populations to Retain Genetic Diversity . . . . . . 200
5.9 Applying Genetic Information in Conservation . . . . . . . . . . . . . . . . . . . . . 203
5.9.1 General Considerations . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 203
5.9.2 Genetic Analysis Can Clarify Relatedness, Taxonomy, and
Phylogeny . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 203
5.9.3 Genetic Analysis Can Define Management Units of Fragmented
Populations . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 203
Contents xxv
5.9.4 Genetic Analysis Can Determine Rates of Gene Flow Among

Populations . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 204
5.9.5 Genetic Analysis Can Expose Exploitation of Protected
Species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 205
Appendix 5.1: A Summary of Mathematical Equations Used in Common
Estimations of Genetic Diversity and Effective Population Size . . . . . . . . . . . . . . 207
6 The Conservation of Populations: Theory, Analysis, Application . . . . . . . . . . 211
6.1 Defining Populations and Population Processes . . . . . . . . . . . . . . . . . . . . . 211
6.1.1 What Is a Population? . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 211
6.1.2 Population Demography . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 212
6.1.3 Stochastic Perturbations – Density-Independent Factors of
Population Growth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 214
6.1.4 Density Dependent Population Regulation . . . . . . . . . . . . . . . . . . 217
6.2 Populations and Metapopulations: Complexities of Population
Subdivision and Fragmentation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 220
6.2.1 Origins of Metapopulation Theory . . . . . . . . . . . . . . . . . . . . . . . 220
6.2.2 The Definition and Development of Metapopulation Concepts . . . 221
6.2.3 Does Metapopulation Theory Predict Behavior of Real
Populations? The Case of the Growling Grass Frog . . . . . . . . . . . 223
6.3 Detecting Populations for Conservation Management . . . . . . . . . . . . . . . . 224
6.3.1 The Problem of Detection . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 224
6.3.2 Occupancy Theory and Modeling . . . . . . . . . . . . . . . . . . . . . . . . 229
6.3.3 Developing Technology and Applications in Occupancy
Modeling . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 230
6.4 Minimum Viable Populations . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 233
6.4.1 General Considerations . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 233
6.4.2 Trend Analysis and Factor Resolution: Systematic Approaches
for Identifying Causes of Population Decline and Strategies
for Restoration . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 233
6.4.3 Saving a Population from Extinction: The Case
of the Black-Footed Ferret . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 236
6.5 Population Viability Analysis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 240
6.5.1 Conceptual Foundations . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 240
6.5.2 Developing a Conservation PVA– The Western Prairie
Fringed Orchid . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 242
6.5.3 Incorporating Stochasticity . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 246
6.5.4 Evaluating Elasticity . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 247
6.5.5 Applications for Animal Populations – Bonelli’s Eagle
in Western Europe . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 247
6.5.6 Evaluating a PVA . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 247
6.6 Applying PVA Results in Conservation Management . . . . . . . . . . . . . . . . 249
6.7 From Population Viability Analysis to Population Viability
Management . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 251
6.8 The Problem of Recovery: Protecting Conservation – Reliant Species . . . . 252
6.8.1 What Is a Conservation–Reliant Species? . . . . . . . . . . . . . . . . . . 252
6.8.2 The Kirtland’s Warbler: A “Success Story” of Conservation-
Reliance . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 253
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7 The Conservation of Terrestrial Habitat and Landscape . . . . . . . . . . . . . . . . 261

7.1 A Foundational Understanding of Habitat . . . . . . . . . . . . . . . . . . . . . . . . . 261
7.1.1 What Is Habitat? . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 261
7.1.2 How Do We Measure Habitat Use? . . . . . . . . . . . . . . . . . . . . . . . 262
7.2 Heterogeneity, Landscape Gradients and Patch Dynamics . . . . . . . . . . . . . 266
7.2.1 Habitat Heterogeneity, Gradients, and Patchiness . . . . . . . . . . . . . 266
7.2.2 Habitats and Landscapes: Measuring Scales of Space
and Time . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 267
7.2.3 How Do We Predict Habitat Change? . . . . . . . . . . . . . . . . . . . . . 268
7.3 Dimensions of Destruction: Understanding Habitat Loss, Fragmentation,
Isolation and Degradation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 271
7.3.1 Defining Our Terms . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 271
7.3.2 Isolating Consequences of Habitat Fragmentation and Effects
of Edge . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 273
7.3.3 Habitat Alteration Through Effects of Edge: First Principles . . . . . 274
7.3.4 Environmental Characteristics of Edges . . . . . . . . . . . . . . . . . . . . 275
7.3.5 What Lies Between? Managing Matrix Habitat . . . . . . . . . . . . . . 277
7.3.6 Loss and Fragmentation: Experimental Isolation of Separate
Effects . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 278
7.3.7 A Larger Perspective: Long-Term Studies of Habitat Loss and
Fragmentation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 280
7.4 Theories and Models of Loss and Fragmentation . . . . . . . . . . . . . . . . . . . 281
7.4.1 Neutral Landscape Models . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 281
7.4.2 Percolation Theory: Defining the Critical Threshold of
Fragmentation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 282
7.4.3 Can Percolation Theory Explain the Real World? Models
and Field Studies . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 284
7.5 Conservation Through Protected Areas . . . . . . . . . . . . . . . . . . . . . . . . . . . 289
7.5.1 Algorithms of Reserve Design . . . . . . . . . . . . . . . . . . . . . . . . . . 289
7.5.2 GAP Analysis and Reserve Design . . . . . . . . . . . . . . . . . . . . . . . 291
7.5.3 Reserve Design and Habitat Suitability . . . . . . . . . . . . . . . . . . . . 291
7.5.4 Determining Appropriate Reserve Size . . . . . . . . . . . . . . . . . . . . 292
7.6 Preserving Habitats in Human-Modified Landscapes . . . . . . . . . . . . . . . . . 295
7.6.1 Intermediate Disturbance: Where Does Conservation
Matter Most? . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 295
7.6.2 Conservation in Agricultural and Commercially Forested
Landscapes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 295
7.6.3 Mitigation in Forest Environments . . . . . . . . . . . . . . . . . . . . . . . 298
7.6.4 Mitigating Human Effects to Avoid Habitat Loss and Range
Displacement: The Case of the Line Creek Elk . . . . . . . . . . . . . . 298
8 The Conservation of Aquatic Systems . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 307
8.1 Conservation Challenges of Aquatic Habitats . . . . . . . . . . . . . . . . . . . . . . 307
8.1.1 Reservoirs of Global Biodiversity . . . . . . . . . . . . . . . . . . . . . . . . 307
8.1.2 Basic Properties of Aquatic Environments . . . . . . . . . . . . . . . . . . 308
8.1.3 Threats to Freshwater Ecosystems . . . . . . . . . . . . . . . . . . . . . . . . 309
8.1.4 Chemical and Biological Degradation . . . . . . . . . . . . . . . . . . . . . 311
8.1.5 Dams, Levees, and Flood Plains: Flow, Impoundments, and
Connectivity . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 312
8.1.6 Consequences of Dams on Fish Biodiversity and Community
Composition . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 313
Contents xxvii
8.1.7 Dams as Barriers to Population Persistence and Reproduction . . . . 316

8.2 Management of Freshwater Habitats for Conservation . . . . . . . . . . . . . . . . 317
8.2.1 Managing Chemical and Physical Inputs to Aquatic Systems . . . . 317
8.2.2 Lake Systems as Alternative Stable States . . . . . . . . . . . . . . . . . . 318
8.3 Managing Freshwater Systems at Landscape Levels . . . . . . . . . . . . . . . . . 319
8.3.1 Protected Areas for Freshwater Systems . . . . . . . . . . . . . . . . . . . 319
8.3.2 Coarse-Filter Approaches for Regional Representation . . . . . . . . . 320
8.4 Wetlands, Pools and Ponds . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 322
8.4.1 What Are Wetlands? . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 322
8.4.2 Managing Wetlands for Conservation . . . . . . . . . . . . . . . . . . . . . 324
8.4.3 The Special Case of Forest Pools: Critical Elements
for Amphibian Biodiversity . . . . . . . . . . . . . . . . . . . . . . . . . . . . 325
8.4.4 Engagement of Legislators and Stakeholders in Forest Pool
Conservation: A US Case History . . . . . . . . . . . . . . . . . . . . . . . . 326
8.4.5 Big Impacts of Small Habitat: Pond Biodiversity . . . . . . . . . . . . . 326
8.5 Policies and Practices that Protect Freshwater Habitats . . . . . . . . . . . . . . . 328
8.5.1 Connecting Stakeholders, Scientists, and Policy Makers . . . . . . . . 328
8.5.2 Forming an Issue-Driven Coalition: The Healthy Waterways
Partnership . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 329
8.6 Marine Habitats and Biodiversity . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 332
8.6.1 Destruction of Benthic Environments . . . . . . . . . . . . . . . . . . . . . 332
8.6.2 Pollution in the Water Column . . . . . . . . . . . . . . . . . . . . . . . . . . 333
8.6.3 Habitat Destruction in Shallow Water Environments – The Plight
of Seagrass . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 335
8.6.4 Habitat Destruction and Marine Biodiversity:
Threats to Coral Reefs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 337
8.6.5 Rehabilitation Techniques for Coral Reefs . . . . . . . . . . . . . . . . . . 340
8.7 Overexploitation of Marine Populations . . . . . . . . . . . . . . . . . . . . . . . . . . 343
8.7.1 The Collapse of Marine Fisheries . . . . . . . . . . . . . . . . . . . . . . . . 343
8.7.2 The Surplus-Yield Theory: Great Whales and Ecological
Function . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 346
8.7.3 Bycatch: The Preeminent Threat to Large Marine Vertebrates . . . . 347
8.7.4 Reducing and Mitigating Effects of Bycatch . . . . . . . . . . . . . . . . 349
8.8 Preserving Marine Habitats and Biodiversity through Protected Areas:
The Marine Reserve . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 349
8.8.1 Management Context, Goals and Strategies in Marine
Reserves . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 349
8.8.2 Protection at Ecosystem Levels: Australia’s Great Barrier Reef
Marine Park . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 350
8.8.3 Co-management – Can Shared Authority Provide Better
Conservation? . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 351
8.8.4 Marine Protected Areas and Commercial Fisheries . . . . . . . . . . . . 353
9 Conservation Through Ecosystem Management . . . . . . . . . . . . . . . . . . . . . . . 359
9.1 The Concept of Ecosystem Management . . . . . . . . . . . . . . . . . . . . . . . . . 359
9.1.1 Resource Management and Ecosystem Management . . . . . . . . . . . 359
9.1.2 How the Spotted Owl Started Ecosystem Management . . . . . . . . . 362
9.1.3 The Modern Context: EBM in Contemporary Conservation . . . . . 364
9.2 Shaping Decision-Making Processes in EBM . . . . . . . . . . . . . . . . . . . . . . 367
9.2.1 Criteria that Define Ecosystem-Based Management . . . . . . . . . . . 367
9.2.2 The Role of Adaptive Management . . . . . . . . . . . . . . . . . . . . . . . 370
xxviii Contents
9.2.3 Evaluating Ecosystem-Based Management as a

Performance-Based System . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 373
9.3 Scientific Foundations of Ecosystem-Based Management . . . . . . . . . . . . . 377
9.3.1 The Problem of Location – Where Is the Ecosystem? . . . . . . . . . . 377
9.3.2 Do Protected Areas Protect Ecosystems? . . . . . . . . . . . . . . . . . . . 377
9.3.3 Using Watersheds to Define Ecosystem Limits, Boundaries,
and Processes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 379
9.3.4 Knowing the System– What Data Should Be Collected
for EBM? . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 380
9.4 Implementing Management Decisions –Tools of Ecosystem
Management . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 388
9.4.1 Ecosystem Modeling . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 388
9.4.2 Managing Ecosystem Processes . . . . . . . . . . . . . . . . . . . . . . . . . 391
9.4.3 Managing Nature’s Ecosystem Engineers: Herbivores and
Herbivory . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 393
9.5 Creating and Managing Governance Systems of Ecosystem-Based
Management . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 396
9.5.1 Managing Through Collaboration with Stakeholders . . . . . . . . . . 396
9.5.2 Linking Interest with Identification: Australia’s Tully-Murray
Watershed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 396
9.5.3 Costs of Bad Governance: Managing Time from
Recommendation to Action . . . . . . . . . . . . . . . . . . . . . . . . . . . . 400
9.5.4 Creating a Working Framework for Ecosystem-Based Governance:
The Case of the Great Barrier Reef . . . . . . . . . . . . . . . . . . . . . . . 402
9.5.5 Relational Governance: Managing Stakeholder Interactions by
Building Trust . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 403
9.5.6 Stakeholders as Managers: Ecosystem Management from the
Bottom Up . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 406
10 Values and Ethics in Conservation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 411
10.1 Does Conservation Science Need Conservation Ethics? . . . . . . . . . . . . . . . 411
10.1.1 Conservation Biology – Regulatory Science or Value-Laden
Mission? . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 411
10.1.2 Value – Property of Nature or Product of Thought? Problems
of Plastic Trees . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 412
10.2 The Necessity of Value Judgments in Conservation . . . . . . . . . . . . . . . . . 414
10.2.1 Recognizing Management Actions as Value Judgments . . . . . . . . 414
10.2.2 Values and Ethics – Foundational Definitions . . . . . . . . . . . . . . . 415
10.2.3 How Values Inform Management . . . . . . . . . . . . . . . . . . . . . . . . 417
10.3 How Do We Categorize Conservation Values? . . . . . . . . . . . . . . . . . . . . . 418
10.3.1 An Overview of Value Categories . . . . . . . . . . . . . . . . . . . . . . . . 418
10.3.2 Instrumental Values . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 418
10.4 Moral Value: Assigning Intrinsic Values in Conservation . . . . . . . . . . . . . 421
10.4.1 Where Does Intrinsic Value Reside? . . . . . . . . . . . . . . . . . . . . . . 421
10.4.2 Establishing Intrinsic Value Through Moral Extensionism . . . . . . 422
10.5 Conservation Value and Practice in Religious Traditions . . . . . . . . . . . . . . 428
10.5.1 Intrinsic Value in the Judeo-Christian Tradition . . . . . . . . . . . . . . 428
10.5.2 Beginning in the Middle – The Historical Roots of Our
Ecologic Crisis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 429
10.5.3 Biblical Teaching and Application in Conservation . . . . . . . . . . . 429
10.5.4 Islamic Teaching on Conservation . . . . . . . . . . . . . . . . . . . . . . . . 431
10.5.5 Conservation in Hinduism . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 431
Contents xxix
10.5.6 Conservation Teachings in Buddhism . . . . . . . . . . . . . . . . . . . . . 432

10.5.7 Indigenous Belief Systems in Conservation . . . . . . . . . . . . . . . . . 433
10.6 Practical Applications: Faith-Based Contributions to Conservation . . . . . . . 436
10.6.1 “Goal Rational” Versus “Value Rational” Conservation . . . . . . . . 436
10.6.2 Jewish and Christian FBOs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 437
10.6.3 Laudato Si – Pope Francis’s Call to Global Conservation . . . . . . . 437
10.6.4 Conservation FBOs in Islam . . . . . . . . . . . . . . . . . . . . . . . . . . . . 439
10.6.5 Conservation Activism in Hinduism . . . . . . . . . . . . . . . . . . . . . . 440
10.6.6 Conservation FBOs in Buddhism . . . . . . . . . . . . . . . . . . . . . . . . 440
10.6.7 Future Roles and Contributions of FBOs in Global
Conservation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 441
10.6.8 Saving the Cedars of Lebanon . . . . . . . . . . . . . . . . . . . . . . . . . . 441
11 Conservation Economics and Sustainable Development . . . . . . . . . . . . . . . . . 449
11.1 The Role of Economics in Conservation . . . . . . . . . . . . . . . . . . . . . . . . . . 449
11.1.1 Thinking Like an Economist . . . . . . . . . . . . . . . . . . . . . . . . . . . . 449
11.1.2 Ecosystem Services . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 450
11.1.3 Stock-Flow Resources and Fund-Service Resources . . . . . . . . . . . 451
11.1.4 Nonexcludable and Nonrival Goods . . . . . . . . . . . . . . . . . . . . . . 452
11.2 Microeconomic Approaches to Conservation Dilemmas . . . . . . . . . . . . . . 454
11.2.1 Fundamental Assumptions of Supply and Demand . . . . . . . . . . . . 454
11.2.2 The Challenge of Externalities . . . . . . . . . . . . . . . . . . . . . . . . . . 455
11.2.3 Cost Benefit Analysis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 456
11.3 Methods for Valuing Ecosystem Goods and Services . . . . . . . . . . . . . . . . 456
11.3.1 Should We Price Nature? . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 456
11.3.2 Revealed Preference Methods . . . . . . . . . . . . . . . . . . . . . . . . . . . 457
11.3.3 Stated Preference Methods . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 460
11.4 The Role of Moderating Institutions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 461
11.4.1 Institutions from Economic Perspective . . . . . . . . . . . . . . . . . . . . 461
11.4.2 Government-Market Interactions . . . . . . . . . . . . . . . . . . . . . . . . . 463
11.4.3 The Role of Property Rights in Conservation . . . . . . . . . . . . . . . . 467
11.5 Ecological Economics . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 472
11.5.1 Historical Challenges to Neoclassical Economics . . . . . . . . . . . . . 472
11.5.2 Characteristics of Ecological Economics . . . . . . . . . . . . . . . . . . . 473
11.5.3 Implications of Challenging Business-as-Usual . . . . . . . . . . . . . . 475
11.6 Broader Linkages Between Economics and Development . . . . . . . . . . . . . 478
11.6.1 The Origins of Sustainable Development . . . . . . . . . . . . . . . . . . . 478
11.6.2 Integrated Conservation and Development . . . . . . . . . . . . . . . . . . 479
12 The Legal Foundations of Conservation Biology . . . . . . . . . . . . . . . . . . . . . . . 489
12.1 Law and Policy as a Framework for Conservation . . . . . . . . . . . . . . . . . . . 489
12.1.1 Nexus Between Conservation Science and Policy . . . . . . . . . . . . . 489
12.1.2 Defining Terms: Legal Frameworks and Linkages to Policy . . . . . 491
12.2 Foundational Conservation Law in the United States . . . . . . . . . . . . . . . . . 493
12.2.1 Common Characteristics of Effective Conservation Law . . . . . . . . 493
12.2.2 The US National Environmental Policy Act (NEPA) . . . . . . . . . . 493
12.2.3 The US Endangered Species Act (ESA) . . . . . . . . . . . . . . . . . . . . 500
12.3 International Conservation Law . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 510
12.3.1 Understanding Key Terms . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 510
12.3.2 Stockholm: The Beginnings of Modern International
Conservation Law . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 512
xxx Contents
12.3.3 Protection of Endangered Species: The Convention

on International Trade in Endangered Species of Wild Fauna
and Flora (CITES) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 513
12.3.4 Combining Conservation and Development in International
Agreements . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 515
12.3.5 The Process of Creating and Enforcing International
Conservation Law . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 518
12.4 The Challenge of Interdependence on a Global Stage . . . . . . . . . . . . . . . . 521
12.4.1 The Nature of Legal Interdependence Among Nation-States . . . . . 521
12.4.2 Case History I: Tuna and Dolphins . . . . . . . . . . . . . . . . . . . . . . . 522
12.4.3 Case History II: Shrimp and Sea Turtles . . . . . . . . . . . . . . . . . . . 523
12.4.4 Case History III: Brazilian Biodiversity and Genetic Resources . . . 524
12.4.5 Outcomes and Future Prospects . . . . . . . . . . . . . . . . . . . . . . . . . 525
13 Conservation as Vocation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 531
13.1 Conservation as Vocation – First Steps . . . . . . . . . . . . . . . . . . . . . . . . . . . 531
13.1.1 Articulating Your Personal Mission in Conservation . . . . . . . . . . . 531
13.1.2 Foundational Elements of Conservation Education . . . . . . . . . . . . 532
13.1.3 Making the Transition from Student to Colleague . . . . . . . . . . . . . 533
13.2 Reaching a Wider Audience . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 536
13.2.1 A Professional Network of Contacts and References . . . . . . . . . . . 536
13.2.2 Conservation as a Social Process: Involvement in Professional
Societies . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 537
13.3 Graduate Education in Conservation Biology . . . . . . . . . . . . . . . . . . . . . . 538
13.3.1 Independent Evaluation for Graduate School – The Graduate
Record Exam . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 538
13.3.2 Choosing a Program . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 539
13.3.3 Choosing a Project, Graduate Professor and Mentor . . . . . . . . . . . 540
13.4 Innovative Educational Approaches . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 540
13.4.1 The Need for Non-traditional Education . . . . . . . . . . . . . . . . . . . 540
13.4.2 An Intentionally Interdisciplinary Approach . . . . . . . . . . . . . . . . . 541
13.4.3 Intentionally Creative Thinking – New Paths Out of Old Ruts . . . . 542
13.4.4 Interdisciplinary Study Through Program-Level Innovation . . . . . 543
13.4.5 Systemic Pathways to Creative Education . . . . . . . . . . . . . . . . . . 545
13.4.6 Relational Skills in Conservation: Learning How to Lead . . . . . . . 547
13.4.7 A Career in Conservation Social Sciences . . . . . . . . . . . . . . . . . . 548
13.5 Entering a Vocational Setting: How Do I Get a Job? . . . . . . . . . . . . . . . . . 551
13.5.1 Choose Courses for the Job, Not the Degree . . . . . . . . . . . . . . . . 551
13.5.2 Choosing a Vocational Setting – Should I Take this Job? . . . . . . . 555
13.5.3 How Can I Excel in my Work and Nurture Professional
Relationships? . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 556
13.5.4 How Do I Overcome Barriers? Inclusion and Diversity in
Conservation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 557
13.5.5 How Do I Learn to Recognize Opportunity? . . . . . . . . . . . . . . . . 560
13.6 Becoming an Effective Advocate for Conservation . . . . . . . . . . . . . . . . . . 563
13.6.1 Professional Expressions of Advocacy . . . . . . . . . . . . . . . . . . . . . 563
13.6.2 An Alternative View of Advocacy . . . . . . . . . . . . . . . . . . . . . . . 563
13.6.3 Examining Outcomes: Implications of Alternative Views of
Advocacy . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 564
13.6.4 Can Conservation Biologists Not Be Advocates for
Conservation? . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 566
Contents xxxi
13.6.5Making Advocacy Intentional – Avoiding Inadvertent

Advocacy . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 566
13.6.6 Avoiding Conflicts of Interest in Advocacy . . . . . . . . . . . . . . . . . 567
Glossary . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 571
Index . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 589
The History and Distinctions of Conservation Biology
1
Disciplines are not logical constructs; they are social crystallizations which occur when a group of people agree
that association and discourse serve their interests. Conservation biology began when a critical mass of people
agreed that they were conservation biologists.
Michael Soulé (1986:3)
Keywords 65 million years, it was time for academics and conserva-

Conservation history · Origins of conservation · Conser- tionists to join forces to save threatened and endangered
vation traditions · Washburn expedition · Resource con- species (Gibbons 1992). Soulé’s words sparked both contro-
servation ethic · Wilderness ideal · Extractive reserves · versy and criticism, but few were left unmoved. That
Conservation science · Arcadian vision · Distinctions of meeting, now ambitiously called the First International Con-
conservation biology ference on Conservation Biology, led to new beginnings. A
landmark publication that would become a foundational
statement of the new discipline’s identity, Conservation Biol-
ogy: An Evolutionary-Ecological Perspective (Soulé and
Overview Wilcox 1980) resulted. A new scientific organization, the
In this chapter you will learn about: Society for Conservation Biology, was created, and a new
discipline, conservation biology, was born.
1. The origins and history of conservation and conser- Officially, the formal organization and incorporation of
vation biology. the Society for Conservation Biology (SCB) was still years
2. The conceptual distinctions of conservation away. As Soulé noted later,
biology. For the record, the Society for Conservation Biology originated
3. The role of conservation biology in the world of about 5 PM on May 8, 1985, in Ann Arbor, Michigan [USA] at
the conclusion of the Second Conference on Conservation Biol-
conservation. ogy. An informal motion to organize such a society was approved
by acclamation, following reports by chairpersons (Jared Dia-
mond and Peter Brussard) of two ad hoc committees. Those
committees met during the conference to discuss the need for
such a society and for a journal and I was asked to help from
there. With the help of many people and organizations . . . we
1.1 Perspectives for an Inquiry into drafted a constitution and discussed matters of policy and publi-
cation (Soulé 1987:4).
Reflecting on the motives that drew the first members of
1.1.1 A Remarkable Meeting SCB together, Soulé remarked, “The Society is a response by
professionals, mostly biological and social scientists,
In 1978, a group of academic scientists, zookeepers, and managers and administrators to the biological diversity crisis
wildlife conservationists attended a banquet at the San that will reach a crescendo in the first half of the twenty-first
Diego Wild Animal Park. There biologist Michael Soulé century. We assume that we are in time, and that by joining
made an impassioned plea to his colleagues: with world together with each other and with other well-intentioned
extinction rates estimated to be at their highest levels in
# Springer Nature Switzerland AG 2020 1

F. Van Dyke, R. L. Lamb, Conservation Biology, https://doi.org/10.1007/978-3-030-39534-6_1
2 1 The History and Distinctions of Conservation Biology
persons and groups, the worst biological disaster in the last

65 million years can be averted. . .Although we have varying
philosophies, we share a faith in ourselves, as a species and as
individuals, that we are equal to the challenge. . .For these
reasons we join together in professional alliance, in the ser-
vice of each other, but also in the service of the less articulate
members of our evolutionary tree” (Soulé 1987:4–5). Two
years after the events Soulé describes, the first annual
meeting of the SCB was held at Montana State University
in Bozeman (USA) in June 1987.
There were inevitable growing pains for the new move-
ment. Soulé noted, “The ecologists and biogeographers
didn’t think geneticists had much to contribute to conserva-
tion, and the wildlife managers didn’t think that the academic
eggheads had anything relevant to say” (Gibbons 1992:20).
But, in time, this disparate, cross-disciplinary and interna-
tional group of scientists learned not only to speak to one Fig. 1.1 Percentage of scientific publications in environmental sciences
another, but to work together with mutual esteem and com- and ecology that contain the search term “conservation” between 1985
mon purpose. By 2006, membership had climbed to over and 2012. (Reprinted by permission from Springer Nature, from
Dalerum, F., 2014. Identifying the role of conservation biology for
11,000 worldwide, the discipline boasted a number of under-
solving the environmental crisis. AMBIO 43, 839–846. # 2014 Royal
graduate textbooks (Hunter 2002; Pullin 2002; Primack Swedish Academy of Sciences)
2004, 2006; Groom et al. 2006), dozens of titles on more
advanced aspects of conservation problems, and hundreds of
colleges and universities offering graduate and undergraduate many cooperative and cross-disciplinary efforts in a myriad
studies in the discipline. of scientific studies and conservation initiatives. It has
The growth of the field was, in the words of founding generated funding for research and management that now
member Stanley A. Temple, wildlife ecologist at the Univer- totals in the hundreds of millions of dollars. The Conserva-
sity of Wisconsin-Madison (USA), “incredible” (Gibbons tion Biology Institute, founded in 1997, is one such benefi-
1992). But as science journalist Ann Gibbons noted prophet- ciary that has been foundational in developing broadly used
ically, “. . .despite the burgeoning funding for conservation web-based conservation data sharing systems such as the
biology and the demand from students for more classes in the Protected Areas Database of the US (PAD-US), Data Basin,
field, the real test of the vision Michael Soulé elaborated in and Data Basin Climate Center. Despite this expansion,
1978 is yet to come. The test is not whether conservation the question remains: Will conservation biology be able to
biology can convince its academic critics that it is ‘real preserve biodiversity? That is, as Gibbons put it, “the true
science,’ or improve its diplomatic relations with traditional test.”
conservationists, or generate funding from federal and private
sources. The true test will be whether the field can actually
preserve biodiversity” (Gibbons 1992:22). 1.1.2 The Emergence of Conservation Biology
The intervening years have answered Gibbons first three as a Professional and Scientific Discipline
questions definitively. Throughout the world, in scores of
undergraduate and graduate university programs, at the Conservation biology did not emerge as a distinct and differ-
United Nations, in national and provincial government ent discipline because foresters, wildlife biologists,
agencies, in non-governmental conservation organizations ecologists, and fisheries managers decided to stop being all
and citizens groups, in dozens of academic journals and these and other things. Conservation biology was, as biolo-
professional meetings, conservation biology is unquestion- gist David Ehrenfeld noted in retrospect, “not defined by a
ably recognized as “real science” and the percentage of discipline but by its goal – to halt or repair the undeniable
scientific publications in environmental science and ecology damage that is being done to ecosystems, species, and the
that contain the search term “conservation” has grown at an relationship of humans to the environment . . .” (Ehrenfeld
unprecedented rate (Dalerum 2014) (Fig. 1.1). As conserva- 1992:1625). Conservation biology experienced remarkable
tion biology has not only improved diplomatic relations growth in numbers and influence, particularly in its early
with older, more traditional disciplines, it also has forged years, because it filled a critical gap: a common ground, not
1.2 The Origins of Conservation 3
previously in existence, where scientists concerned with 1.2 The Origins of Conservation
extinction and how to prevent it could meet, talk, and work
together, and arising out of the perceived failure of older 1.2.1 What Is “Conservation”?
disciplines to address contemporary conservation problems
(Meine et al. 2006). To understand the origins and history of conservation, one
In its contemporary context, Michael Soulé defined con- must begin by defining conservation by its goals, not its
servation biology as a “crisis discipline” whose goal was to impacts. Human beings always have affected their physical
provide principles and tools for preserving biodiversity environment and its non-human species, but a group of
(Soulé 1985). By 1970, biologist David Ehrenfeld was people living in relative equilibrium with their environment
already articulating the connection between biological are not necessarily conservationists. They may be wasteful
knowledge, political action, and ethical responsibility. but cause no damage because their environment produces
“Biologists,” said Ehrenfeld, “are beginning to forge a disci- more than they need. Or they may be inefficient, desiring,
pline in that turbulent and vital area where biology meets the but not succeeding, to take more from the environment than
social science and humanities. The need is now very great for they do. If their efficiency improves, they may do increasing
a scientifically valid presentation of the biological problems harm. In this case, their practice of “conservation” will be
that are more relevant to the life of modern man” (Ehrenfeld inversely related to their technological capabilities. This type
1970:vii). Conservation biology made such perception more of passive “conservation” will be unsustainable if the envi-
specific, more urgent, and more scientifically based. Soulé ronment becomes less productive or if the people become
expressed the urgency dramatically by saying that conserva- more efficient in exploiting it (Alvard 1993).
tion biology is to biology what surgery is to physiology or Genuine conservation can occur only when humans
war to political science. It must respond to emergency intentionally use resources at less than maximum sustainable
situations with incomplete information, and its goal is a rates or forego the use of some resources altogether. This
predetermined desirable outcome, not simply an accumula- kind of conservation is motivated by appreciating an intrinsic
tion of scientific information or an enlightened discussion value of the resource itself or from the desire to provide a
about interesting problems. Although urgent, short-term long-term supply of the resource for others, including others
goals in conservation change, the long-term goal is the per- still to come in future generations. These motivations are not
sistence and viability of functioning ecosystems (Soulé mutually exclusive, and they are primarily ethical in nature.
1985). From its inception, conservation biology understood Throughout history, human beings have shown themselves
itself as synthetic, eclectic, and multidisciplinary. It did not capable of embracing both motives, but often have lacked the
draw all its theories and models from biology and it had the will to do so. In fact, humans have shown the opposite
effect of breaking down the dichotomy of pure versus applied capacities for intensely selfish motives to get as much of a
science (Soulé 1985). As an emerging discipline, conserva- resource as they can for their own needs and pleasures. The
tion biology stressed the intrinsic value of biodiversity (the history of conservation is one of ethical conflict as well as
value of biodiversity in and of itself), not simply its utilitarian scientific discovery. Conservation has benefits for humans
values. but requires restraint and incurs costs. Conservation that
Today conservation biology is still an arguably new and involves neither restraint nor cost is not conservation. If one
distinctive discipline, but few, if any, of the scientific is careless in defining conservation, it is easy to make histor-
paradigms it employs are original, nor are the problems it ical generalizations that are not true. One false generalization
has attempted to solve. There were conservationists, is that people with little technology are good conservationists
biologists and conservation dilemmas long before there was and people with high levels of technology are not. Another is
conservation biology, and there were many individuals who that hunter-gatherer societies conserve the environment and
practiced conservation as biologists. Although conservation agricultural societies degrade it. This kind of thinking distorts
biology had an identifiable “birth” as a professional society the historical context of conservation and misguides present
and subsequently as a professional discipline in the applications of a conservation ethic.
mid-1980s, the foundational soil from which it rose is much Throughout the world, early human societies in Europe,
older and deeper. In this chapter, we will examine those Asia, Africa, Australia, and the Americas radically affected
foundations and how conservation biology germinated their physical environment and the species in it. They began
from them. To better understand the origins and distinctions to “manage” ecosystems even with little technological devel-
of the science of conservation biology, we will consider opment, primarily through the use of fire. All early peoples
where our ideas about conservation have come from, and used fire, and with this single tool changed landscapes,
how they have contributed to the formation of this “new” exterminated species, and created cultivable areas (Pyne
discipline. 2001). Even primitive cultures that were able to employ
effective combinations of fire and cultivation could create

significant ecosystem effects. For example, the environmen-
tal historian J. R. McNeill notes that “Amazonian peoples
apparently painstakingly created their own patches of fertile
soil (for farming), the so-called ‘dark earths’ that make up as
much as ten percent of the rain forest region” (McNeill
2003:23). Similarly, the place names given to some locations
by native North Americans, such as Head-Smashed-In Buf-
falo Jump (Pyne 2001:31) in south-west Alberta, Canada,
attest to hunting methods, with appropriate knowledge of
landscape ecology, that contributed to effective population
manipulation of selected species, but not for the intention of
conserving them. Such population and ecosystem “manage-
ment” did have some of the same effects as conservation
management programs today – altering the distribution of Fig. 1.2 The sockeye salmon (Oncorhynchus nerka), a species that was
historically managed sustainably by the Tlingit and Haida tribes of
habitats, changing the composition of plant and animal Alaska through careful division of property rights, site-specific knowl-
communities, and providing benefits to human beings as a edge of habitat and resources, community-based rules of proper fishing
means of subsistence. But practices of this kind cannot be methods and site selection, and an intrinsic valuation and appreciation of
called “conservation” because they were not used to cause the the sockeye. (Photo courtesy of Ryan Hagerty, US Fish and Wildlife
Service)
persistence of, or “conserve” any of the species they affected.
This kind of human-centered “management” has instead been
correlated with extinctions of many species of large yitsati could use his power to enforce the regulations and to
mammals, especially in North America (Pyne 2001). punish violators, but sanctions were rare because fishing
Some hunter-gatherer societies did achieve remarkably rights could not be transferred. As a result, there were few
sustainable and sophisticated forms of resource management conflicts among users. These conservation practices, and the
that do meet even the most rigorous definitions of “conserva- sockeye salmon they conserved, collapsed in the late nine-
tion.” One such example is found in the allocation of fishery teenth and early twentieth centuries when other cultures, first
resources among Alaska Natives (USA). There are five spe- from Russia and then the United States, ignored the Tribes’
cies of salmon (Oncorhynchus spp.) (Fig. 1.2) that commonly established system and allowed anyone to place traps and
spawn in Alaskan rivers, but the sockeye salmon (O. nerka) weirs at the mouths of rivers.
has long been the most prized by Native Americans.
Sockeyes arrive earliest and remain longest in their spawning
streams, have the lowest historical variation in annual return 1.2.2 Ancient Traditions of Conservation
numbers, the highest nutritional value and, in the opinion of
some, the best taste (Leal 1998). The Tlingit and Haida As human societies shifted from hunting and gathering plants
Tribes, two groups of coastal indigenous Alaskan peoples, and animals to growing and domesticating them, they
placed a high value on the sockeye. Sockeye were scarce but continued to “manage” nature, but seldom conserve it. In
definable as property resources because sockeyes migrate ancient Greece, already an advanced pastoral and agricultural
only in stream systems that include a freshwater lake. The society, the Greek philosopher Plato compared present land
combination of scarcity and definable resource boundaries conditions to the past in one of his Dialogues through the
enabled the Tlingit and Haida to make rules limiting access to character Critias. Critias, in a conversation with the philoso-
particular streams to an individual clan or house group. pher Socrates, laments of the land that
Streams were assigned by the yisatii, the clan’s eldest male, of old its yield was most copious as well as excellent. . .By
as were limits on how many fish could be taken, and at what comparison with the original territory, what is left now is, so to
say, the skeleton of a body wasted by disease; the rich, soft soil
times. Management decisions were informed by the yisati’s has been carried off and only the bare framework of the district
intergenerational knowledge and life-long experience of both left. . .the soil got the benefit of the yearly ‘water from Zeus,’
the salmon and the stream, and therefore linked to time- and which was not lost, as it is today, by running off a barren ground
place-specific knowledge of resource constraint. The yisati’s to the sea; a plentiful supply of it was received into the soil and
stored up in the layers of nonporous potter’s clay. Thus, the
knowledge of family and clan structure also ensured that the moisture absorbed in the higher regions percolated to the
number of individuals consuming the resource was appropri- hollows, and so all quarters were lavishly provided with springs
ate to the sustainable level of production of the resource and and rivers. Even to this day the sanctuaries at their former
based on the size of the stream and the number of people sources survive to prove the truth of our present account of the
country (Hamilton and Cairns 1961:1216–1217).
required to set traps and weirs across the stream’s mouth. The
Before Plato, Judaism extended the principle of Sabbath,

or appointed rest, to include the land. Israel recorded God’s
command, through his prophet Moses, that the land must
receive a rest from cultivation every seventh year.
. . .in the seventh year the land is to have a Sabbath rest, a
Sabbath to the Lord. Do not sow your fields or prune your
vineyards. Do not reap what grows of itself or harvest the grapes
of your untended vines. The land is to have a year of rest (The
Bible, Leviticus 25:4–5, New International Version).
The Israelites, like the Greeks, abused their land and did
not keep this commandment, but it was not forgotten. Israel’s
prophets stated that failure to observe the land Sabbath was
one of the reasons for Israel’s eventual exile from Palestine.
Fig. 1.3 The “environmental impact triangle” displaying three key
Speaking of the defeat of Judah and the destruction of
elements of human environmental impact on nature: the interaction of
Jerusalem by Nebuchadnezzar in 586 B. C., the writer of the characteristics of the local natural environment (nature), the kind of
the Second Book of Chronicles records that Nebuchadnezzar local human economy employed by the human community, and the
carried into exile to Babylon the remnant, who escaped from perceptions of and attitudes toward nature by humans. Only when
human perception begins to view nature as something of value in itself,
the sword, and they became servants to him and his sons until
or something to be sustained beyond immediate need for future
the kingdom of Persia came to power. The land enjoyed its generations, does “conservation” emerge as a consistent practice in the
Sabbath rests; all the days of its desolation it rested, until the human community. (Developed from concepts described in A. McEvoy
seventy years were completed in fulfillment of the word of the 1988)
Lord spoken by Jeremiah (The Bible, II Chronicles
36:20–21, New International Version). Such case histories, although diverse in locality and
In China, the practice of setting aside “sacred” or fengshui worldview, reveal common patterns in human interaction
forests around every village, first developed over a with nature. Taken together, they reflect the principle that
1000 years ago during the Song Dynasty (960–1279 AD), human interaction with nature is determined primarily by the
was part of a religious practice that attempted to maintain characteristics of the surrounding environment, the character-
or create a spatial framework positively regulating spiritual istics of the local human economy that obtain needed and
power inherent in the physical landscape, promoting har- desired resources from nature, and human ideas and attitudes
mony between human and heavenly realms. Such a practice about nature (Fig. 1.3). In Plato’s example, hilly and moun-
had important practical implications. As one family geneal- tainous terrain (the state of nature) combined with herding
ogy from Sichuan noted, “When building manors and and farming (the human economy) with an assumption that
mansions, the gentry will not fell trees” (Coggins the land exists for growing crops and raising animals (the
2003:199). A village history from Anhui warns, “Every human attitude toward nature), led to rapid soil erosion and
family must take care of the mountains and waters around. loss of productivity. In the case of ancient Israel, a divine
Plant trees and bamboo as shelters. . .Keep an eye on the revelation offered a means to change this customary under-
environment and protect it from damage. This is a chore for standing of “the land” (nature) from a commodity to be used
people of one hundred generations to undertake” (Coggins to an entity enjoying a “right” to rest. Although the “new”
2003:199–200). The practices of fengshui declined in modern idea was not embraced and obeyed, it provides an objective
China under Communism, which disavowed the spiritual manifestation of the abstract concept that human attitudes
realities that fengshui addressed, and, more recently, free toward nature must regard the land with intrinsic value if
market capitalism, which has emphasized the preferences of human communities are to move from exploitive forms of
individuals above the good of the community. “management” to meaningful conservation.
These historical cases from Greece, Israel, and China are Changes in human attitudes toward nature affect human
among many examples worldwide which demonstrate that, treatment of it. Although humans have “managed” their
despite being given a teaching from a respected scholar, a physical environment for millennia, they began to conserve
command of God, or a charge from ancestors, people often nature only as they embraced certain ideas about nature. One
neglect and degrade the world around them. Neither Plato’s is that non-human creatures, and even the physical landscape
remembrances of a better landscape, nor religious commands itself, possess intrinsic value, that they are “good” in and of
that revealed land as an object of God’s care and concern, nor themselves. The sources of this perception are varied
sacred traditions of ancestors and kin have been sufficient to (Chap. 10), but, whenever humans begin to perceive nature
keep people from selfish behavior that ruined their and natural objects as good in their own right, they begin to
environment. treat them with increased respect. The perception of intrinsic
value leads to considering the interests and needs of the thing killing predators, introducing and translocating game animals
valued. When such things are considered as having “a good to increase their densities, and manipulating habitat.
of their own,” humans begin to consider what would be Such activities did not achieve a consistent approach to
“good for them.” That is, what human action would produce conservation because they operated within established social
conditions that would be “good” for nature? When humans boundaries of class, rank, and economic status characteristic
begin to ask this question, they begin to take actions that are of feudal societies and aristocracies. Sometimes their long-
the beginnings of “conservation.” term effects were exactly the opposite of “conservation,” as
the goals driving these practices made no consideration of the
species inherent value. In the United Kingdom, killing game,
1.2.3 Conservation as Expression of Privilege cutting trees, or even trespassing in royal reserves was a
serious offense, and violators often were deported to
Many early endeavors in conservation were not achieved by Australia, which the British originally used as a penal colony.
ethics, but enforced by punishment. In many cultures, con- Ironically, this practice helped to create a strong anti-
servation began with efforts to preserve nature by prohibiting conservation sentiment in that country. As historian John
some or all human use of resources in particular areas, except McCormack noted, “the idea of game laws was anathema to
for a privileged few. European and Asian royalty and other many Australians” (McCormick 1989:7). The abundance of
wealthy individuals set aside land as hunting and forest Australian fauna suggested that it should be everyone’s right
preserves, forbidding “common people” to kill game animals, to kill animals without hindrance. Thus, “because Australia
or even gather sticks within preserve boundaries. Violators was a free and expanding young society, it would reject the
were imprisoned or killed for their trespasses. Indeed, the notion of conservation of fauna as a hated relic of the feudal
word “forest,” a term of European origin, originally referred past from the Old Country” (McCormick 1989:7).
to areas where nobility had exclusive rights to game and Many countries within the Majority World, with long and
timber, controlling how much was taken and how much complex histories of colonialism, have also experienced
remained. William of Normandy established one such “conservation” as an expression of western privilege that
reserve, the New Forest, in England in 1085. Regarding his simultaneously disregards local human history and
affection for his New Forest and its creatures, one contempo- downplays the socio-political realities of a given space. As
rary wrote that “he loved the stags as dearly as though he had environmental anthropologists Stephen Nuget and Anja
been their father” (quoted in Holdgate 1999:2). Perhaps that Adams have noted, the tropical forests of Brazil have often
is why he did not allow anyone to hunt the stags without his been treated as if they were “practically uninhabited except
permission, and punished any common people who did, by fauna and by quasi-natural populations of humans”
whom he apparently loved less. making this so-called “primitive” and “natural” place of
With similar rationale and motivation, Chinese royalty particular interest to conquistadors, corporate industrialists
protected some game preserves with walls and guards, and, and conservationists alike (Anderson and Berglund 2003:5).
during the Qing Dynasty (1644–1911), even established This view ignored “already existing, complex social, ecologi-
a government agency called the “Bureau of Imperial Gardens cal, and political systems” and was used to justify the dispos-
and Hunting Parks” where royal reserves were divided session, displacement, and in some cases the slaughter of
into three categories: (1) Hunting Enclosures, (2) Enclosures indigenous populations from the fourteenth century to the
for Provision for the Royal Household, and (3) Enclosures present (Anderson and Berglund 2003:5). Even with the rise
for Military Training (Coggins 2003:11). Although such of community-based conservation in current conservation
prohibitions were primarily expressions of royal prerogative, efforts, which has sought to integrate rights and interests of
they also represented an early recognition of the limits of local populations into conservation objectives (Chap. 11), there
resource use, namely, that natural resources, such as forests often has been suspicion of international conservation efforts
or animal populations, could not produce sufficient resources because they were viewed, especially by local or indigenous
for exploitation by everyone. Unless exploitation was lim- residents, as purporting western privilege and prioritization of
ited, the resource would degrade. “nature” at the expense of local people and their needs and
Over time, cultural expressions of conservation in some desires. We shall see more of how these perceptions affected
cultures advanced from preservation by prohibition to active the development of conservation throughout the world as we
management of natural resources. In Europe, wealthy continue through this chapter, but first we must explore a more
individuals employed gamekeepers, whose function was to foundational question. How did people begin to re-envision
ensure an abundance of favored species for hunting. their relationship with nature in ways that replaced power and
Gamekeepers accomplished this, in part, by keeping out privilege with attitudes that would support more genuine and
vagrants and poachers, but also through such activities as lasting approaches to conservation?
1.2.5 Conservation as Knowledge – The

Point of Engagement Question Invitation to Study and Appreciate
Ongoing conservation work must be careful not to Nature
repeat past narratives of power and privilege in the
effort to protect non-human species and their habitats. In 1789, an obscure British clergyman, Gilbert White, pastor
Can you think of examples you know or experiences of the Church of Saint Mary’s in the village of Selborne,
you have had in which the conservation of “nature” England, published his records of over 20 years of natural
failed to perceive the needs of local people and their history study and observation. His book’s humble and
communities, or appreciate their own understanding of homely title was The Natural History and Antiquities of
appropriate nature conservation? Selborne (Fig. 1.4). No one then could have foreseen its
enormous success and popularity. More than 200 years
later, White’s work remains one of the classics of natural
1.2.4 Conservation as Right Relationship
history, continuously in print from the time of its publication,
with Nature – The Arcadian Vision
among the most published books in the English language and
recently ranked among the 100 greatest nonfiction books of
On the island of Peloponnesus, in what is today modern
all time (McCrum 2017).
Greece, was the province of Arcadia. According to Greek
The Natural History and Antiquities of Selborne is, in many
mythology, Arcadia was the domain of Pan, god of the forest,
ways, simply a well written and engaging naturalist’s journal
and his court of dryads, nymphs, and other spirits of nature.
of his local surroundings. White never ventures far from home,
Arcadia’s beauty, as well as the presence of spiritual beings,
writes about common and supposedly insignificant creatures,
gave it a reputation as an earthly paradise. Its human
and presumes no scientific credentials, authority, or mandate to
inhabitants were known for their simple, pastoral way of
justify his work. Yet his careful studies, exact and direct
life, living in harmony with the environments around them.
observations, simple but elegant experiments, and pleasant,
Over time, European art and literature transformed the
congenial warmth and humor made significant contributions
concept of Arcadia into an idealized place where both people
to science and drew thousands of people, formerly uninterested
and nature lived in harmonious simplicity, uncorrupted by
in nature, into more careful and intentional study of their
the vices of civilization, as portrayed in William
surrounding world. White, now considered England’s first
Shakespeare’s play, A Midsummer Night’s Dream, where
ecologist, grasped the importance of earthworms for soil pro-
an Arcadian-like realm is ruled by a fairy king and queen.
ductivity, was the first to describe the harvest mouse as a
Another example of the Arcadian ideal comes from the
species distinct from other kinds of mice, and noted the timing
English poet William Wordsworth, who wrote many poems
of hibernation in a tortoise. In the last example, one can see an
extolling the beauty and virtues of nature, and the value of
example of White’s inimitable style:
living in harmony with it. We can see this vision in his poem
about a shepherd, Michael, of the Lake District in England A land-tortoise, which has been kept for thirty years in a little
walled court belonging to the house where I now am visiting,
who lived and worked among
retires under ground about the middle of November, and comes
Fields, where with cheerful spirits he had breathed
The common air; hills, which with vigorous step
He had so often climbed; which had impressed
So many incidents upon his mind
Of hardship, skill, or courage; joy or fear;
Which like a book preserved the memory
Of the dumb animals, whom he had saved,
Had fed or sheltered, linking to such acts
The certainty of honorable gain; (Wordsworth 1975:63).
Although such an idealized view of nature did not stimu-

late actions or policies that we would today regard as conser-
vation, it created the perception that humans and nature were
meant to live in harmonious relationship marked by mutual
affection between humans and non-humans, and that nature
was to be valued for its own sake. The Arcadian view of
nature provided a foundation for the beginnings of a conser-
vation mentality in Western Europe and North America that Fig. 1.4 “The magpie,” an illustration in the first edition (1789)
would eventually grow to express itself in more specific and of Gilbert White’s The Natural History and Antiquities of Selborne, by
tangible ways. the noted Swiss artist Samuel Hieronymus Grimm (Public
domain image)
forth again about the middle of April. When it first appears in the cottage. . .” (McFarlane 1987:77). The nineteenth century
spring it discovers very little inclination towards food; but in the English poet William Morris captured the spirit of this long-
height of summer grows voracious: and then as the summer
declines its appetite declines; so that for the last six weeks in ing for the simple country life and the pleasures of rural
autumn it hardly eats at all. Milky plants, such as lettuces, nature in his poem, “The Earthly Paradise.”
dandelions, sow-thistles, are its favourite dish. In a neighbouring
village one was kept till by tradition it was supposed to be an Forget six counties overhung with smoke,
hundred years old. An instance of vast longevity in such a poor Forget the snorting steam and piston stroke,
reptile! Forget the spreading of the hideous town;
Think rather of the pack-horse on the down,
White’s writings offered no specific appeals to “conserve” And dream of London, small, and white, and clean,
nature. There was no incentive to do so because White simply The clear Thames bordered by its gardens green. . .(Morris 1905)
wrote about the world as he knew it, one in which there was
no scarcity of life around him and no apparent threat to its Gilbert White’s work was essential in providing a detailed
continuance. Grounded in the Arcadian view that nature was picture of what such a “back to the land” life might be like,
to be enjoyed for its own sake, and that an attitude of sim- and, in doing so, created a cultural climate in England recep-
plicity and humility were the prerequisites for such enjoy- tive to the first and earliest organized conservation efforts in
ment, White’s work encouraged ordinary people to “make the the western world, including the first organizations formed
acquaintance” of nature, to pay attention to and be careful specifically for conservation purposes. Among these were the
observers of the non-human world. In his own words, White Commons Preservation Society, initiated in 1865, formed to
frames this invitation to the reader in a poem he places at the prevent urban areas from encroaching on natural woodland
beginning of his book. and heath ecosystems (Guha 2000:16). Three years later in
1868 another conservation organization, the Association for
See, Selborne spreads her boldest beauties round the Protection of Sea Birds (APSB), was formed, and its
The varied valley, and the mountain ground, formation is a story worth telling (Information Box 1.1).
Wildly majestic! What is all the pride,
Of flats, with loads of ornaments supplied ?--
Unpleasing, tasteless, impotent expense, Information Box 1.1: Origins of the First Conserva-
Compared with Nature’s rude magnificence.
Arise, my stranger, to these wild scenes haste;
tion Organizations
Following the example of Gilbert White, many English
As years passed, readers of White, particularly in the citizens, including many members of the clergy,
United Kingdom, began to see the world that White became devoted students of natural history. Like
described, studied and loved, as increasingly imperiled by White, clergy often were particularly attracted to such
the unrestrained expansion of industrial development. As a study because, understanding the world as a “good”
result, one of the first expressions of conservation in England, creation of God (Genesis 1), they believed that such
and later much of western Europe, was the “Back to the study would lead them, and their congregations, to a
Land” movement (Guha 2000:6), a perspective that idealized fuller grasp of God’s providential character and good-
rural life reflected in the Arcadian perspective. White’s work, ness toward all that he had made. One individual who
appearing during the early stages of the British industrial exemplified this pattern was the Reverend Francis
revolution, struck an emotional chord with English and Orpen Morris (Information Box Fig. 1.1). Beginning
other European readers who were beginning to see the darker his work of overseeing the church at Nafferton in East
side of progress apparent in industrial development and Yorkshire, England, in 1844, Morris began to develop
attendant pollution. The nineteenth century English social a reputation as a naturalist through his books on natural
critic and ethicist John Ruskin described British industrial history, especially bird identification. Through such
progress as “the frenzy of avarice. . . daily drowning our work Morris became acquainted with many of
sailors, suffocating our miners, poisoning our children, and England’s best scientists, including John Cordeaux, a
blasting the cultivable surface of England into a treeless prominent naturalist of the time. Cordeaux was partic-
waste of ashes” (quoted in Cook and Wedderburn ularly interested in the study of bird migration, and
1908:137). Anthropologist Alan McFarlane noted of naturally focused his investigations on seabirds and
conditions during that time, “England was the most shorebirds along the Yorkshire and Dunham coasts.
industrialized country in the world, yet one where the yearn- Independently, both Morris and Cordeaux became
ing for the countryside and rural values was most developed. alarmed at the large numbers of sea birds being killed
Its strangely anti-urban bias was shown in the prevalence of by unregulated sport hunting, particularly in the spring.
parks, the ubiquity of flower gardens, the country holiday
industry, the dreams of retirement to a honeysuckle (continued)
Information Box 1.1 (continued) Information Box 1.1 (continued)

landowners to prevent hunter access to sensitive areas,
and then added political and social clout by gaining the
support of his own superior, the Archbishop of York, as
well as that of several members of Parliament. Barnes-
Lawrence’s contacts in Parliament then sponsored a
bill, the Sea Birds Preservation Act, which passed par-
liament in June 1869 and enforced a closed season from
1 April to 1 August. The first successful prosecution
under the Act took place in Bridlington on 10 July 1869
when Mr. Tasker, of Sheffield, was fined ₤3 19s for
shooting 28 birds (Dyson 1997).
The formation of the APSB marked the beginning of the

establishment of societies to protect specific plants and
animals. The formation of the British Royal Society for the
Protection of Birds (BRSPB) followed in 1889. In the United
States, the BRSPB was preceded by the formation of the
American Ornithologists Union in 1883 and the National
Audubon Society in 1886. Although the formation of the
Information Box Fig. 1.1 Francis Orpen Morris, a British
APSB marked the beginning of species-specific conservation
clergyman who was instrumental in founding the world’s first efforts through organized conservation advocacy, a second
organization for species’ conservation, the Association for the major form of conservation, the creation of parks and nature
Protection of Sea Birds. (Photo courtesy of Ash Midcalf and preserves from existing landscapes, would be radically
Birdcheck, public domain)
affected and transformed by developments in the United
States, creating a model for land- and ecosystem-based con-
servation that would come to be imitated throughout the
At that time, bird shooting and egg collecting were
world. This was the concept of the modern National Park.
widespread and unregulated. By the 1860s, in the
18 miles of coastline between Bridlington and
Scarsborough where Morris worked and lived, it was
estimated that 120,000 birds were annually trapped or 1.2.6 Conservation as Preservation
shot between April and August. of Landscape – The Washburn Expedition
Alarmed by these events, Morris presented a petition Goes to Yellowstone
to the House of Commons calling for a heavy tax on
guns, a move he hoped would curtail the loss of birds in In Europe, centuries of human impact on the natural land-
England, especially the loss of seabirds and shorebirds scape left few areas unchanged by human presence, and
in coastal areas. Morris engaged the help of his vicar unaffected areas that remained were often small and remote.
(bishop), the Reverend Henry Frederick Barnes- There was little opportunity to establish large areas for con-
Lawrenece, along with Cordeaux, to find ways to stop servation purposes, so reserves set aside for species or habitat
the slaughter of sea and shore birds. Barnes-Lawrence, protection were small. The first known nature reserve was the
who knew Morris as both a fellow Christian clergyman Karpf Stock, a mountain in Switzerland where the Canton
and bird lover, decided to attack the problem by forming (Council) of Glaus forbade hunting by anyone in 1576
an organization specifically dedicated to sea bird pro- (Holdgate 1999:3). At a similar scale, the Dutch Prince
tection. To form such an association, Barnes-Lawrence William of Orange set aside The Wood of the Hague in
used his influence to convene a meeting of local clergy 1576. By 1826, the English had established Walton Park as
and naturalists in 1868 to establish the Association for the worlds’ first bird sanctuary. But in these and other cases,
the Protection of Sea Birds (APSB). At the same time, the areas preserved were small and not unaltered by human
Barnes-Lawrence worked to secure the support of local habitation or development. In a remote region of the western
United States, an 1870 expedition would initiate an entirely
(continued)
new paradigm in conservation at a regional landscape level: of the area and what ought to be done about them. Hedges
the US National Park. suggested that they should not abandon the area to commer-
In 1807, John Colter, who left the famous Lewis and Clark cial development. Rather, Yellowstone should be set aside as
expedition in 1806 to become an independent trapper and a “national park” for the enjoyment of US citizens, as well as
explorer, found a region of incredible scenic beauty and visitors from around the world (Frome 1987). It was an idea
unbelievable thermal wonders, including geysers, “pots” of never before proposed, that a government should preserve the
boiling mud, petrified forests, and hot springs in what is today best part of its natural heritage so as to make that heritage
northwestern Wyoming. His reports of the area were not accessible to everyone, a uniquely democratic vision of con-
initially believed but led others to name the area “Colter’s servation. What US historian and environmental novelist
Hell.” A former military scout, Jim Bridger, explored the area Wallace Stegner called “the best idea we ever had” (Stegner
23 years later in 1830. He returned with even more incredible 1998:137) rose from the sparks of an evening campfire to
stories, including tales of a river fed by geysers and thermal become an ideal emulated round the world – the ideal of the
springs that “got hot on the bottom” (what is called today the national park.
Firehole River), cliffs of black glass (obsidian), and springs Convinced of the “rightness” of their vision, leading
belching sulfurous steam. But his reports, often embellished members of the expedition returned to the eastern United
(Bridger spoke facetiously of “petrified birds that sing States to initiate a campaign for Yellowstone’s protection.
petrified songs in petrified trees”), were treated with doubt A scientific expedition the following year confirmed their
and referred to as “Jim Bridger’s lies” (Frome 1987). reports, complemented by the stunning photographs of
Despite public skepticism, persistent rumors of the William Henry Jackson and dazzling watercolor paintings
region’s incredible natural wonders eventually led to a num- of Thomas Moran (Fig. 1.5), both of whom had accompanied
ber of visits to the area by other explorers and early settlers, the party. On March 1, 1872, Congress approved a bill
and their reports continued to build its reputation. Finally, in creating Yellowstone National Park “as a public or pleasuring
August of 1870, an official expedition led by the Surveyor ground for the enjoyment of the people” (Petulla 1977). By
General of Montana, Henry Washburn, with US Army Lieu- 1916, Congress would create the National Park Service,
tenant Gustavus Doane commanding a military escort, left directing it to “conserve the scenery and the natural and
Fort Ellis, Montana and proceeded south toward “Colter’s historic objects and the wild life therein, and to provide for
Hell.” The Yellowstone River and its surrounding area, the the enjoyment of the same in such manner and by such means
“Yellowstone Country,” were so called because of the yel- as will leave them unimpaired for the enjoyment of future
low- and copper-colored formations of rhyolite, a volcanic generations” (National Park Service Organic Act 1916).
rock that characterized the river’s spectacular canyons Today, much of the worldwide efforts in conservation biol-
(Clepper 1966; Frome 1987). The expedition members ogy rest on the legitimacy of designing and establishing
enjoyed themselves immensely. Speaking of the trout they conservation reserves, and on the assumption that such
caught, one wrote, “Few of them weighed less than two reserves are an important part of national and cultural heri-
pounds, and many of them over three. They had not been tage. Such a foundational premise of conservation is now so
educated up to the fly, but when their attention was respect- familiar it is taken for granted. In truth, the idea of landscape-
fully solicited to a transfixed grasshopper, they seldom failed scale conservation reserves, first expressed in the US concept
to respond” (Anonymous 1871). of the National Park, is a radical and recent concept. It created
Members of the Washburn expedition found the area even an enduring paradigm for conservation throughout the world
more amazing than earlier tales they had heard and meticu- and was widely exported as a model for other countries.
lously chronicled and sketched what they encountered. One The Yellowstone Model, however, was strongly
member, Cornelius Hedges, described a thermal spring he influenced by North American landscape characteristics, as
observed in a meadow. “This spring, with two others, was well as by perspectives and prejudices of US culture. The
situated in about an east and west line, and at the upper side of expedition members were primarily concerned with the
the basin, which opened south, toward the creek. The central enjoyment of their fellow citizens, not with the preservation
one of these three was the largest of all, and was in constant, of the Yellowstone Ecosystem. They envisioned a form of
violent agitation, like a seething caldron over a fiery furnace” “development”, facilitated by the railroad that would make
(Anonymous 1871). Speaking of these and similar features, Yellowstone accessible to millions with relatively little cost
Hedges remarked, “If a person should be cast into one of or effort. One member expressed these aspirations this way.
these springs, he would be literally immersed in a lake of As an agricultural country, I was not favorably impressed with
burning brimstone” (Anonymous 1871). the great Yellowstone Basin, but its brimstone resources are
Seated around the flames of a campfire on a September ample for all the matchmakers in the world. A snow-storm in
evening at the junction of the Firehole and Gibbon Rivers, the September, two feet deep, is hardly conducive to any kind of
agriculture or stock-raising;. . . When, however, by means of the
members of the expedition discussed the remarkable wonders
1.3 Foundations and History of Conservation in the United States 11
Fig. 1.5 Castle Geyser, Upper

Geyser Basin, Yellowstone
National Park, painted by Thomas
Moran, official artist of the
1871 US Geological Survey
Expedition to Yellowstone.
(Courtesy of the US National Park
Service, Yellowstone National
Park)
Northern Pacific Railroad, the falls of the Yellowstone and the

geyser basin are rendered easy of access, probably no portion of 1.3 Foundations and History
America will be more popular as a watering place or summer
resort than that which we had the pleasure of viewing. . . (Anon- of Conservation in the United States
ymous 1871).
1.3.1 Conservation as Moral Mission – John
The members of the Washburn Expedition perceived the Muir and Theodore Roosevelt
beauty and uniqueness of the Yellowstone landscape, but
not of the people living in it. There was no consideration as The American conservationist, John Muir (Fig. 1.6), although
to how the Park’s establishment would affect or include the generally credited as the person most responsible for the
Native Americans who were living there and had claims to development of the US National Park Service and System,
land and resource ownership in Yellowstone. spent most of his years as a conservation activist concerned
Modern conservationists may be forgiven for smiling at with the practical problem of how to save the Sierra Nevada
the thought of seeing Yellowstone National Park used as a Mountains of the US state of California from logging,
giant supply depot for match making, but it is wise to remem- mining, hydrological development, and other forms of com-
ber that even well-educated people seldom challenge cultural mercial exploitation in the late nineteenth and early twentieth
patterns of thinking in which they have been raised. The US century. Prior to his work in conservation, Muir invented and
would not expressly establish a national park to protect an manufactured mechanical equipment, but nearly lost his eye-
“ecosystem” until the formation of the Florida Everglades sight in an Indianapolis, Indiana carriage factory when a file
National Park in 1934. With all its cultural blind spots, the he was using slipped and went through one eye. Eventually
US vision of the national park was a watershed event in the recovering his sight but shaken by this near tragic experience,
development of conservation biology, for it formalized the he retired to the wilderness in an attitude of repentance and
concept of setting aside land areas of landscape scale specifi- reexamination of his life. After traveling on foot for over a
cally to preserve natural settings, and the animals and plants thousand miles through parts of Florida, Georgia, Kentucky,
within them. The creation of national parks, and later national and Tennessee, he eventually turned west and reached the
wildlife refuges and national forests, in the United States Sierras. There, Muir had an intense religious experience,
began to change the way in which people, and, perhaps just gaining a sense of profound fulfillment and exaltation, as
as importantly, their government, interacted with nature. The well as an intense oneness with the land around him (Petulla
US conservation effort, which began as a moral crusade, 1977). But, seeing destruction and degradation of the Sierras
grew and changed to become a policy and priority of the and other natural areas everywhere, Muir at once began a
federal government itself. public campaign to save the American wilderness.
extraction of its resources as commodities for human use and

material goals, but the appreciation of it intrinsic values and
aesthetic qualities through which the human spirit was
transformed, a place where one could draw near to and
commune with God (Callicott 1990). Muir shared this view,
and made it clear in his writings that people who used nature
as a place for religious worship, aesthetic contemplation,
inner healing, rest, and relaxation were making a “better”
(that is, morally superior) use of nature than those who cut
the trees, dammed the rivers, mined the minerals, or plowed
the soil. Nature, Muir believed, was to be preserved in an
undisturbed state so that the higher uses and values could be
appreciated and enjoyed. As he worked out the practical
implications of the transcendentalist view, Muir became a
Fig. 1.6 US conservation activist John Muir, founder of the Sierra Club fierce advocate of protecting nature from development and
and champion of the Preservationist Approach to conservation. Muir, exploitation precisely so that it could be enjoyed in these
who framed conservation as a question of moral choice, was instrumen- higher and better ways. Muir was instrumental in framing the
tal in establishing the US National Park Service. (Courtesy of the US debate in conservation around the essential question: What is
Library of Congress, ca. 1902)
the best and “highest” use of nature and natural resources?
This question still resides at the core of conservation debate
Muir was an eloquent and persuasive writer, and his
today and remains a key to understanding the development of
articles in major newspapers and national magazines urged
conservation biology as both a scientific discipline and cul-
Americans to (temporarily) leave the cities and enjoy the
tural force.
wilderness. As his experience had been religious, his writing
used religious language to express his concerns, as
exemplified by the title of one of his early and most influen-
1.3.2 Conservation as Utilitarian Purpose –
tial articles in the Sacramento (California) Record-Union,
Gifford Pinchot and Sustainable Yield
“God’s First Temples: How Shall We Preserve Our Forests?”
(Petulla 1977). His worldview, combative nature, moral
1.3.2.1 The Federal Government Empowers
absolutism and relentless opposition to “development” in
Conservation as Science and Democratic
his beloved Sierras are best expressed in his own words.
Ideal
The battle we have fought, and are still fighting, for the forests is Even as Muir was framing the key question of the great
part of the eternal conflict between right and wrong, and we
conservation debate as a moral appeal, the end of the western
cannot expect to see the end of it. . .The smallest forest reserve,
and the first I ever heard of, was in the Garden of Eden; and frontier in the United States had begun to affect a profound
though its boundaries were drawn by the Lord, and embraced change in US environmental attitudes. Settlers had been
only one tree, yet even so moderate a reserve as this was faced with a hostile, often life-threatening environment on
attacked. And I doubt not, if only one of our grand trees on the
the frontier that could be made livable only by strenuous
Sierra were reserved as an example and type of all that is most
noble and glorious in mountain trees, it would not be long before effort and significant environmental alteration. Yet, for all
you would find a lumberman and a lawyer at the foot of it, its hardship, westward expansion gave an impression that
eagerly proving by every law terrestrial and celestial that the there would always be new lands to settle and more natural
tree must come down. So we must count on watching and striving
resources to use. The passage of the US Homestead Act of
for these trees, and should always be glad to find anything so
surely good and noble to strive for (Muir 1896:276). 1862, which offered a free 160 acres (65 ha) of western land
to any person who would build a house on the acreage and
Muir was the father of a new school of thought and live in it for five years, provided motivation for thousands of
activism in US conservation, the philosophy of US citizens to begin dreaming of a new life in the West. The
preservationism. Muir and other preservationists condemned end of the US Civil War in 1865 brought peace to the nation,
the destruction of nature to satisfy what they considered the released most men from military service, reunited thousands
greedy appetite of materialism. In doing so, Muir and his of families throughout the country, and provided an opportu-
allies were drawing on the ideas of earlier American writers nity for people to begin making plans for moving west. And
like Ralph Waldo Emerson and Henry David Thoreau the completion of the transcontinental railroad in 1869 gave
(Fig. 1.7) who had refined the Arcadian Ideal into the more ordinary citizens a means of transport to western areas for-
formal philosophy of Romantic Transcendentalism, a view merly inaccessible to them. Armed with such means, motive,
that argued that the highest and best use of nature was not the and opportunity, US citizens began to occupy formerly
Fig. 1.7 Walden Pond, a 26 ha

glacial kettle lake in the US state
of Massachusetts. The writer,
transcendentalist, and philosopher
Henry David Thoreau resided on
the northern shore of Walden
Pond for two years (1845–1846)
during which time he attempted to
live quietly on and from the land
itself. His experience inspired
Thoreau’s classic work, Walden,
or Life in the Woods, a book in
which Thoreau, in relating his
experiences, encourages respect
for and personal communion with
nature, manifested in simplicity of
life and direct connection to the
land. (Courtesy of the US Library
of Congress, ca 1908)
remote western areas. As the frontier came to an end and the pursuit of the second began at almost the same time. In 1873,
density of human populations began to grow even in formerly just one year after establishing Yellowstone, the US Con-
desolate areas, US citizens now found themselves living in a gress, in an effort to aid western homesteaders, passed the
nation of increasingly well-defined physical and environmen- Timber Culture Act, which permitted the clearing of up to
tal limits. At the same time, use and exploitation of natural 160 acres of timber if the owner replanted trees on 40 of those
resources had begun to shift from individual to corporate acres. Although intended to help individual families, the law
effort and from simple exertions of human and animal labor was used most effectively by timber companies to clear large
to increasingly sophisticated applications of advanced tracts of forests with minimal reforestation. In 1891, at the
technologies. Freed from many of the former limitations, urging of many scientific and professional societies, Presi-
corporate interests in mining, lumber, fishing, and grazing dent Benjamin Harrison, aided by his Secretary of the Inte-
began to rapidly exploit and extract resources over large rior, John W. Noble, persuaded Congress to pass a bill
areas. For example, the US state of Michigan, the nation’s repealing the Timber Culture Act and granting (in a relatively
largest producer of timber as recently as 1900, had removed unnoticed rider on the bill) presidential authority to set aside
most of its commercially valuable timber by 1920, effectively forest reservations (Petulla 1977). The Forest Preservation
“cutting” itself out of the lumber business (Dickman and Act, as it came to be called, was designed to prevent western
Leefers 2003). In states further west, similar trends were in US forests from suffering the same fate as their counterparts
motion. in the states of Michigan, Minnesota, and Wisconsin.
The establishment of Yellowstone as a US national park As many private citizens were becoming increasingly
had set a legal and political precedent for the federal govern- vocal in their opposition to the destructive use of natural
ment to take a more active role in conservation, but the exact resources, particularly on public lands, the power of the
nature of that role remained undefined. As government influ- federal government and its involvement in conservation was
ence in conservation began to increase, it began to take two growing. The assassination of President William McKinley
separate, sometimes conflicting, paths. One led to increasing in 1901 brought his young Vice-President, Theodore
influence in establishing parks and nature preserves that were Roosevelt, to the Oval Office (Fig. 1.8). Roosevelt, an active
to be protected from exploitation and disturbance because outdoorsman, began to take an interest in western lands,
of their own intrinsic value, the other to increasing authority particularly in the use of forest and wildlife resources. By
in management and administration of lands where resources this time, John Muir had formed the Sierra Club, whose
would be used and harvested sustainably for the public good. purpose expressed Muir’s own philosophy of
The establishment of Yellowstone Park was a monumen- Preservationism, “to enlist the support of the people and the
tal step for the federal government down the first path, but government in preserving the forests and other features of the
Fig. 1.9 Gifford Pinchot, American forest scientist and administrator

who developed the US Forest Service and its philosophy of Sustained
Yield and Multiple Use during the administrations of US President
Fig. 1.8 Theodore Roosevelt, the US President who made conservation Theodore Roosevelt. (Photo by Francis Benjamin Johnson between
a national priority during his presidency from 1900 to 1908. (Courtesy 1890 and 1910. Courtesy of the US Library of Congress)
of the US Library of Congress, ca 1903, Rockwood Photo Company)
Sierra Nevada Mountains” (Petulla 1977). In his campaign to of logging and mining in the western US with similar hostil-
save the Sierras, especially Yosemite Valley, Muir had con- ity. They were not only wasteful, but undemocratic. To
vinced the federal government to make Yosemite, then a state Roosevelt, it was clear that a handful of individuals and
preserve, into a national park in 1890. Muir’s influence, their companies were reaping most of the profits from natural
subsequently empowered by Roosevelt’s position and actions resources that rightfully belonged to all citizens. Thus, he saw
as President, gained acceptance for the concept of national federal regulation as appropriate and necessary because he
parks throughout the West and laid the foundation for the viewed the federal government as the only institution power-
eventual establishment of a government agency, the National ful enough to oppose corporate timber and mining interests
Park Service, to oversee their management. With the estab- that threatened to destroy the landscape.
lishment of Pelican Island, Florida, as a wildlife sanctuary in On this basis, Roosevelt used the provisions of the Forest
1902, Roosevelt also began an ambitious program to create Preservation Act aggressively and often by setting aside large
federally protected wildlife refuges throughout the US, forested areas in the western states. Although the administra-
establishing 52 such preserves during his administration, tion of such lands was logically the domain of the Depart-
along with 16 national monuments and five national parks. ment of the Interior, Roosevelt distrusted that agency, fearing
The refuges Roosevelt preserved were the beginning of what that corporate interests in timber and mining had corrupted
would grow to become the National Wildlife Refuge System, it. Determined to protect the new forest reserves, Roosevelt
today administered by the US Fish and Wildlife Service created a new federal agency, the US Forest Service. He
(USFWS), although the USFWS itself would not exist as a placed it in the Department of Agriculture (which he consid-
federal agency until many years later. ered less corrupt) and put his own man, Gifford Pinchot, in
Ironically, the pursuit of the second pathway of govern- charge (Fig. 1.9). This appointment brought conservation
ment involvement in conservation, one that led to increasing efforts in the US under the influence of conservation practices
government oversight and jurisdiction of management of the developed earlier in European settings.
sustainable use and harvest of natural resources, also was
furthered under Roosevelt, aided by his association with 1.3.2.2 German Influences in Conservation –
John Muir. Alerted and persuaded by Muir and others of Forest Monocultures and Maximum Yields
increasing environmental degradation in the American In Europe, with its longer history of human settlement and
West, Roosevelt increasingly viewed the large corporations landscape cultivation, natural resources like forests were
that were unfairly profiting from western lands with hostility managed by human manipulations in ways that were not
and distrust. Known as “The Trustbuster” for his zeal in fundamentally different from commercial agriculture. The
breaking up industrial monopolies concentrated in the eastern most advanced techniques were practiced by Germans, who
United States, Roosevelt came to view the corporate practices founded the disciplines of silviculture (tree growing) and
forest management in the 18th and 19th centuries. Under a worldwide turning to science and technique-oriented man-
strong central government during the reign of Fredrick the agement, an influence that was keenly felt and exemplified in
Great (1740 to 1786) that practiced close supervision of state the development of conservation in the United States. In
forests, German foresters began to move from traditional 1886, the US Government made its first appointment of a
“area-based” concepts of management that focused on divid- scientifically trained forester to serve in government-directed
ing forest units by landscape features to a “yield-based” forest management. The position was that of Chief Forester
management approach. The yield-based management system, of the US Department of Agriculture, and the appointee was
which worked best with stands containing only one tree not a US citizen, but, again, a German forester, Bernard
species, was based on direct estimates of volume and weights Fernow. Fernow believed that commercial production of
of trees of different ages. By studying growth patterns that timber was the primary purpose of a forest and referred to
could be determined on small, experimental plots, German forests as part of the “great economy of nature.” His views
foresters were able to develop highly accurate and predictive would set the course of forest management and conservation
standard yield tables for every species of commercially in the United States for years to come and contributed to a
important tree. Using such tables, foresters could accurately serious split in the ranks of the US conservation movement.
estimate not only the wood mass of individual trees but of
entire stands, even over fairly long time periods. Armed with 1.3.2.3 The Rise of the Resource Conservation
such predictive power, German foresters began to develop Ethic
the concept of sustained yield forestry which managed the Within the US Department of Agriculture, the energetic and
forest in such a way that the same harvest could be reliably talented young forest scientist, Gifford Pinchot, rose rapidly
removed every year without loss or interruption of forest to prominence in developing forest management and policy.
productivity. Pinchot was a US born scientist but had been trained in
Encouraged by their successes, German scientists enthusi- German traditions of scientific forest management under the
astically promoted their approaches to forest management mentorship of none other than Dietrich Brandis. Pinchot also
abroad, not only in Europe but also in its colonies and in was a younger contemporary, and, for a time, close friend of
the US and Canada. So esteemed were the German forest John Muir. Pinchot was not a preservationist like Muir and
scientists of that time that even countries with respected did not subscribe to the Romantic-Transcendentalist ethic of
scientific communities of their own often “imported” German nature preservation for the sake of moral values. He saw
foresters for forest management, consultation, and practice. timber in the new reserves as an exploitable resource to be
In British-dominated India, the first Conservator of Forests used with careful application of scientific management.
who was appointed in Bombay (modern Mumbai) in 1847, Pinchot, in the United States, as well as Brandis in India,
and another in Madras in 1856, were both British citizens. saw that high rates of deforestation led to soil erosion, loss of
However, by 1864, as the British sought to consolidate their soil productivity, water pollution, and even changes in local
forest management in India under one administrative unit, climate such as desiccation and drought. Muir believed that
they appointed a German, Dietrich Brandis, as the first Forest such environmental degradation was best stopped by preserv-
Superintendent of India, empowering his work a year later ing the land from exploitive use, but Pinchot and Brandis saw
with the passage of the Indian Forest Act. Under the authority the solution in scientific forest management. As a result,
of the Act, Brandis was able to organize all Indian forest Pinchot can justifiably be called the father of a new ethic,
management under a single administrative unit, the Forest the Resource Conservation Ethic, sometimes simply
Department of India, and applied German silviculture referred to as “resourcism” – a view distinct from the
philosophies and methods to forest management throughout Romantic-Transcendentalist Ethic. Pinchot crystallized the
the subcontinent. He would lead the agency for almost philosophy of the movement in a simple, memorable slogan
20 years, during which time it would come to control nearly (which he credited to a contemporary, W. J. McGee): “the
one-fifth of India’s land area (Guha 2000:26). Thus, even as greatest good for the greatest number for the longest time”
the British were imposing “colonial conservation” on India, (Callicott 1990).
they were themselves subservient to German concepts of The Resource Conservation Ethic rested on two intellec-
forest management. German approaches eventually would tual pillars. The first was equity – resources should be justly
be challenged in the US and throughout the developing and fairly distributed among present and future generations.
nations of Africa and Asia, but they effectively ruled global The second was efficiency – resources should not be used
conservation practice for much of the nineteenth century and wastefully. Pinchot and others who advocated for a resource
first half of the 20th. conservation ethic were not, like Muir, concerned with the
The sustained yield forestry developed by the Germans “best” use of nature (which they called “natural resources”) in
was a classic example, but only one example, of an overall the sense of moral superiority. They were concerned with the
practice and philosophy of conservation marked by a “fair” (i.e., democratic) and “sustainable” (i.e., longer-term)
use of nature. They believed that all interests in resource use,

both consumptive and non-consumptive, should be consid-
ered and, when possible, satisfied. Time and again Pinchot
made clear that, in his view, conservation did not mean
protecting or preserving nature. It meant wise and efficient
use of natural resources, informed by scientific study and
practice, with the goal of controlling nature to meet human
needs over the long term (Nash 1989). This view ultimately
led to the end of the friendship between Muir and Pinchot, but
neither man wavered in his convictions. Pinchot’s view of
nature as an array of resources to be used productively for the
common good led him and others to advance the concepts of
multiple use (using the same systems to supply or satisfy
many different needs) and maximum sustained yield
(maintaining a constant level of extractive use approximately
equal to the rate of renewal of a renewable resource, such as
timber). After his career in the Forest Service had ended, Fig. 1.10 Aldo Leopold, US conservationist and founder of The Wil-
Pinchot helped to establish the Yale School of Forestry at derness Society, a chief intellectual architect of the “Wilderness Ideal” in
conservation at his “shack” near Baraboo, Wisconsin (USA). His
Yale University, the first of its kind in North America, and
experiences there inspired much of his classic work on land
one deeply embedded in German philosophies and techniques ethics, posthumously published in A Sand County Almanac (Chap. 2).
of forest management for maximum sustainable yield. (Photo courtesy of the Aldo Leopold Foundation Archives)
Although sustained yield and multiple use concepts were
being developed and practiced in other parts of the world, probably the greatest laboratory that nature furnishes on the
they were, in the early twentieth century, nowhere more clearly surface of the globe” (Holdgate 1999:6). Had Doane been a
manifested in land management policies than in the United man of science, he might have had opportunity to act on this
States. Both were to become important planks in the foundation remarkable insight and begin unique and original
of modern forestry and models for other nations in their devel- investigations in Yellowstone or other wilderness areas.
opment of national forest management programs. By the early Instead, it remained for a future scientist of the twentieth
twentieth century, the rise of independent nation states through- century to articulate the value of wilderness for both scientific
out the world, combined with the growing centralized power of and cultural purposes.
the national governments and the development of ecology as a Aldo Leopold (Fig. 1.10), born and raised in Iowa
scientific discipline, permitted the management of lands at (USA) in the 1880s when that state still had many
national scales. It began to make sense to speak of “national characteristics of a wilderness, was the son of highly
forests” and “national grasslands” as resources managed by a educated German immigrants, and trained as a forester in
national government for the good of its citizens (Guha the best German traditions of that science at Yale University.
2000:27), and to manage such lands according to strategies of After graduation, Leopold joined the US Forest Service
“multiple use” and “sustained yield.” where he rose with stunning rapidity through the ranks,
Even as the US federal government was intensifying the gaining the position of Supervisor of the Carson National
extraction and use of resources on public lands for utilitarian Forest in New Mexico at age 24. Along with his obvious
purposes, and at the same time establishing an increasing intellect and energy, Leopold displayed even keener interest
number of national parks and wildlife refuges, another impor- and insight toward wildlife than he did toward trees, an
tant paradigm in conservation was beginning to emerge in the orientation not unnoticed by his superiors. Increasingly
United States that would influence conservation worldwide. stimulated by opportunity and encouragement to conduct
This was the concept of the Wilderness Ideal. investigations of game populations in national forests,
Leopold, as result of his ongoing studies, became convinced
that, like forest management, the practice of game manage-
1.4 Aldo Leopold and the Formation ment could be performed scientifically.
of the “Wilderness Ideal” Leopold’s experiences and reflections culminated in his
classic textbook, Game Management, published in 1932,
With an incredible depth of vision, Lieutenant Gustavus which in turn led the University of Wisconsin at Madison
Doane, leader of the military escort of the Washburn Expedi- to offer him a faculty position in the field he had created.
tion, remarked after his return that Yellowstone should not be Leopold accepted the offer and helped to organize one of the
viewed simply as a public pleasuring ground, but that “as a first academic departments in wildlife management in the
field for scientific research it promises great results. . .It is United States.
1.5 The Emergence of Global Conservation 17
The synthesis of Leopold’s experience in the Forest Ser- on public lands, and the “Wilderness Ideal” were not all
vice and his studies in academia left him disillusioned with entirely US inventions, but they were concepts that flourished
the German models and concepts of intensively managed in US culture and government, and subsequently exported
monocultures as the ideal form of forest conservation. In and adapted globally. However, as a wider world conserva-
later years, following a trip to Germany and an inspection tion movement began to take shape in the twentieth century,
of its forests and forestry management practices, Leopold, the flaws in such models appeared when they were uncriti-
dismayed by the German obsession with spruce, lamented, cally accepted and applied in other contexts. There was often
“never before or since have the forests of a whole nation been a mixed set of goals undergirding these transnational efforts,
converted into a new species within a single generation.” The and in some cases, western socio-cultural assumptions
Germans, Leopold wrote to a colleague in disgust, had replaced context-specific scientific assessment and cultural
“taught the world to plant trees like cabbages” (quoted in sensitivity, to the detriment of both human and non-human
Guha 2000:55). Leopold was not only disillusioned with populations. As conservation became a global concern and
German-inspired principles of forest monocultures, but also conservationists a global community, new models emerged
with the philosophical underpinnings of the Resource Con- to make conservation viable in contexts vastly different from
servation Ethic that supported it. Leopold became convinced the US and European experience.
that such an ethic was inadequate, principally because it was
untrue. The land was not, as management science taught, a
Point of Engagement Question
collection of separate, compartmentalized products that could
Which conservation framework do you believe is most
be managed for commodity output. Rather, the land was a
prominent in our society today? What contemporary
system of interdependent processes. The outcome of those
laws and policies are in place that manifest values
processes, when they functioned properly, was sustained
espoused by such a framework?
production of the commodities associated with the processes,
such as soil, water, timber, wildlife, and forage for wild and
domestic animals. This vision of managing land as a system
rather than as a storehouse of commodities had profound
ethical implications for land management and treatment,
implications that Leopold would articulate years later in his 1.5 The Emergence of Global Conservation
most well-known book, A Sand County Almanac, a work that
would ultimately inspire a new ethical philosophy in conser- 1.5.1 Multilateral Treaties – The Beginnings
vation called ecocentrism (Chap. 10). of International Conservation Efforts
Shortly after his appointment at the University of
Wisconsin in 1935, Leopold helped to form The Wilderness 1.5.1.1 Conservation Driven by Shared
Society, a group of scientists, scholars, and conservation Commercial Interests
activists who shared the conviction that remaining roadless Even as new forms of conservation in philosophy and prac-
areas, still relatively untouched by mining, grazing, logging, tice were emerging in the United States in the late 19th and
and roads, had value if preserved in their current state. early 20th centuries, international paradigms of conservation
Leopold’s claim that “Wilderness is the raw material out of were developing throughout the world. As international com-
which man has hammered the artifact called civilization” merce and trade increased in the nineteenth century, particu-
(Leopold 1966: 264) reflected a long-held conviction. In larly in the West, nations found increased cooperation a
1924, he had been instrumental in getting the US Forest necessity in evolving international dealings, international
Service to establish its first wilderness area, the Gila Wilder- agreements and conventions (treaties). Such conventions
ness, in New Mexico. In this way, Leopold led the US in addressed resources that could be disputed, “natural” or not,
making yet another substantive contribution to global conser- and attempted to govern the use of natural resources, espe-
vation practice, the manifestation of the Wilderness Ideal as cially migratory animal species, that crossed international
an important means of conserving landscapes, habitats, and boundaries. One of the earliest of these agreements was the
biodiversity, ultimately culminating, after his death, in the Convention Respecting Fisheries, Boundary, and the Resto-
passage of the US Wilderness Act of 1964. The Wilderness ration of Slaves established between the United States and
Act created the means to develop and administer an entire Great Britain in 1818. Although this treaty had some effect in
system of roadless, uninhabited areas, and in the Act’s own protecting some commercial fish stocks, its primary purpose
words, “untrammeled by man” and “for the permanent good was to provide clear allocation of boundary and property
of the whole people” (Public Law 88–577). rights between the two nations. Subsequent agreements
The “Yellowstone Model” of the national park, the “max- between Britain and France on fisheries (1867), between
imum sustainable yield” approach to resource management European nations on salmon fishing in the Rhine River
(1886), and between the US and Britain on the taking of fur legislative consequence of the treaty was the Migratory Bird
seals in the Bering Sea were all conventions driven by the Treaty Act (MBTA), ratified in the United States in 1918.
same purpose. Multilateral and bilateral treaties addressing The Act prohibited taking or killing of migratory birds with-
migratory species date to the late nineteenth century out a permit and imposed strict penalties and liabilities for
(Holdgate 1999). violations. Indirectly, the MBTA was the forerunner of many
Initially international cooperation was motivated by future US laws that empowered state and federal agencies to
concerns over trade, and the dawning awareness that com- regulate hunting and punish violators of game laws, and
merce that depended on migratory species needed those many of these eventually came to serve as models for other
species to have sustainable populations and sustainable levels nations.
of harvest. The Treaty Concerning the Regulation of Salmon As the US federal government began to take an increasing
Fishery in the Rhine River Basin, signed in 1885 by the role in conservation during the late nineteenth century, it also
Federal Republic of Germany, Switzerland, and The began to act to secure international provisions for conserva-
Netherlands, prohibited the use of fishing methods that tion, particularly where such provisions affected US
blocked more than half of a watercourse, prescribed concerns. Given its worldwide interests, influence, and
specifications for fishing nets, provided for closed seasons ambitions, the US also took initiative in creating international
and regulation of fishing hours, and promoted captive- organizations designed to influence, if not govern, many
breeding. These were all measures designed to enhance the aspects of conservation policy, such as the International
sustainability of the salmon population by reducing its har- Association of Game, Fish, and Conservation
vest, removing impediments to migration and spawning, and Commissioners in 1902 and the American Committee for
enhancing wild populations by supplementing their numbers International Wildlife Protection in 1930.
with captive ones. Many of these US efforts took place during the adminis-
tration of Theodore Roosevelt, who viewed conservation as a
1.5.1.2 International Protection of Migratory moral ideal. “There can be no greater issue,” he stated, “than
Species that of conservation in this country. Conservation is a great
Given that one of the oldest problems in conservation has moral issue, for it involves the patriotic duty of insuring the
been that of managing species that cross international safety and continuance of the nation. . .[I] do not intend that
borders, it is no surprise that some of the earliest treaties our natural resources shall be exploited by the few against the
related to conservation addressed the conservation of migra- interests of the many, nor do [I] intend to turn them over to
tory species, including: (1) species that bred on the shore or in any man who will wastefully use them by destruction, . . .the
rivers leading to the ocean, but spent their adult life in the sea; rights of the public to the natural resources outweigh private
(2) migratory marine species that traveled over ocean areas rights and must be given its first consideration” (quoted in
across national boundaries; (3) terrestrial species that Holdgate 1999:8).
migrated from breeding to non-breeding areas across interna- Had Roosevelt remained in office, it is likely he would
tional boundaries; and (4) non-migratory terrestrial species have moved his national concerns for conservation to the
that lived near international boundaries and routinely crossed international level and made the United States a leader in
them in the course of normal movements. international conservation. However, having taken a public
From treaties that were concerned with commerce and pledge not to seek a third term as President, he resigned from
trade, agreements began to develop that recognized more office at the end of his second administration. That resigna-
intrinsic values of the species themselves. In 1902, the Inter- tion, the departure of Gifford Pinchot from the Forest Service
national Convention for the Preservation of Useful Birds was in 1910, the death of John Muir in 1914, and the onset of
signed by 12 European nations. Although it did not protect all World War I sent conservation to the shadows of interna-
species of birds (for example, birds that ate crop plants were tional politics, although it continued to develop nationally as
not considered “useful”), it was an important first step in an issue in many countries.
international conservation of migratory species. One of the National leaders in the twentieth century increasingly
most effective and enduring treaties protecting migratory perceived that international cooperation would be the only
birds followed soon after: The Convention between the sure way to achieve international results, in conservation or
United States and Great Britain for the Protection of Migra- anything else. The need for increasing international coopera-
tory Birds (1916), which protected birds flying between the tion to protect migratory species from overexploitation began
US and Canada. Originally motivated by a desire to conserve to be replaced in importance by the emerging awareness of
waterfowl, the treaty established hunting regulations and the needs of future generations through sustainable resource
closed seasons for ducks and geese. It also prohibited hunting use. Motivated by a growing level of accountability to their
of migratory “insectivorous” birds (most songbirds), and citizens, government officials in democratic countries began
established refuges for selected species (Holdgate 1999). A to realize that they must act in the best interests of not only
the present generation, but of the generations to come. Such

realization was an early, emergent perception of the modern
concept of sustainability, but, for the first half of the twentieth
century, there was no place to even discuss such a concept,
let alone promote it through international agreement. A per-
manent forum for international cooperation had to be
established if conservation was to become an internationally
coordinated effort, rooted in recognized international law.
1.5.2 Forums for International Conservation –

The UN and the IUCN
Although nations increasingly began entering into bilateral

and multilateral agreements throughout the first half of the
twentieth century, there was no recognized international body
that could propose or create international laws or treaties
aimed at conservation, nor was there any established interna-
tional organization that served to connect the varied and
diverse efforts of conservation beginning to arise throughout
the world. Some European nations, such as Great Britain, Fig. 1.11 Paul Sarasin, Swiss scientist and international statesman of
had, in the past, been able to create organizations, laws, and conservation, who founded the Swiss League for Nature Protection and
policies of international effect because their influence and whose vision of an international conservation network eventually led to
the formation of the International Union for the Conservation of Nature
jurisdiction applied to all British colonies globally. (IUCN). (Photo courtesy of Image Archive ETH-Bibliothek Zurich,
Organizations such as The Society for the Preservation of under CC BY-SA 4.0)
Wild Fauna and Flora of the Empire (1903), while thoroughly
“British,” affected species protection in Africa, Asia, and on the state of world conservation. Unfortunately, the Com-
South America. However, as colonial domination by mission gradually slipped into lethargy and inaction, and
European powers declined, there was increasing need for a finally fell apart. Sarasin did not give up, but he was unable
more representative international body to address global con- to revive the Commission despite constant and strenuous
servation law and policy. It was a scientist and political efforts over the next 15 years. Sarasin died in 1929, with no
activist of Switzerland, Paul Sarasin, who first conceived of sign that his vision of an international conservation organiza-
an international organization whose task would be to attempt tion would ever become a reality. Inspired by Sarasin’s work,
to coordinate and unite the efforts of conservation similar initiatives were launched in the 1930s, but these also
organizations worldwide. withered and died without effect. The onset of WWII ended
In 1909, Paul Sarasin (Fig. 1.11), with his cousin Fritz, further discussion or development of Sarasin’s vision.
founded the Swiss League for the Protection of Nature (or, in In October of 1944, with the end of WWII in sight,
its original language, Ligue Suisse pour la Protection de la US President Franklin Delano Roosevelt proposed a
Nature). An active and respected zoologist, Sarasin urged the meeting of “the united and associated nations [for] the first
Congress of Zoology in the following year to establish “a step towards conservation and use of natural resources.” In a
Committee charged to establish an international or world memo to his Secretary of State, Cordell Hull, Roosevelt
Commission for the protection of nature. . .throughout the wrote “I am more and more convinced that conservation is
world, from the North Pole to the South Pole, and covering a basis of permanent peace” (quoted in Holdgate 1999:15).
both continents and seas” (Holdgate 1999:11). Roosevelt did not live to see his ideas bear fruit, but his vision
The Congress did establish an ad hoc committee, but such and initiatives contributed to the formation of the United
a grand scheme as Sarasin’s needed more than committee Nations (UN) in 1948. The formation of the UN brought
meetings to succeed. Persistent and determined, Sarasin with it the formation of two UN programs, the United Nations
eventually was able to bring together representatives from Educational, Scientific, and Cultural Organization
16 European nations and the US at a meeting in Berne, (UNESCO) and the United Nations Environmental
Switzerland in 1913. There the representatives agreed to Programme (UNEP), both with strong interests in conserva-
establish a Consultative Commission for the International tion. UNESCO and UNEP began to provide the international
Protection of Nature that would assemble and publish issues forums and multi-national networks that brought
conservation to the international agenda (more on these nature” (Holdgate 1999:33). This was tall order for an orga-
developments in Chap. 12). nization with no endowment, budget, or full time employees.
With the establishment of an international forum for Nevertheless, the IUPN put forward an entirely new organi-
global conservation issues, UNESCO’s first director, British zational model, the Governmental and Non-Governmental
biologist Julian Huxley, revived Sarasin’s vision of an inter- Organization (GONGO), whose members and contributors
national, non-governmental conservation organization dedi- would not be individuals but nations and organizations.
cated to networking the global conservation effort. Although Supported, at least in spirit, by UNESCO, the IUPN pursued
Sarasin’s International Commission for the Protection of two simple but critical objectives. First, create a working
Nature had not survived, the Swiss League had, and its worldwide network of conservationists who would use the
representatives, along with leading conservationists from Union as their primary mechanism of information exchange.
other countries, pressed Huxley to revive the Commission Second, convene conferences that would regularly reinforce
under the auspices of the UN. Working with an existing such exchanges. Eventually articulating its mission more
organization, the International Council of Scientific Unions clearly through carefully framed goal statements, the Union
(ICSU), leaders of the Swiss League and others labored with began its work. Its first major strategic success was the
Huxley over a period of 2 years to convene an international creation of the “Survival Service” in 1950. Working with a
conference at Fontainebleau, France in 1948. The conference grant of US$2500, the Survival Service began to recruit
was attended by representatives of 23 governments, volunteer scientific experts to write reports on the conditions
126 national institutions and eight international of endangered species around the world. Although it had no
organizations. On October 5, 1948, most of the delegates money to fund field investigations, its reports drew increasing
present signed a formal act creating the International Union respect, and alarm, in the international community as the
for the Protection of Nature (IUPN). The Union was to Union patiently and persistently documented the plight of
“encourage and facilitate cooperation between Governments global endangerment. Working directly with government
and national and international organizations concerned representatives from many nations, the Union began to have
with, and persons interested in, the ‘Protection of success in its appeals to national leaders to work to conserve
Nature’. . .promote and recommend national and interna- endangered species within their borders. The growth and
tional action. . .” to preserve wildlife and “collect, analyze, development of the Union deserves special attention (Infor-
interpret and disseminate information about the ‘protection of mation Box 1.2).
Information Box 1.2: The Birth of the IUCN Information Box 1.2 (continued)
Now, more than 70 years later, International Union for the known as the World Conservation Union, has effectively
Protection of Nature (IUPN) has changed almost every- achieved the vision that Paul Sarasin pursued but never
thing about itself, including its name (now the Interna- lived to see: a worldwide network of conservation efforts
tional Union for the Conservation of Nature, or IUCN), to preserve biodiversity. In addition to its direct efforts,
its logo (Information Box Fig. 1.2), and its programs (the IUCN’s networking and information services have made
Survival Service became the Species Survival Commis- thousands of other conservation organizations more effec-
sion). Its once disparate reports now are published com- tive and stimulated the creation and development of many
prehensively as the world famous Red Data Books, the new organizational initiatives. In multiple ways and by a
authoritative standard for the global “Endangered variety of measures, IUCN has been and remains today
Species List” of plants and animals. What the Union has one of the world’s most important actors in the develop-
not changed is its mission. Today the IUCN, now better ment of a truly worldwide conservation effort.
Information Box Fig. 1.2 The evolution of the official logo of the early IUCN staff, adopted in 1954, the “Letter Block” design
International Union for the Conservation of Nature (IUCN), from the adopted in 1977, and more recent emblems adopted in 1992 and
original “Flaming Artichoke” or “Brussels Sprout,” as nicknamed by 2008. (Figures courtesy of the IUCN)
(continued)
For all its achievements, current members of the Union on it (Guha 2000:41). As one Indian noted, remarking on the
(now the International Union for the Conservation of effect of scientific forestry on the actual conservation of
Nature or IUCN) would be the first to admit that organiza- native biodiversity and cultural practices,
tional development is not the same as conservation success. Small landowners who could not subsist on cultivation alone
The threats to biodiversity that it was established to combat used to eat wild fruits and [berries] and sell the leaves and
have grown more serious. The future of conservation will not flowers of the flame of the forest and the mahua tree. They
be determined only by scientific expertise, but by engage- could also depend on the village ground to maintain one or
two cows, and two to four goats, thereby living happily in their
ment of ordinary citizens with models of conservation appro- own ancestral villages. However, the cunning European
priate to and workable in their own nations and cultures. To employees of our motherly government have used their foreign
achieve this, the world conservation effort must adapt its brains to erect a great superstructure called the forest depart-
traditional and, in some circles, revered, historical models ment. With all the hills and undulating areas and also the fallow
lands and grazing grounds brought under the control of the
of nature preservation to changing conditions and forest department, the livestock of the poor farmers do not even
circumstances. We turn back to those models of scientific have place to breathe anywhere on the surface of the earth
conservation and sustained yield, the “Yellowstone Model” (quoted in Guha 2000:39).
of the national park, and the “Wilderness Ideal” to see how
It was not merely scientific conservation that stood
their adaptation, and, in some cases, replacement, in other
accused. As noted earlier, the Yellowstone Model of the
cultures has driven and quickened the development of con-
national park was easily exportable to nations that possessed
servation in recent decades.
large land areas relatively unaffected by human development,
especially where such land was either uninhabited or where
the inhabitants had no standing as citizens and therefore
1.5.3 New Expressions of Resource could be forcefully relocated. Not surprisingly, the first
Management, National Parks and Nature countries to imitate the US model were Australia (1879),
Preserves Canada (1885), South Africa (1890), and the Scandinavian
countries of Europe (beginning with Sweden, which
In India, sacred forests had been protected from hunting,
established its first national park in 1903). The concepts of
logging, and other forms of destructive use for over
national parks and wilderness areas, which these countries
2000 years based on the Hindu belief that each forest was
adopted, viewed the human presence as destructive, some-
the dwelling place of a deity (Apffel-Marglin and Parajuli
thing to be excluded if the “real” values of nature were to be
2000). Such protection was usually effective even though the
preserved. This destructive view of human impact, however,
forests were often surrounded by areas of dense human
was not one that could be exported to the densely peopled and
habitation (Holdgate 1999:2). British colonization of the
much altered landscapes characteristic of many countries
Indian subcontinent in the 18th and 19th centuries brought
worldwide. The US model was inapplicable in settings like
with it very different rationales for forest conservation. Under
western Europe, where humans had been resident on land and
the successive administrations of German forest scientists,
affected it for centuries, such that there were no large unde-
Dietrich Brandis, Wilhelm Schlich, and Bertold von
veloped tracts left, or in Central America, where there was
Ribbentrop, government foresters in India confidently little undegraded land remaining. In countries where indige-
adopted intensive, sustained yield management, converting
nous peoples had lived on the land without “developing” it
thousands of acres of diverse, species-rich native Indian
for millennia, the “Yellowstone Model” was not only inap-
forests to single-species plantations of economically desir- propriate but unjust, forcibly removing thousands of
able trees. Although such an approach maximized timber
individuals from their homes, native landscapes, and cultural
production, the associated revenues and other benefits went
practices in the name of conservation. As environmental
primarily to industry and government, not to local citizens. scholar Ramachandra Guha noted regarding the establish-
The German model, outstanding as a timber production para-
ment of game reserves and national parks in the Republic of
digm, destroyed the resource base for local extractive
South Africa, “Where did the African fit into all this? To be
economies and subsistence agriculture which had developed precise, nowhere. . .In game reserves Africans were barred
over many centuries. Unfortunately for local citizens, Indian
from hunting, while in national parks they were excluded
independence from Great Britain did not immediately change
altogether, forcibly disposed of their land if it fell within the
forestry practices. By the 1990s, one study determined that boundaries of the designated sanctuary” (Guha 2000:46–47).
130 years of state (whether Indian or British) forest manage-
Another pillar of North American conservation, the “Wil-
ment had left the forests in worse shape than when “scientific
derness Ideal,” also proved problematic in other cultural
forestry” first appeared. The investigators noted ruefully that, settings. In the United States, designated wilderness areas
by this time, 22% of the nation’s land was still controlled by
were established on much the same grounds as national
the Forest Department, but less than half of that had any trees
Table 1.1 Contrasts between key conservation issues in wilderness preservation and biodiversity conservation
Issue Wilderness preservation Biodiversity conservation
Objective Landscapes without humans Biological diversity at all levels of organization
Justifications Aesthetic Intellectual interest; present and future utility
Targets National parks; wilderness preserves High-biodiversity regions; representative sample of biodiversity
Obstacles Economic interests; overconsumption; human Economic interests; overconsumption; human encroachment; invasive
encroachment; invasive technologies technologies; habitat fragmentation; human exclusion, in some cases; diversion
of scarce resources from conservation to wilderness
Strategies Legislation; habitat purchase Diverse methods
Reprinted by permission of Oxford University Press, from Sarkar, S., 1999. Wilderness preservation and biodiversity conservation—keeping
divergent goals distinct. BioScience 49, 405–412. # 1990 American Institute of Biological Sciences
parks, except without developments for the comforts of Valley in California, which would eventually become, in
visitors. US conservationists tended to conflate wilderness large part through Muir’s efforts, Yosemite National Park
and park preservation, and the absence of permanent human in 1890, Muir said that “Indians walked softly and hurt the
presence, with biodiversity preservation. As conservation landscape hardly more than the birds and squirrels” (Nabban
biologist Sahotra Sarkar has noted, two problems are created 1995). But Muir and his allies were dedicated to preserving
in developing countries when the Yellowstone Model and the the park, not the Indians, and did not perceive that the
Wilderness Ideal are employed uncritically. First, local indigenous people had any inalienable right to residence or
populations are displaced and their economies disrupted or resource use in Yosemite, no matter how long they might
destroyed. Second, conservationists fail to see that high have lived there. In fact, most Native Americans had left the
levels of biodiversity may be in areas with high densities area by the time Yosemite was established, primarily from
and long land tenures of indigenous peoples, whose historic earlier conflicts with white settlers and gold prospectors. This
practices of farming and resource extraction are beneficial to pattern of displacement and deportation of native people from
biodiversity enhancement (Sarkar 1999). As such, conserva- national parks was repeated consistently in the United States,
tion strategies must be rethought in cultural context, and their and subsequently throughout the world, especially in Africa,
objectives clarified, because wilderness preservation and bio- Asia and Australia. Displacement and deportation were also
diversity conservation approach similar issues with very dif- justified by the rationale that wildlife needed to be
ferent goals, strategies, justifications and targets (Table 1.1) “protected” from the hunting and trapping practices of indig-
(Sarkar 1999). enous people. Aristocratic hunters in Great Britain, for exam-
ple, who had formed the previously mentioned Society for
the Preservation of Wild Fauna and Flora of the Empire in
1.5.4 Conservation as Preservation of Culture 1903, were concerned with the decline of wildlife in British
and Livelihood – The Extractive Reserve colonies, particularly in Africa. Many of the original
members believed that such declines were the result of deci-
The Yellowstone Model for conservation reserves was an mation by “savage” hunter-gatherers who captured animals
inspiring ideal actualized with practical success in species through inhumane practices of fire, poison, traps, snares and
and landscape preservation, but it was not without intellectual pits. Society members considered that conservation would be
flaws or social injustice. From its initial implementation in best achieved by segregating wildlife into protected areas,
the founding of Yellowstone National Park, US conserva- abolishing hunting practices of indigenous people, and
tionists and preservationists saw reserve lands as “empty relocating and concentrating their communities in govern-
space,” unoccupied by people. This perception was false. ment settlement areas (Neumann 2002), a pattern of thinking
Native Americans were resident in Yellowstone and other with many parallels to the original US philosophy of national
US national parks formed in the nineteenth century and were parks and the preservation of the wildlife within them which
deported when the parks were created. Although the United justified removal of native human residents.
States recognized many Native American tribal groups as This pattern of forced removal was also followed in South
sovereign nations, the US government, through treaties with America, where displacement of indigenous peoples was
such groups, acquired possession of Native American lands sometimes explicitly incorporated into government policy
through “sales” associated with the treaties, or in some cases to advance national interests, especially the interests of
simply dispossessed Native Americans in favor of other non-indigenous citizens. Ruled by a military government
competing national interests. Some conservation leaders, until the 1970s, Brazil’s leaders in this regime described the
such as John Muir, did not see Native Americans as a threat Brazilian Amazon region as “a land without men, for landless
to preservation or conservation. Speaking of the Yosemite men,” a slogan intended to inspire and encourage
non-indigenous Brazilians to move to and occupy the Ama-

zon Basin as part of a program of national development. In
this view, the “men” (as well as women and children) native
to the Brazilian Amazon did not exist. As early as 1973 Brazil
began to demarcate indigenous territories under the 1973
Statute of the Indian, but initially such demarcation did not
necessarily prevent Brazilians from other parts of the country
from working or living in the indigenous territories. Ironi-
cally, the perception of indigenous territories began to
change, not so much from actions taken by indigenous peo-
ple, but from non-indigenous Brazilians working in and near
indigenous reserves. The most influential of these were the
rubber-tappers.
Traditionally, rubber was gathered from trees on individ-
ual estates of wealthy landowners. The landowner assigned a Fig. 1.12 Chico Mendes, Brazilian rubber tapper and union leader who
pioneered the development of the Extractive Reserve as a new way to
particular group of workers, the rubber-tappers, to a living
create protected areas designed to safeguard livelihoods of indigenous
space called a colocacão (literally “a putting, placement”) people and workers dependent on products of natural ecosystems.
within the estate and close to the rubber trees. This arrange- (Photo taken by M. Smith, reprinted under CC BY-SA 4.0)
ment, like sharecropping in US agriculture, created a “debt
bondage” system of employment that tended to perpetuate advocacy for establishment of an “extractive reserve” dedi-
the poverty of the rubber tappers and deepen their indebted- cated to protecting the livelihood of rubber tappers. As
ness to and dependence on the landowner. But as rubber Mendes explained, “. . .the Indians have a right to an area, a
prices collapsed worldwide in the 1960s, so did this system forest reserve for Indians, and the rubber tapper ought to have
(Schwartzman 2018). Rubber tappers became independent, a forest reserve for the rubber tappers. . .” (quoted in
working within Amazonian forests collecting rubber and Allegretti 2002:398). Thus, Mendes and other union leaders
other forest products and selling them to independent buyers. changed their strategy from simply protecting their own right
Not surprisingly, as independent traders, the rubber tappers to work to forming coalitions with indigenous people to
eventually formed a trade union, the Xapuri Rural Workers’ establish reserves in which both native people and rubber
Union, in 1977. tappers could practice a traditional way of life.
Concurrent with these events, the Brazilian government Such an approach was increasingly successful, not only
was continuing to pursue policies to “develop” the Amazon politically in Brazil but in drawing global attention to the
region through increasing accessibility to the area through problem of deforestation, framing the conflict not simply as
more road building, which in turn facilitated higher levels of an effort to preserve biodiversity but as a way to preserve the
timber harvest, deforestation, and conversion of land use to dignity of human culture and traditional livelihoods. But as
cattle ranching. Union leaders began to realize that continued the success of the movement grew, so did the strength of
deforestation would deprive rubber tappers of their opposition. From the late 1970s through the early 1990s,
livelihoods, and so began to oppose such policies. One many union leaders were assassinated by or at the behest of
union leader, Chico Mendes (Fig. 1.12), began to see that ranchers, including Mendes in 1988.
such a strategy would have to embrace forest conservation. Although Mendes’s life ended with assassination, his
Initially rubber tappers would confront or surround cause and vision continued to flourish. By 1990, the first
employees of ranchers cutting trees in the forest and persuade Extractive Reserve, Alto Jurá, was established in the
or prevent them from continuing their work. This produced Brazilian state of Acre in the southwestern portion of the
some success in negotiating with the ranchers, such that Brazilian Amazon, where the rubber tappers movement had
ranchers began to give small parcels of land to individual first originated (Gomes et al. 2018). Both rubber tappers and
rubber tappers. But, as Mendes began to realize, “We discov- indigenous people continued their protests but became more
ered as time went on. . .that what was important wasn’t tactically sophisticated and politically engaged in their
making agreements with the ranchers and getting a piece of efforts. Marina Silva (Fig. 1.13), a former rubber tapper,
land. What was important was to struggle for the conserva- was elected to Brazil’s federal senate in 1994 and reelected
tion of our forest, the defense of the rubber trees, and finally in 2002, and in 2003 appointed Brazil’s Minister of the
the defense of our landholdings. . .” (quoted in Schwartzman Environment. Silva became a major influence in the develop-
2018:60). ment and expansion of Extractive Reserves. By 2018, Brazil
At this point, reserves were beginning to be established in had established 51 federal extractive reserves in Amazonia
Brazil for indigenous people (“indigenous reserves”), and covering 12 million hectares (nearly 30 million acres) and
Mendes and other rubber tappers transferred the concept to 25 state-established reserves covering two million hectares
representing indigenous people are now increasingly present

at and contributing to global conservation meetings (Paulson
et al. 2012). Thus, today, especially in countries in the
Majority World, both governments and international bodies
are beginning to incorporate concern for indigenous
landholdings and culture into their design of conservation
reserves. Nevertheless, the Forest Peoples Programme, one of
the best known and globally effective organizations
representing indigenous people, has observed that, so far,
the inclusion of principles in conservation planning to con-
sider the needs of indigenous peoples has still not been
widely applied (Paulson et al. 2012).
As conservation has become an international effort and
has grown more cognizant of human interaction with nature,
Fig. 1.13 Marina Silva, a former rubber tapper who became Brazil’s new models of nature preservation like the Extractive
Minister of the Environment and played a leading role in the develop-
ment of Extractive Reserves in the Brazilian Amazon. (Photo courtesy Reserve facilitated by international programs like UNESCO,
of L. Cabral, reprinted under CC BY 2.0) UNEP, IUCN, and others have replaced traditional
approaches. The Man and Biosphere Program, begun in
1970 under the auspices of UNESCO, is one approach
(nearly five million acres) (Gomes et al. 2018) (Fig. 1.14) intended to provide a means for establishing the equivalent
Silva also played a leading role in establishing Brazil’s Chico of “world national parks” in a manner very different from the
Mendes Institute for Biodiversity Conservation in 2007 Yellowstone Model. The Program was designed “to establish
(Gomes et al. 2018). Over the same period, Brazil also a scientific basis for the improvement of relationships
became an international leader in the establishment of between people and their environments” (UNESCO 2019).
demarcated indigenous lands, which, by 2005, covered an Out of such effort has come the concept and implementation
area of 820,000 km2 (about twice the size of California, of the Man and Biosphere Reserve. Biosphere Reserves are
the third largest state in the US). One of the best examples land areas designated for conservation with the intent of
is the Kayapó Indigenous Territory, an area of over 11 million fostering “the harmonious integration of people and nature
ha inhabited by only 4500 people (Mittermeier et al. 2005). for sustainable development through participatory dialogue,
In light of these and other events, we can now justifiably knowledge sharing, poverty reduction, human well-being
view Mendes’s own words about his understanding of improvements, respect for cultural values and by improving
his work in the rubber tappers’ movement as prophetic. society’s ability to cope with climate change” (UNESCO
“At first I thought I was fighting to save the rubber trees, 2019). Such language was intentionally crafted to not focus
then that I was fighting to save the Amazon forest. Now I exclusively on indigenous cultures or traditional land uses,
realize that I’m fighting for humanity” (quoted in Haberman, but to also include opportunity for more current forms of
2016:1). natural resource use and its incorporation into contemporary
local economies. For this reason, Man and Biosphere
Reserves (686 in 2019) typically include areas dedicated
1.5.5 Indigenous People, Integrated exclusively to nature conservation through exclusion of
Development, and Conservation Concern development or extraction of natural resources, as well as
areas of permanent human residence and sustainable resource
With the development of the rubber tappers’ movement and use. For example, in the W Region Boundary Reserve that
the increased establishment of both Indigenous and Extrac- includes land in the African nations of Benin, Burkina Faso,
tive Reserves, indigenous people in Brazil, and subsequently and Niger, there are resident human communities in rural
around the world, have intensified their demands for the right villages (Benin and Niger) and cities (Burkina Faso), with
to live on their own traditional lands and use them in tradi- ongoing economic activities including agriculture, animal
tional ways, and public, social, cultural and governmental husbandry, craft production, and tourism (UNESCO 2019).
awareness of indigenous people and their relationship to the In areas designated for human residence and development,
land has increased. Such intensifying demands and increas- Biosphere Reserves often permit and engage a great deal of
ing awareness have led to a discernible and growing connec- local participation in the management of the preserve, not
tion between the work of conservation and the issues of social simply relying on centralized government control or scien-
justice, to the point that individuals and organizations tific expertise.
1.6 Return to Start: What Is the Place of Conservation Biology in the World. . . 25
Fig. 1.14 Spatial distribution and temporal phases of establishment of C.V. Gomes, from Gomes, C.V., Alencar, A., Vadjunec, J.M., Pacheco,
federal and state-level Extractive Reserves in Brazilian Amazonia. Through L.M., 2018. Extractive Reserves in the Brazilian Amazon thirty years after
2018, Brazilian Extractive Reserves protected over 35 million acres (14 Chico Mendes: social movement achievements, territorial expansion and
million hectares) of Amazonian forest. (Reprinted by permission from continuing struggles. Desenvolvimento e Meio Ambiente 48)
As Yellowstone Model reserves have been faulted for ecosystem the reserve was supposed to protect (Ma et al.
their failure to protect indigenous people, Man and Biosphere 2009).
Reserves have been criticized for sometimes uncritical and
only weakly restrained incorporation of modern practices of
agriculture and resource use that have compromised their 1.6 Return to Start: What Is the Place
ability to protect native plants and animals from exploitation. of Conservation Biology in the World
In areas within Biosphere Reserves “zoned” principally for Conservation Effort?
conservation, efforts in species protection have often been at
least somewhat effective. In the zones open to more intensive 1.6.1 The Emergence of Conservation Biology
development, conservation efforts have sometimes suffered. from the Applied Sciences
For example, in China’s Yancheng Biosphere Reserve,
populations of the endangered red-crowned crane (Grus If conservation is only a moral or political cause, then it
japonensis) (Fig. 1.15) have declined in the face of develop- hardly needs a separate science to bolster it. The classical
ment activities in the “transition zone” that have degraded disciplines of biology, chemistry and physics, supported by
wetland habitats, leading to an increased concentration the more recent but now well established applied sciences
of cranes in the interior “core zone” of the refuge, which like forestry, wildlife management, and fisheries manage-
has consequently required replacement of natural wetlands ment, would be sufficient to inform policy makers and
with artificial wetlands designed to produce more food for activists to make the right decisions and support the right
cranes (Fig. 1.16). Although such created habitats have effec- causes. Indeed, some of the world’s most influential
tively supported cranes and other wetland dependent bird conservationists have strengthened, and in some cases even
species, they have compromised the integrity of the wetland founded, such classic or applied disciplines. What, then, is
Fig. 1.15 The endangered red-crowned crane (Grus japonensis),

whose numbers have declined with increasing development in the
Yancheng Biosphere Reserve in eastern China, particularly in the
“buffer” and “transition” zones of the reserve which were designed to
integrate human development activity with biodiversity conservation.
(Photo taken by A. Rae, reprinted under CC BY-SA 2.0)
the function of conservation biology as a professional scien-

tific discipline, and does it possess sufficient distinctions to
endure in the years to come?
Through the work of Aldo Leopold and others, applied
sciences in resource management gained academic
respectability in state universities after the 1930s. The most
pervasive and influential of these disciplines included for-
estry and silviculture, fisheries management, (outdoor) recre-
ation management, range management, and wildlife ecology
(the modern version of game management). In addition, the
traditionally “pure” discipline of ecology increasingly fea-
tured studies of species or systems with clear implications for
conservation. The inaugural issue of the Journal of Wildlife
Fig. 1.16 (1) Relationship between percentage of red-crowned cranes
Management (JWM), the official journal of The Wildlife (Grus japonensis) in the core (a, b), buffer (c, d) and transition (devel-
Society (TWS), defined wildlife management as “the practi- oped) zones relative to the total number of cranes (a, c, e) and percentage
cal ecology of all vertebrates and their plant and animal of developed land (b, d, f) in the Yancheng Biosphere Reserve, China.
associates” (Bennett et al. 1937). In fact, the lead article of (2) Percentage of cranes in natural (solid bars) and artificial (hollow
bars) habitats in the Yancheng Biosphere Refuge, China. Artificial
the inaugural issue actually conflated wildlife management habitats include aquaculture ponds, farmland, saltworks, and managed
and conservation biology as one and the same, saying “In the reedbeds. (Reprinted by permission from Wiley, from Ma, Z., Li, B., Li,
new and growing field of conservation biology. . .” W., Han, N., Chen, J., Watkinson, A.R., 2009. Conflicts between
(Errington and Hamerstrom 1937:3). Early issues of JWM biodiversity conservation and development in a biosphere reserve. Jour-
nal of Applied Ecology 46, 527–535. # 2008 British Ecological
in the 1940s showed promise of embracing this definition, Society)
featuring a number of multiple-species studies and non-game
studies (Bunnell and Dupuis 1995). However, in the decades
that followed, JWM and TWS became increasingly
dominated by studies of game mammals and birds. was an emphasis on studies of individual species of interest to
Applied sciences like wildlife management, forestry, conservation. The other was to stress the study of individual
fisheries management, range management and others were types of habitats. Using this kind of thinking, conservation
hindered from embracing studies of biodiversity and multiple was essentially a case-by-case effort. Effective conservation
species in other ways besides their emphasis on economically was based on knowing everything possible about the natural
valuable species and commodity uses of resources. Their history of the species of interest and then preserving as much
paradigm of conservation science, developing initially in of its habitat as possible. Refuge design was therefore based
the United States, rested primarily on two approaches. One on preservation of habitat for a particular species, whether it
Fig. 1.17 Three species, snow

goose (Chen caerulescens, a),
Kirtland’s warbler (Setophaga
kirtlandii, b) and Joshua tree
(Yucca brevifolia, c), which have
been the focus of special
management areas, refuges, or
national monuments in the US
designed primarily for their
benefit. (Snow goose photo
courtesy of US Fish and Wildlife
Service. Kirtland’s warbler photo
courtesy of US Forest Service,
Huron-Manistee National Forest.
Joshua tree park photo courtesy of
H. Schwalbe/US National Park
Service)
was, in the US, the Kirtland’s warbler (Setophaga kirtlandii) managers also increased their focus on conservation issues.
on specially purchased land in Michigan, the snow goose Some wrote textbooks on “biological conservation” and
(Chen caerulescens) at DeSoto National Wildlife Refuge in started a journal with the same name.
Iowa, or the Joshua tree (Yucca brevifolia) at Joshua Tree Although the applied sciences were becoming more inclu-
National Monument in California (Fig. 1.17). Such effort was sive in their definition and conception of “wildlife,” tensions
intelligent and sincere, but did little to establish general between the applied sciences and the conservation movement
principles that could be applied to all species or to produce were growing. With prophetic insight, Leopold had foreseen
unifying theories of refuge design. the beginnings of these conflicts decades earlier and predicted
Despite these problems, the applied sciences in general, the outcome. In an essay titled, “Land Health and the A-B
and wildlife management in particular, prospered in the cli- Cleavage” (Leopold 1966), he wrote about a bifurcation
mate of a growing environmental and conservation move- common to academic specialties in resource management.
ment of the 1960s and 70s. Legally supported by laws like the “Group A,” wrote Leopold, “regards the land as soil, and its
US Endangered Species Act and aided by associated funding, function as commodity production; another group (B) regards
studies in wildlife management became increasingly impor- the land as biota, and its function as something broader”
tant as sources of scientific information for recovery plans for (Leopold 1966:258–259). Resource management fields such
threatened animal populations. Wildlife ecology, although as wildlife management, forestry, and range management,
continuing to emphasize studies of animals that were hunted (the A group) did not fully embrace either the values
or trapped for profit or recreation, expanded to include espoused by Leopold and others or the growing emphasis
specialties such as “nongame wildlife management” and on nongame species (the B group). Although many biologists
“urban wildlife management.” Wildlife ecologists and in wildlife management and other sciences respected
Leopold’s ethical position that valued all species in the con- worldwide extinctions and loss of species, conservation biol-
text of their communities, the bulk of money and effort ogy, in its modern form, emerged concurrently with the
consistently went toward enhancing populations of species concept of and concern for the preservation of biodiversity
with commercial or recreational value for humans (Soulé (Chap. 2). To engage such an overwhelming objective, con-
1985). servation biology has, from its inception, understood itself as
The 1960s and 1970s saw the development of major new synthetic, eclectic, and multidisciplinary. It did not draw all
ideas in population biology and community ecology, such as its theories and models from biology and it had the effect of
the theory of island biogeography. Many scientists began breaking down the dichotomy of pure versus applied science
testing the predictions of the new paradigms in the problems (Soulé 1985). With its emergence in the 1980s, conservation
of conservation. But the results of their experiments were not biology identified itself with seven distinctions that reframed
always appreciated in traditional applied sciences such as relationships between conservation science and conservation
wildlife ecology, fisheries management, forestry, and range practice (Table 1.2).
management. These disciplines had become The first and most important emergent distinction of con-
departmentalized in major universities and isolated from servation biology was its focus on the preservation of biodi-
one another. Their isolation led to alienation, and even hos- versity (the entire range of all species, their habitats, and their
tility. The natural exchange of ideas and infusion to phylogenetic lineages), not on the management of individual
applications in conservation problems was inhibited (Soulé species. Core disciplines that inform conservation biology’s
1986). attempts to achieve this goal are, according to Soulé, ecology,
Traditional academic disciplines in resource management systematics, genetics, and behavior. Related disciplines in the
were also limited in other ways. For the most part, they did applied sciences, such as wildlife ecology, fisheries manage-
not understand or effectively respond to the growing chorus ment, and forestry also draw much of their source data from
of voices in the developing field of environmental ethics that similar backgrounds. However, the latter fields have tradi-
claimed that all species, not merely game animals and fish, tionally selected subjects for research based either on com-
livestock, or plants of commodity value, possessed intrinsic mon characteristics or common applications for their
values, not merely utilitarian values. Conversely, active management. Conservation biology, in contrast, focuses on
conservationists (Leopold’s “B” group) were failing to infuse the study and preservation of the diversity of life itself.
their land ethic into resource management and the academy. Conservation biology’s second founding distinction was
Stress on Leopold’s A-B cleavage was increasing, and the that of understanding itself as a scientific discipline that was
pressure could only be relieved by a split. both value laden and value driven. Integral to conservation
biology’s identity was its explicit recognition of Michael
Soulé’s four “normative postulates” (Soulé 1985). The first
1.6.2 The Intellectual Inception was that diversity of organisms is good, and its negative
of Conservation Biology corollary that the untimely extinction of populations and
species is bad. Second, ecological complexity is good. The
Conservation biology has been described as a science of second postulate assumes the first, but adds value to the
scarcity and abundance, the “application of biology to the preservation of ecosystem diversity. Thirdly, evolution is
care and protection of plants and animals to prevent their loss good, or more precisely, that it is desirable to maintain the
or waste” (Meffe and Carroll 1997). Born out of the crisis of genetic potential of populations that permits adaptation and
Table 1.2 Seven core disciplinary distinctions of conservation biology

Core Distinctions Explanation that conservation biology. . .
Basis in preserving Focuses on the study and preservation of biodiversity, rather than the management of individual species
biodiversity
Value laden and value Is committed to valuing biodiversity, regardless of its utilitarian value
driven
Mission- and advocacy- Emphasizes intention and action to save species and habitats
oriented
Crisis-oriented Requires rapid investigation and response, even before risk or replication studies can be performed
Integrative and multi- Synthesizes information across disciplines (biology, ecology, ethics, politics, and other disciplines)
disciplinary
Concerned with Seeks preservation of genetic information and processes that promote speciation for future biodiversity, not just
evolutionary time conservation for present organisms
Adaptive Treats management options as experimental and imprecise, where outcomes may be risky or unpredictable
Table design by R. Lamb and F. Van Dyke
innovation in a changing environment. Soulé’s final postulate ecologists ‘live in a world of wounds’ (Leopold 1966), then
was that biotic diversity has intrinsic value, regardless of its conservation biologists could be said to perform triage daily
utilitarian value. Stating this as a normative value made it in a ward full of chronically hemorrhaging patients” (Beever
explicit that conservation biology, from its inception, was 2000: 907). Conservation biology must respond to emer-
committed to the study and understanding of all species and gency situations with incomplete information. Although
their relationships. Although conservation biology can be urgent, short-term goals in conservation change, the long-
described by its interest in biodiversity, it is defined by its term goal is the persistence and viability of functioning
commitment to the value of biodiversity. ecosystems (Soulé 1985). Conservation biology’s need for
The emergence of conservation biology as a formal scien- rapid investigation and response does not provide it with the
tific discipline has been interpreted as the mandate for con- luxury of long reflection and multiple replications of studies
servation in the scientific community, as well as a mandate before action must be taken. Most scientific disciplines have
for science to attempt the noble (but perilous) quest of viewed premature application of scientific results as worse
assimilating moral principles into scientific study and appli- than no action and have emphasized the importance of
cation. A foundational principle of conservation biology is minimizing risk and maximizing reliability. In conservation
that species possess intrinsic value as entities that ought to biology, however, failure to act when a population is declin-
exist and persist in the world. As conservation biologist John ing or a habitat is being degraded may ensure the extinction
Robinson explained, “. . .incorporating values into our sci- of the species or the loss of its environment. Conservation
ence is necessary and means that CB should build a case for biologists are more willing to tolerate the risk of inappropri-
the conservation of biodiversity, not disinterestedly ate action than the irreparable losses or environmental dam-
investigate. . .the consequences of biodiversity loss” age that may be associated with no action.
(Robinson 2006:661). Conservation biology is also different in its tendency to
A third distinction of conservation biology, strongly cross disciplinary lines. This is conservation biology’s fifth
related to the second, is that it has been historically mission- distinction, its integrative and multi-disciplinary nature.
and advocacy-oriented. In fact, conservation biology has Although rooted in the core discipline of biology, studies in
been explicitly defined as a mission-oriented discipline com- conservation biology routinely cross disciplinary boundaries
prising both pure and applied science (Soulé and Wilcox among major taxa, such as plants and animals, vertebrates
1980:1). This sense of mission in conservation biology is a and invertebrates, and between biological and physical pro-
natural consequence of being a value-laden discipline. Given cesses. Further, because it is value driven and mission ori-
the intrinsic value of species, conservation biology perceives ented, it routinely investigates and addresses issues of ethics,
that the best and highest application of scientific knowledge human behavior and culture, law, politics, and sociology with
about species is to ensure their preservation. But conservation the aid social science. By linking conservation initiatives to
biology does not confine this effort to research and manage- diagnostic tools, such as the social–ecological systems frame-
ment. Rather, many conservation biologists assert the impor- work, conservation scientists are able to better integrate
tance of scientists being able to communicate the social considerations into conservation initiatives, making
spontaneous inner experience and appreciation for the them more sustainable over the longer term (Moon et al.
creatures they investigate (Naess 1986), claiming that no 2014).
one has more expertise, or right, to express a love for nature A sixth distinction is that conservation biology is a science
than those who have given their lives to its study (Soulé concerned with evolutionary time. In its emphasis on the
1986). Indeed, environmental philosopher Arne Naess, in preservation of biodiversity, conservation biologists seek
his keynote address at the Second International Conference not merely the preservation of present types of organisms,
on Conservation Biology in 1985, told his audience of con- but the preservation of their genetic heritage (representing
servation biologists that they had “obligations to announce their evolutionary history) and the preservation of ecosystem
what has intrinsic value” (Naess 1986, emphasis his). As a processes that promote adaptation, innovation, and speciation
discipline, conservation biology’s emphasis on action to save to maintain and enhance future biodiversity. Geneticist
species and habitats and its declared intention to announce Richard Frankham has repeatedly suggested that the way in
the values of nature encourages, indeed often demands, that which conservation biologists define a species genetically
its practitioners act as focused advocates (Rohlf 1995) which will have a profound impact on the financial, legal, and
can be defined as a person or group reporting data concerning biological implications of their work (Pertoldi et al. 2007;
an area in which he or she has expertise as well as deeply held Frankham et al. 2012).
convictions, and who works to ensure that the information A seventh distinction is that conservation biology is an
presented is correctly interpreted and rightly applied. adaptive science. Although the recent paradigm of adaptive
A fourth distinction is that conservation biology is crisis- management is not unique to conservation biology, it is a
oriented. In the words of Aldo Leopold and Erik Beever, “If concept decidedly at home in this discipline. Compared to
more traditional, management-oriented disciplines in the life are a key but too often neglected dimension of conservation”
sciences that have tended to see management actions and (Kareiva and Marvier 2012:963).
their responses in a cause-and-effect relationship, conserva- Many of those active in conservation today would affirm
tion biology is characterized by its tendency to treat manage- these convictions by designating the discipline as conserva-
ment actions themselves as experiments. It has tended to both tion science, and by so doing explicitly recognizing that
expect and accept a higher degree of uncertainty associated ecological dynamics cannot be separated from human
with the response of a system or a population. The response is dynamics, and that as such the disciplines that inform and
then treated more as an experimental result than a perfectly define conservation science are different from the traditional
predictable outcome, and the management strategy itself is conception of the fields informing conservation biology
adaptively revised in light of the results obtained. (Fig. 1.18). Increasingly, conservation efforts are being com-
One cannot claim to be an adaptive science and cease to bined with efforts in poverty reduction (Robinson 2006) and
adapt, even when it comes to founding principles and conservation efforts like the Biodiversity in Development
distinctions. So it is in conservation biology, where now even Project (2001) have argued that conservation in the absence
the seminal principles that gave it birth and vitality are being of economic development will fail. The need for biological
transformed as conservation efforts affect and are affected by and conservation priorities to be assessed within economic
conditions in the world today. and sociopolitical context is not a new idea (Meine et al.
2006; Pressey and Bottrill 2008), but the proposed change for
the name of the discipline is not simply cosmetic. During its
emergence in the 1980s, most conservation biologists saw
Since the Enlightenment, Western science has
human development as the primary threat to biodiversity
presented itself as value neutral. Is being value neutral
preservation. Stopping such development to protect species,
an expression of objectivity or apathy in conservation?
primarily by setting aside areas to be kept in a near-pristine
Does a scientific idea become less true if one feels
state, was one of conservation biology’s most important goals
personally connected to and involved in the idea’s
as well as one of its most favored conservation strategies.
implications? Do you believe that conservation biology
Many conservationists today desire, in fact, insist, that
will continue to treat its “normative postulates” as
“conservation biology” make explicit recognition that it is
important elements in its work, or adopt a more
not concerned only with the welfare of non-human nature,
“value-free” approach characteristic of many other
but that a key goal of their work is the improvement of human
sciences? Why?
well-being through the management of the environment
(Robinson 2006; Kareiva and Marvier 2012). This perspec-
1.6.3 A Time of Transition: Protecting Nature tive has certainly shaped the development and establishment
from People to Protecting Nature of Indigenous Reserves, Extractive Reserves, and Biosphere
for People Reserves that we have previously discussed. But such con-
cern and consideration is not new to conservation biology.
Conservation biologists have long recognized that political, From its inception, one of the discipline’s distinctions was its
social, cultural and economic factors and influences had to be recognition that a close linkage had to be formed between
integrated into successful conservation efforts. However, biodiversity conservation and economic development, and
contemporary critics of the historic formation and conceptual that one of conservation biology’s goals would be to improve
development of conservation biology have charged that, in its that relationship. Furthermore, its founders recognized, from
traditional understanding, people played only one of two the beginning, that its work had to be fully engaged in policy
roles. Most people, in their ordinary activities of life, posed formation while still remaining a credible source of objective
threats to biodiversity preservation. A smaller number of scientific information (Meine et al. 2006). We have already
people (mostly western scientists) were the protectors or seen how insensitivity to human presence in any landscape
“saviors” of biodiversity, primarily through safeguarding it leads to both social injustice and environmental degradation,
from human intrusion and disturbance. Many conserva- and more examples will follow in subsequent chapters, espe-
tionists today want to make fundamental changes in this cially those concerned with anthropogenic influence
perception. As Peter Kareiva and Michelle Marvier express (Chap. 3), climate change (Chap. 4), conservation values
it, “conservation is fundamentally an expression of human and ethics (Chap. 10) and conservation economics and sus-
values. . .For better or worse, people’s attitudes and actions tainable development (Chap. 12).
help to shape and reshape the world that will be left behind Although these and other contrasts between “old” conser-
for future generations. Therefore, the psychology and ethical vation biology and “new” conservation science are some-
reasoning that underlie people’s actions and views of nature times overstated by contemporary conservationists, such
Fig. 1.18 (a) Fields contributing to conservation biology as originally effort to protect nature. (Reprinted by permission from Oxford Univer-
conceived and depicted by Soulé (1985) and (b) a more recent concep- sity Press, from Kareiva, P., Marvier, M., 2012. What Is Conservation
tion of fields contributing to “conservation science” by Kareiva and Science? BioScience 62, 962–969. # 2012 by American Institute of
Marvier (2012) intended to reflect a broader and more interdisciplinary Biological Sciences)
concerns are relevant given that the areas of the world with managed aquatic ecosystems and agroecosystems. But the
the greatest biodiversity are and will continue to be areas that role, place and size of novel and anthropogenic ecosystems
experience the highest rates of human population growth and has grown and now occupies a large place in the landscape of
the greatest increases in resource consumption following a global conservation efforts, as we will examine in detail in
“business-as-usual” approach. Therefore, many conserva- Chap. 3.
tionists today want to see increased attention to the protection
of and provision for ecosystem services, economic equity,
Synthesis
social justice, and human rights as integral elements of con-
Conservation biology is inexorably wedded to conser-
servation planning, precisely the kinds of concerns that led to
vation, a movement that is informed by science, yet
the design and formation of the first Extractive Reserves in
seeks normative ends and purposes as “good.” The
Brazil (Sect. 1.5.4). Kareiva and Marvier are, again, two
initial attraction of scientists to conservation biology’s
practitioners who express the conviction most strongly.
most explicit professional expression of itself, The
“Conservation as it was defined by Soulé is reactive and on
Society for Conservation Biology, was indicative of
the defensive; its goal is to minimize losses and, to the extent
the growing dissatisfaction scientists felt with being
that this is possible, to maintain the world as it once was”
mere “informers” about the state of the biodiversity
(Kareiva and Marvier 2012:967). Such a description is more
crisis instead of actors who could do something about
harsh than accurate, but does convey the strength of convic-
it. To be able to persist as a distinct discipline, conser-
tion held by some conservationists about the need to move
vation biology needed more than a crisis in biodiversity
conservation efforts towards greater integration with human
and a few new scientific ideas. To survive and grow, a
welfare. It also correctly expresses the perception that effec-
discipline requires a unique conceptual framework and
tive conservation today will often take place within human-
a set of identifiable intellectual distinctions. But it also
altered and human-inhabited landscapes, and so must be
needs human relationships formed not only out of
prepared to operate in novel ecosystems shaped by anthropo-
mutual interest, but in shared hope that collective
genic influences (Kareiva and Marvier 2012). This perception
aspirations can actually be realized, aspirations which
also is not a new idea in conservation biology, as, from its
are in each members interest to achieve. “Disciplines,”
inception, founding members of the discipline were engaged
in defining biodiversity values on private lands, as well as in (continued)
noted Michael Soulé, “are not logical constructs; they that they can succeed. As conservation biologist Eric
are social crystallizations which occur when a group of Beever noted, conservation biologists must give hope
people agree that association and discourse serve their to others “. . .because the success of conservation
interests. Conservation biology began when a critical efforts depends on how we are perceived by decision-
mass of people agreed that they were conservation makers and the public at large. Although we must alert
biologists,” (Soulé 1986:3). these groups to impending ecological challenges, we
The roots of conservation historically have been must also give them reasons for hope. If we do not,
grounded in moral arguments about the intrinsic value resistance to conservation biology and a negative per-
of nature, the proper response of humans to nature, and ception of its practitioners could marginalize our efforts
the rights and needs of future generations to enjoy at all scales. . .” (Beever 2000:907). Yet, conservation
nature and the resources it provides. Although the suffers under a cloud of hopelessness, and the problem
conservation movement dare not be ignorant of the of hopelessness in conservation, and particularly
best scientific information and insight, conservation among conservation biologists, is widely perceived.
biology dare not be ignorant of its own origins, and Conservation biologists Ronald Swaisgood and James
the motivations of individuals like Gilbert White, John Sheppard have argued that there is currently a culture
Muir, Theodore Roosevelt, Aldo Leopold, Paul of hopelessness among conservation biologists that
Sarasin, Michael Soulé, and others who, although compromises our ability to mobilize conservation
well informed of scientific theories that could make action among the general public and draw others into
conservation effective, also understood logical and the conservation noble field (Swaisgood and Sheppard
moral arguments that made conservation worth doing. 2010). They write, “. . .people must believe they can
Conservation biology is a product of such past and exercise some control over the situation – in effect,
continuing efforts and must be presented and under- they must believe they are empowered to make a dif-
stood as a unity of facts, theories, and values linked by ference” (Swaisgood and Sheppard 2010:629). Conser-
common purpose. Indeed, the ecological economist vation biologists must believe this just as strongly, in
Herman Daly rightly called conservation “a policy in fact, more strongly, than “ordinary” people do, and
the service of a purpose” (Daly 1999:694) and we will both must be empowered by such hope. As conserva-
not dispute his insight. Purpose must exist if we are to tion scientist and environmental ethicist Kyle S. Van
experience value in conservation work and communi- Houton has noted, “Conservation needs a cultural legit-
cate that value to others. Therefore, conservation again imacy that inspires enthusiasm, allegiance, and per-
resolves fundamentally to a moral endeavor. Scientific sonal sacrifice – in other words, actual changes in
knowledge and expertise support and empower moral human behavior. Such a vision does not provide a
purpose. They cannot replace it. straight line path to easy answers, it offers a description
Conservation’s increasing emphasis on integrating of ethics currently estranged from conservation sci-
biodiversity conservation with human welfare is not ence” (Van Houtan 2006:1371).
wrong, but it is a complement to, not a replacement Although there is much hard work to be done to
for, appreciation of and affection toward the natural stem the tide of species extinctions, ecological col-
world in its own right. Conservation cannot be accom- lapse, and a warming climate, conservation biologists
plished without moral and economic restraint. No one can play a crucial role in regularly confronting and
can benefit from or control what they refuse to limit. overcoming despair. There is joy in conservation suc-
The benefits of a healthy environment, and the joy of its cess, and it is in the hope that long-term solutions to
vibrant and diverse life, are rewards that come to those these and other current problems are possible that con-
whose limits make room for the lives of other kinds of servation biologists work so tirelessly to achieve it. The
creatures in their world. chapters that follow will identify the controlling ideas,
The work of conservation biology, and its unifying scientific theories and definitive studies that today char-
purposes, must also be emboldened by hope. Psychol- acterize and define conservation biology and its efforts
ogy, the science that is perhaps best qualified to under- to preserve biodiversity at genetic, population, and
stand the bases and motivations of human hope, is not system levels. Within their content, we will present a
just a tool for conservationists to apply to solution-oriented framework for conservation engage-
non-conservationists to get them to believe that conser- ment that outlines the knowledge, tools, applications
vation can succeed. They must also apply psychologi- and contexts needed to engage conservation challenges
cal understanding to themselves if they are to believe with perseverant hope. We are convinced that this is the
(continued) (continued)
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Biodiversity: Concept, Measurement,
and Management 2
But now there is something new, never before seen on Earth during its billions of years of evolving species.
Humans have begun to set conservation of the biodiversity on Earth as a moral and social goal.
Holmes Rolston III (2012)
Keywords of natural habitats to systems designed to maximize human

Biodiversity conservation · Species concept · Species benefit; their use of energy and its resulting waste products;
diversity · Endemism · Surrogate species · Biodiversity their high rate of population growth; their ability to move
distribution · Biodiversity hotspots · Protected areas · themselves, and all sorts of plants and animals, from one
Functional conservation areas · Rarity · Conservation region of the world to another; their capacity and supporting
planning technology to harvest (kill) large quantities of plants and
animals in a short time; their industrial-level production of
many substances that (unintentionally) destroy many kinds of
life; and their effects on changing the global climate.
Overview Solving a problem with these root causes is overwhelm-
In this chapter you will learn about the: ingly daunting, but the attempt to stop biodiversity loss has
been a foundational motivation for the practice of conserva-
1. Concept and definition of biodiversity in conservation biology from its inception. Indeed, the field of conserva-
tion biology. tion biology now understands its two major missions to be
2. Relationship between biodiversity and ecosystem that of conserving the biodiversity of the Earth and
function. safeguarding the benefits (ecosystem services) that intact
3. Current state of world biodiversity and causes of ecosystems provide to human welfare through their func-
biodiversity loss. tional operation (i.e., through what scientists refer to as
4. Methods of understanding, valuing, and measuring ecosystem properties). Reflecting on conservation biology’s
biodiversity. beginnings, Michael Soulé noted that an increasing percep-
5. Global distribution of biodiversity and global tion of an accelerating and global loss of species – the
patterns of species abundance. “extinction crisis” – was a major factor in conservation
6. Methods for prioritizing, preserving and managing biology’s emergence as a distinct discipline (Soulé 1986:4).
biodiversity. This shift in focus among conservation biologists from the
problem of “endangered species” to the problem of “loss of
biodiversity” might, at first, sound like an exercise in
biological semantics, but the change in emphasis is
2.1 Biodiversity and Conservation Biology significant.
Conservation biology’s historical origins were informed
One of the most pervasive environmental changes of recent
by discoveries in basic sciences such as genetics and popula-
history is the global loss of the Earth’s diversity of life
tion demography but began to be manifest in practice in
(Fig. 2.1). Biodiversity loss is a systemic problem embedded
applied sciences like forestry, wildlife management, fisheries,
in the “normal” ways that human beings live on planet Earth,
and range management. The latter disciplines – emerging as
including and especially their conversion and manipulation

36 2 Biodiversity: Concept, Measurement, and Management
distinctive professional communities from the 1930s through simply as a tragic parade of passing species, but as losses in
the 1950s – were species-specific in their approach to man- genetics, community attributes, and ecosystem properties.
agement and understanding of species’ values. In these With changes in perception of what was being lost came
disciplines, studies of species’ natural history and habitat changes in perceptions of why species were being lost. New
requirements received priority. Resource management research shifted from studying natural history to identifying
sciences began to influence environmental law in the 1960s ecological and evolutionary processes contributing to
and 1970s, and environmental legislation drafted in this emerging patterns of extinction. The extinction crisis created
period reflected a similar emphasis. The US Endangered an urgency to develop an alternative concept to that of
Species Act, with its stress on individual species as primary “endangered species.” That concept was biodiversity.
targets of conservation efforts, is one of the best, but not the
only, example of this type of legislation (Fig. 2.2). Other US
examples, such as the Sea Turtle Act and the Marine 2.2 Biodiversity and Ecosystem Function
Mammals Act, displayed a similar focus.
The emergence of conservation biology from such applied In a special report addressing the role of biodiversity in
management sciences reflected a shift in emphasis and a ecological systems, a committee of 15 distinguished
break with historic perceptions about the nature of the ecologists informed the Ecological Society of America
“endangered species problem.” The traditional view of recent (ESA) that, despite controversy and knowledge gaps charac-
extinctions as a collection of tragic, individual case histories teristic of scientific experimentation, some conclusions about
was replaced by a conviction that the global extinction crisis biodiversity could be asserted. Specifically, some
was caused by fundamental disruptions of ecosystem pro- combinations of species (i.e., biological communities) are
cesses. Extinctions, in this perspective, were not perceived complementary in their patterns of resource use and can
increase average rates of productivity and retention of
nutrients, both of which decline when species are lost from
the community (Hooper et al. 2005). Such increased produc-
tion varies less through time in these systems than in systems
with fewer species (Duffy 2009). Systems with many species
produce more biomass and capture more resources than those
with only a few species, even at different trophic (feeding)
levels and in different habitats (Wilsey and Potvin 2000;
Duffy 2009).
Enhanced biodiversity can also lead to increased system
stability. For example, the greater the diversity of a plant
community, the lower the success rate of invasive plant
species attempting to enter it (Tilman et al. 1997; Naeem
et al. 2000). A greater diversity of species with different
sensitivities to a range of environmental conditions usually
Fig. 2.1 The Monteverde Golden Toad (Incilius periglenes), a species leads to greater stability of ecosystem properties and a greater
indigenous to the cloud forests of Central America, where it has
number of functional ecosystem processes (Tilman et al.
disappeared in the wild during a period of recent climate change and
loss of its critical habitat. (Photo by Charles H. Smith, US Fish and 1997; Naeem et al. 2000) (Fig. 2.3), such that even rare
Wildlife Service, under public domain) species can make important contributions to stability,
Fig. 2.2 The Black-footed Ferret

(Mustela nigripes), a species
nearly exterminated in the wild,
but restored with the support and
protection of the US Endangered
Species Act through captive
breeding and subsequent
reintroduction. (Photo by Ryan
Moehring, US Fish and Wildlife
Service, public domain)
2.2 Biodiversity and Ecosystem Function 37
especially in changing environments (Lyons et al. 2005). In safeguard against risk resulting from changes in environmen-
experimentally established plant communities, plant cover, tal conditions or, in agricultural systems, market demands,
biomass, and productivity decline as species richness because it extends the productive use of a site’s resources
declines (Tilman et al. 1997; Wilsey and Potvin 2000). In over time and provides an effective way to provide multiple
contrast, below ground biomass as well as total biomass not goods and services (Hooper et al. 2005). But this is not an
only increase as species richness increases, but also as species isolated opinion of one committee in one scientific society.
evenness increases (Wilsey and Potvin 2000) (Fig. 2.4). The There is now widespread consensus that “biodiversity loss
ESA committee concluded that maintaining biodiversity is a reduces the efficiency by which ecological communities cap-
ture biologically essential resources, produce biomass, [and]
decompose and recycle biologically essential nutrients”
(Cardinale et al. 2012:60).
These and other effects on ecosystem functions and
properties affect the amount and quality of ecosystem
services humans can derive from ecological systems. From
a human perspective, ecosystem services are usually placed
in four categories. Provisioning services refer to the
ecosystem’s production of materials humans can use for
food, energy, or structural material. Regulating services are
those that reduce variability and severity of environmental
events, such as a wetland upstream from a human community
that lowers downstream discharge, which in turn reduces
severity and frequency of flooding. Supporting services are
the ecosystem’s provision of physical “platforms,” as well as
essential processes, for human activity, such as soil to grow
crops, oxygen produced from photosynthesis, or a pathway
of transportation for commercial good provided by a naviga-
ble river. Cultural services are the ecosystem’s furnishing of
opportunities for recreational, intellectual and spiritual
renewal, and for acquisition of knowledge (through both
casual experience and scientific inquiry).
Fig. 2.3 In experimental studies of grassland ecosystems, the number Loss of biodiversity reduces the capacity of ecosystems to
of species needed to maximize ecosystem functioning increases with the provide these services. Just as the amount, efficiency, and
number of ecosystem processes considered. (Reprinted by permission
from Wiley, from Duffy, J.E., 2009. Why biodiversity is important to quality of ecosystem functions increases with increasing
the functioning of real-world ecosystems. Frontiers in Ecology and the numbers of species in the ecosystem (biodiversity), so
Environment 7, 437–444. # 2008 Ecological Society of America) decreases in a community’s number of species are associated
Fig. 2.4 Both total and belowground biomass in plant communities Biodiversity and Ecosystem Functioning: Importance of Species Even-
increase as species evenness increases, evidence that biodiversity ness in an Old Field. Ecology 81, 887–892. # 2000 Ecological Society
contributes to an increase in ecosystem productivity. (Reprinted by of America)
permission from Wiley, from Wilsey, B.J., Potvin, C., 2000.
with lower productivity and ecological efficiency (de Baan makes these values of biodiversity explicit by stating, “Bio-
et al. 2013; Aerts and Honnay 2011). Today more than one diversity is essential for food security and nutrition and offers
billion people on Earth rely on wild-harvested products for key options for sustainable livelihoods.”
nutrition and income (Sunderland 2011). As a result, human There are many kinds of human activities, however,
life can be impoverished by biodiversity loss, and the effect is practiced in appropriate ways, that can enhance biodiversity
greater among people of lower incomes who are more (Table 2.1). Agriculture and forestry are among these, as
directly dependent on goods and services of biodiversity for observed in the enhanced biodiversity associated with the
their subsistence. For example, 80% of rural populations in kinds of agriculture and forestry practiced worldwide by
Africa depend on products harvested from local biodiversity farmers with relatively small land holdings. When these
(Pelser and Letsela 2012) and 30–80% of protein intake of systems employ polyculture cropping (growing many crops
rural communities worldwide comes from harvest of wild together on the same site), diverse agroforestry, and mainte-
animals (Fa et al. 2003). But no household, rural or urban, nance of high levels of variety (heterogeneity) in surrounding
can be food secure if its methods of food gathering deplete natural vegetation, the land can support more species than
sources of natural capital needed to produce future food monocultures or homogeneous land uses, and is better buff-
supplies (Sunderland 2011). ered against market fluctuations (Sunderland 2011; Trimble
Biodiversity also is a primary source of medicinal goods and van Aarde 2014). Such diverse production systems pro-
in these communities, with the World Health Organization vide enhanced nutritional benefits and improved livelihoods
estimating that up to 80% of the population in some countries and are an important path to food security for those who
relies on directly-obtained plant and animal products from practice them (Sunderland 2011).
surrounding natural systems for primary health products and As plant agriculture can be managed to enhance biodiver-
care. Loss of biodiversity also has been linked to increased sity, so can pastoral agriculture. In sub-Saharan Africa, many
emergence and transmission of infectious diseases (Keesing pastoral economies, practiced in traditional ways, maintain
et al. 2010) and commercial (“industrial”) agriculture often abundant populations of wild herbivores surrounding domes-
creates or leads to loss of species diversity and ecosystem tic grazing animals (Kinnaird and O’Brien 2012; Trimble and
function (Sunderland 2011). In contrast, agricultural systems van Aarde 2014). Mixed grazing by domestic stock carefully
which enhance biodiversity, both in cultivated crops and controlled by land managers can maintain or create
associated crop-dependent animal species, are more resilient specialized conditions of vegetation structure and function
to environmental change, including climate change, and eco- (in other words, specialized niche spaces) which can help
nomically more resistant to market fluctuations in crop and maintain native plant and animal biodiversity (Pollock et al.
commodity prices, (e.g. Sunderland 2011; Liebman et al. 2013). Together, traditional agricultural and pastoral mosaics
2013). The major international treaty for biodiversity conser- of land use have been shown to be beneficial to many wild
vation, the Convention on Biological Diversity (CBD), species associations (Ratcliffe and Crowe 2001; Konečný
Table 2.1 Human practices that support biodiversity in human modified landscapes. PA ¼ protected areas
Practices that tend to support species diversity and richness in human-modified landscapes
Prefer diversity in selection of crops grown (i.e. polyculture) and land use (i.e. land-use mosaics, over homogenous monocultures)
Encourage traditional agricultural practices over large-scale, mechanized farming
Leave as much remnant natural vegetation as possible and monitor or assist maintenance of keystone structures or species, e.g. large trees
Ensure strict protection for specialist and endemic species and expand PA coverage focused on these groups
Encourage appreciation and understanding of conservation goals among land users
Discourage urban sprawl and maintain and manage urban green spaces
Favor use of native species in gardens and cultivation
Avenues for further investigation into scientific uncertainties and implementation practices
Researching poorly documented combinations of species group, ecosystem type, and land use, e.g. mammals in rangeland agroforests
Moving beyond occurrence data to likelihood of persistence, e.g. how dependent are species in human-modified landscapes on nearby PAs or
remnant habitat?
Investigating the value of reintroduction or rewilding in human-modified landscapes
Going beyond the species level of biodiversity to integrate genetic and ecosystem concepts
Supporting technological and traditional knowledge for cultivating useful native species and investigating the effects of these practices on other
taxa
Creating frameworks for valuing the importance of different species in different landscapes at a local level within a global context,
e.g. commonness versus rarity and specialist versus generalist
Developing policies that integrate and account for local, regional, and global conservation needs and land use systems
Reprinted by permission from Springer Nature, from Trimble, M.J., and van Aarde, R.J., 2014. Supporting conservation with biodiversity research
in sub-Saharan Africa’s human-modified landscapes. Biodiversity and Conservation 23, 2345–2369. # 2014 Springer Science+Business Media
Dordrecht
2.3 Is Conservation Effort Saving Biodiversity? 39
et al. 2010). Thus, biodiversity can be conserved, not only by motions passed by simple majorities against strong opposi-
protecting natural areas, but by encouraging protection of tion, although legally valid, in practice lack legitimacy and
natural vegetation in “unnatural areas” like agricultural authority, especially with dissenting parties” (Stuart et al.
landscapes (Trimble and van Aarde 2014). 2017:2) Through such patience and persistence, IUCN has
developed a reputation for integrity in its science (IUCN’s
“technical” role) as well as in its ability to convene the
2.3 Is Conservation Effort Saving conservation community to address complex and controver-
Biodiversity? sial issues (IUCN’s “convening” role). Through its expert-
driven global consultation processes (its technical role),
2.3.1 Conservation Governance – The IUCN IUCN has developed internationally accepted standards,
and Global Biodiversity Conservation including the IUCN Red List Categories and Criteria (for
species), the Protected Area Management Categories and
The well-known proverb of US political culture, “All politics the IUCN Red List of Ecosystem Categories, among others
is local,” can be applied with equal truth to conservation. (Information Box 2.1). And, in its convening role, IUCN has
Conservation efforts to save rare, often endemic species with led the conservation community to develop major policy
small populations, specialized habitats, and restricted ranges statements and positions on key conservation issues.
must, of necessity, be grounded in local expertise and Among these, the IUCN Red List and Red List Categories
community-based support if they are to have any hope of and Criteria, and its more recently formulated Red List Index
success. Is there any value in even attempting to form a (RLI) are critical benchmarks which deserve further attention
“coordinated global strategy of conservation”? Despite obvi- if we are to understand the status of global biodiversity
ous and inherent difficulties of such a quixotic venture, a conservation today.
world conservation strategy can be said to exist, or, at least,
to be in the process of development, and its effects are
Information Box 2.1: The IUCN and Global
pervasive in conservation efforts.
Standards in Conservation
The acknowledged voice and leader in global conserva-
Through its long-established credibility in scientific
tion is the International Union for the Conservation of Nature
expertise, as well as its unique capacity to bring
(IUCN). Founded in 1948 (Chap. 1), IUCN declares its
together the world conservation community on matters
vision to be that of creating “a just world that values and
of global importance to the protection of biodiversity,
conserves nature”, and states that its mission is to “influence,
the International Union for the Conservation of Nature
encourage and assist societies throughout the world to con-
has contributed published standards and protocols that
serve the integrity and diversity of nature and to ensure that
are now used and followed throughout the world.
any use of natural resources is equitable and ecologically
Students and practitioners need to know about, appre-
sustainable” (Stuart et al. 2017:2). Among thousands of con-
ciate, access and effectively use such resources to work
servation organizations active worldwide, IUCN is unique in
effectively in conservation at any scale, and especially
its membership of representatives from both national
on issues which may involve multiple nations,
governments and non-governmental organizations (NGOs).
governments, and conservation NGOs. Here are some
Such representatives meet every 4 years in the World Con-
of the IUCN’s most important and currently used
servation Congress, a forum established by IUCN to address
contributions.
the state of conservation worldwide. Supported by a perma-
nent staff of nearly 1000 paid employees placed at more than
• Dudley, N. (ed.) 2008. Guidelines for Applying
50 locations globally, as well as thousands of unpaid volun-
Protected Area Management Categories. IUCN,
teer experts, IUCN fulfills a unique place in the conservation
Gland, Switzerland. WITH Stolton, S., P. Shadie
community in both international influence and scientific
and N. Dudley. 2013. IUCN WCPA Best Practice
respect. Although having no explicit authority to make con-
Guidance on Recognising Protected Areas and
servation “laws” binding on any nation, its recommendations
Assigning Management Categories and Governance
and agreements among its members often come to have the
Types, Best Practice Protected Area Guidelines
force of law as international standards for conservation prac-
Series No. 21, Gland, Switzerland. Available at
tice (Stuart et al. 2017). Since 1950, its World Conservation
https://portals.iucn.org/library/node/30018
Congress has adopted more than 1000 motions which have
• IUCN. 2012. IUCN Red List Categories and
become either Resolutions (directed toward the IUCN itself)
Criteria: Version 3.1. 2nd edition. IUCN, Gland,
or Recommendations (directed to third parties). Although a
Switzerland, and Cambridge, UK. Available at
simple majority is enough to adopt motions, Simon Stuart,
https://www.iucnredlist.org/resources/
former Chair of the IUCN Species Survival Commission, and
redlistguidelines
other IUCN representatives explain that IUCN members
“strive to get as close as possible to consensus [because]. . . (continued)
Wild) (Fig. 2.5). Twenty percent (one in five) vertebrate

Information Box 2.1 (continued) species are now classified as threatened, ranging from 13%
• IUCN. 2016. A global standard for the identification of birds to 41% of amphibians (Fig. 2.5). Michael Hoffman, a
of Key Biodiversity Areas. Version 1.0. 1st edition. member of IUCN’s Species Survival Commission, and his
IUCN, Gland, Switzerland. Available at https:// colleagues determined that, in all vertebrate groups, the aver-
portals.iucn.org/library/node/46259 age Red List Index (RLI) of individual species declined
• Bland, L.M., Keith, D.A., Miller, R.M., Murray, N. 0.49% from 1988 to 2008. Although this percentage seems
J. and Rodríguez, J.P. (eds.). 2017. Guidelines for small, it is equivalent to 52 vertebrate species moving one
the application of IUCN Red List of Ecosystems category closer to extinction every year. Over the entire
Categories and Criteria, Version 1.1. Gland, assessment period, 662 amphibians, 223 birds, and
Switzerland. Available at https://portals.iucn.org/ 156 mammals all moved one category closer to becoming
library/node/45794 extinct (Information Box 2.2). Only 7% of listed species saw
improvement, almost all of these involving mammals and
birds. In contrast, 13% of “declines” ended in extinctions
2.3.2 The Current Status of Species (Hoffman et al. 2010). More recent analysis (Rodrigues
Biodiversity et al. 2014) demonstrated that most of the world’s countries
had contributed negatively to these patterns, although there
In one of the most exhaustive assessments of the effects of was considerable variation within regions, just as there were
biodiversity conservation in recent years, 174 conservation important regional differences in what constituted the most
scientists reviewed the status of the world’s vertebrates using important regional threats. Nevertheless, the pattern of
the IUCN Red List, the globally authoritative assessment on decline in vertebrate species remained negative, and more
the conservation status of these species (Hoffman et al. than 50% of the global deterioration in conservation of birds,
2010). The Red List uses six categories to describe a species, mammals, and amphibians was concentrated in <1% of the
from “Least Concern” (LC) to “Extinct” (EX). In between the Earth’s surface area, 3.6% (39 of 1098) of total ecoregions,
extremes are categories that classify various levels of and 4.1% (8 of 195) countries (Australia, China, Colombia,
“Threatened” (Near Threatened, Vulnerable, Endangered, Ecuador, Indonesia, Malaysia, Mexico, and the United
Critically Endangered) or subcategories of “Extinct” (Possi- States) (Fig. 2.6).
bly Extinct, Possibly Extinct in the Wild, or Extinct in the
Fig. 2.5 (a) Trends in the Red List Index (RLI) for the world’s birds, threatened species from further decline, such actions failed to arrest the
mammals and amphibians. (b, c, d) Observed change in the RLI for each overall downward trend in conservation status, reflecting an increasing
respective group (black line) compared to expected trends in RLI if number of threatened species worldwide. (Reprinted by permission from
species that saw an improvement in status due to conservation action the American Association for the Advancement of Science, from
had undergone no change in status (red line). RLI values range from 1.0 Hoffmann, M., Hilton-Taylor, C., Angulo, A., Böhm, M., Brooks, T.
(Least Concern) to 0.0 (extinct). A downward trend in RLI indicates an M., Butchart, S.H., Carpenter, K.E., Chanson, J., Collen, B., Cox, N.A.,
increasing net rate of species extinctions. Shading reflects 95% confi- 2010. The impact of conservation on the status of the world’s
dence intervals. Note that, although conservation actions often prevent vertebrates. Science 330 (6010), 1503–1509 # 2010 AAAS)
Information Box 2.2: Amphibians on the Brink Information Box 2.2 (continued)
of Extinction Causes of amphibian decline are varied, but include
Declines in amphibian populations began to receive over-exploitation, habitat degradation and loss, disease
notice in the 1970s in the western United States, Puerto (especially the fungal disease chytridiomycosis) and
Rico, and northeastern Australia, and had become an “enigmatic declines” in which causes are not well
issue of global concern by the late 1980s, with the First understood (Stuart et al. 2004). Although global,
World Congress of Herpetology in 1989 noting the the worldwide decline of amphibians is not random.
international nature of the problem. Amphibians are Neotropical species are particularly affected, and
notorious for wide fluctuations in annual populations, amphibians in Australia and New Zealand have many
but tests of probabilistic null models showed declines more species suffering enigmatic declines than other
to be more severe and widespread than expected in regions. Over-exploited species are concentrated in
normal variations. Today amphibians are more East and South-east Asia, while reduced habitat species
threatened than birds or mammals, with their 1856 are prominent in South-east Asia, West Africa, and the
globally threatened species representing nearly Caribbean (Information Box Fig. 2.1). Declines are
one-third (32.5%) of all amphibian species. There are also nonrandom taxonomically. Four families are dis-
2468 species (43.2%) experiencing population declines proportionately affected, including Bufonidae (true
and only 28 (0.5%) showing population increases. toads), Leptodactylidae (typical Neotropical frogs),
Such declines are occurring across all habitat types Rheobartachidae (gastric-brooding frogs) and
and, in areas like Central America, are elevation- Ambystomatidae (mole salamanders).
dependent, with the greatest declines occurring at the More recent surveys have shown salamanders to be
highest elevations. Here climate change is having its particularly affected. At sites that have been repeatedly
most pronounced effects on temperature, precipitation, sampled for many years in Guatemala and Mexico
and humidity, creating moisture-scarce environments there were two recorded extinctions and major declines
which place terrestrial amphibians under stress because of formerly abundant species, especially salamanders
they depend on free standing water for drinking, and that were terrestrial habitat specialists. Large declines
because of their permeable skin, through which they were also seen in high elevation species of the “cloud
respire. forest” (mountaintop forests of Central America which
Information Box Fig. 2.1 Global geographical pattern of domi- American Association for the Advancement of Science, from Stuart,
nant causes of rapid decline in amphibian species due to overexploi- S.N., Chanson, J.S., Cox, N.A., Young, B.E., Rodrigues, A.S.L.,
tation (shades of blue), reduced habitat (shades of green), and Fischman, D.L., Waller, R.W., 2004. Status and Trends of Amphib-
unknown causes (shades of red). Darker colors indicate larger num- ian Declines and Extinctions Worldwide. Science 306, 1783–1786.
bers of rapidly declining species. (Reprinted by permission from The # 2004 AAAS)
has yet been significantly affected by conservation initiative.

Information Box 2.2 (continued) For example, for every 10 species suffering deterioration in
typically experience high levels of humidity, forming conservation status from expansion of agriculture, less than
“clouds” of water vapor at their highest elevations), one has improved because agricultural expansion was
several of which had entirely disappeared (Rovito reduced or mitigated (Hoffman 2010). Even “successes”
et al. 2009). The disappearance of “cloud forest” spe- associated with recovering species are unlikely to be
cies has coincided with decreasing humidity and longer maintained without continuing conservation effort, creating
dry periods in these cloud forests, implicating climate a growing category of “conservation reliant species” which
change as a reason for their extirpation. Unless the require ongoing management effort to persist even after they
causes of such declines can be better understood and are no longer considered endangered. Why is current conser-
actions taken quickly to address them, amphibian vation effort failing?
conservationists expect hundreds of amphibian species
to disappear during the next 20–30 years (Stuart et al.
2004). 2.3.3 What Causes Biodiversity Loss?
Many articles and textbooks on conservation biology provide

lists of things that reduce global biodiversity, but analysis is
Despite such pessimistic findings overall, analysis of the
better than list-making when it comes to understanding why
same information, with other data, does indicate that conser-
some factors are more important than others. The two factors
vation effort makes a difference. Hoffman and his colleagues
most responsible for global loss of biodiversity are modifica-
moved beyond tracking changes in the RLI to estimating the
tion of habitat and climate change, both caused primarily by
effect of “no conservation action” on 148 species of wild
human activity, and each of which will subsequently receive
ungulates (hooved mammals) as a “test group” by
attention in its own chapters (Chaps. 4, 7, and 8) in this text,
constructing “counterfactual scenarios” in which the conser-
but both factors reduce the value of biological habitat. Some-
vation laws, policies, and practices undertaken to protect
times referred to as “habitat loss,” the problem is better
these species were removed. From their analysis, Hoffman
described as “degradation,” a reduction in the functional
and his colleagues determined that, without conservation
and structural quality of habitat. Human activities do not
action, “the overall decline in the conservation status of
always destroy habitat, but they often change the structural
ungulates would have been nearly 8 times worse than
and functional qualities of habitat in ways that cause many
observed, corresponding to an average of 12 species
species to be unable to persist in the habitat’s altered
deteriorating by one Red List category per year from 1996
condition.
to 2008 (compared with the observed average of fewer than
Humans impose three kinds of effects on landscapes that
2 species that actually did undergo a deterioration over time).
degrade habitat (de Baan et al. 2013). First come transforma-
Nearly half of all species would have deteriorated in status
tional effects that change the way land is used, as when a
between 1996 and 2008 and 30 would have deteriorated by
prairie is cultivated to produce soybeans. The land is still
2 or more categories. . .” (Hoffman et al. 2015:1309). Con-
present, and the system may be producing as much or more
servation initiatives that made the greatest difference to the
plant biomass than it did before, but its ecological structure
largest number of species were “non-targeted” initiatives,
and processes are simplified. Such simplification reduces
especially habitat protection, rather that actions directed to
opportunity for different species to use resources on the site
particular species to protect them from specific threats (such
in different ways (i.e., to occupy different niches), so most
as overhunting). Despite uncertainty (more honestly, guess-
species are lost in the conversion. But changing land from a
work) in constructing the counterfactual, “no conservation”
natural and functioning plant community to an agricultural
scenarios, the analysis of Hoffman et al. indicates that con-
system has been a common cause of species loss not only
servation efforts matter. But they do not matter enough to
directly through changes in plant structure and function, but
stop the worldwide decline in biodiversity.
often in surprisingly indirect ways. For example, in the Baja
So, despite increases in the amount of land within
region of Mexico, a narrow peninsula of land with salt water
protected areas and adoption of conservation legislation in
(the Pacific Ocean) surrounding it, unsustainable use of
many countries, the state of world biodiversity continues to
groundwater for agricultural development has lowered
deteriorate. For all species, improvements are outnumbered
groundwater levels to the point that groundwater aquifers
by declines because no current conservation actions are stop-
have become susceptible to salt water intrusion. Such intru-
ping, or even slowing, the major drivers of extinction risk. On
sion has resulted in the loss of 22 native species, including
a global scale, none of these drivers (habitat degradation,
13 rare plants, and most species dependent on or associated
climate change, exotic invasions, overharvesting, and others)
Fig. 2.6 Relationship between

each country’s responsibility to
conservation and its contribution
to changes in species’ global
conservation status.
Responsibility measured by: (a)
weighted endemism; (b) weighted
threat. Dashed lines are regression
lines fitted through the origin: (a)
R2 ¼ 0.57; (b) R2 ¼ 0.70 (n ¼
195; p < 0.001). Circle size is
proportional to each country’s
Gross Domestic Product based on
purchasing-power-parity per
capita GDP in 2009. Circle color
indicates the main threat
(or combination of threats) for
selected countries. (Reprinted
under Creative Commons
Attribution, from Rodrigues, A.S.
L., Brooks, T.M., Butchart, S.H.
M., Chanson, J., Cox, N.,
Hoffmann, M., Stuart, S.N., 2014.
Spatially Explicit Trends in the
Global Conservation Status of
Vertebrates. PLoS ONE 9,
e113934. # The Authors)
Fig. 2.7 Box and whisker plot of relative species richness per land use any vegetation on a site formed as a consequence of human impact on the
type, number of data points (n) per land use type, and test-statistics site. “Used forest” refers to forests subject to multiple uses including
(Mann –Whitney U test) of pairwise comparisons of each land use with logging, grazing, or development which can alter forest structure and
reference value of species richness (global average across all biomes and vegetation. Note that, in every category of affected land use, species
taxonomic groups). “Second. vegetation” (secondary vegetation) refers to richness is lower than the reference value. (Reprinted by permission
vegetation associated with secondary succession on a site after a distur- from Springer Nature, from de Baan, L., Alkemade, R., Koellner, T.,
bance event that removed “primary” vegetation more typical or more 2013. Land use impacts on biodiversity in LCA: a global approach. Int J
stable on such sites. More generally, the term may be used to describe Life Cycle Assess 18, 1216–1230. # 2012 Springer-Verlag)
with riparian (streamside) habitat, ponds, and seasonal pools The problem of global biodiversity loss is increasing,
have disappeared from the region (Vanderplank et al. 2014). systemic to all taxonomic categories of life, human-driven,
Humans impose an occupational effect by living on land and difficult to solve. But before we attempt to discover a
previously unoccupied by people. The presence of human solution, we must first better understand the concept of bio-
beings, the resources they remove, the wastes they deposit, diversity itself.
and their patterns of activity leave other species without
needed resources (or the humans collect the species as
resources), exposed to human wastes that are useless or, 2.4 The Problem of Concept: What Is
worse, toxic to them, or the species are repelled by threats Biodiversity?
caused by human activity itself.
Humans impose permanent effects when they disturb an 2.4.1 A Conceptual Definition of Biodiversity
ecosystem in ways from which it cannot easily recover. A
grassland converted to a soybean field may, with proper The first use of the term biodiversity in scientific literature
planning and effort, be later restored to a prairie, and many was by biologist Eliot Norse in a 1980 government report
have. A prairie whose soil is covered with asphalt to create a (Pimm 2001). Norse was ahead of his time. The word did not
parking lot is probably lost for good. attain common use in science until after the (US) National
Some human habitat changes can increase biodiversity, Forum on Biodiversity in 1986 (Thompson and Starzomski
which can occur in some kinds of agricultural activities, or 2007), with its first use in publication appearing in 1988 in a
when humans set aside areas (nature preserves) for other book based on papers presented at that conference (Wilson
species or intentionally manage ecological processes like 1989a, b). In origin, biodiversity is a contraction of
fire or flooding for the benefit of other species. But, in most “biological diversity” (Wilson and Peter 1989), but today
cases, humans change land for human benefit, not nature’s the word not only has multiple definitions, but multi-
benefit. As a result, human use almost always results in dimensional concepts and applications. To those engaged in
reductions in species richness on the affected site (Fig. 2.7) the study of natural history, biodiversity represents the biotic
(de Baan et al. 2013). elements of nature that can be described and classified. The
2.4 The Problem of Concept: What Is Biodiversity? 45
tool for such description and classification is taxonomy. To defined by patterns of association and structure of vegetation.
environmental activists, biodiversity is an intrinsic, value- In landscapes, the entities might be landforms, landscape
laden quality of natural systems that should be preserved elements, or landscape arrangements, the last two potentially
for its own sake, expressed by using the tools of environmen- including human-dominated or human-created landscapes
tal ethics. To conservation biologists, biodiversity is an such as agricultural or urban landscapes. Ecosystem diversity
aspect of living systems to be descriptively characterized would integrate habitat and landscape elements into function-
and explained, a measurable parameter providing insight ing species processing matter and energy within defined
into community structure, environmental processes, and eco- spatial boundaries, and with characteristic communities of
system functions, determined by the tools of site-specific plants and animals. For example, if we think of forest eco-
survey and study (Burch-Brown and Archer 2017). So before system biodiversity, examples might include tropical
we attempt a summary definition of biodiversity, we must rainforests, temperate deciduous forests, or boreal forests.
recognize the “thought styles” and philosophical Although biodiversity can be applied as a concept and
commitments that are entangled in the word, understand measured as a parameter at multiple levels, it is often
how they will affect the definition (Mayer 2006), and take employed at the level of species in science, management,
stock of our own perspective before we speak of it. If we do and policy. To better appreciate this application and under-
not do these things, we will produce, first, ambiguity, and, standing of biodiversity, we will examine the species concept
then, confusion about what we are trying to conserve in more detail.
and why.
Of the myriad definitions of biodiversity, one of the best
remains the structural and functional variety of life forms at 2.4.2 Biodiversity and the Definition of Species
genetic, population, community, and ecosystem levels
(Sandlund et al. 1992). This definition is especially helpful There are at least 26 different definitions of species used in
to scientific investigation in that it focuses on the two ideas current science (Frankham et al. 2012), but the roots of the
that make biodiversity a workable concept for study – that species concept extend beyond the history of science to the
biodiversity is the entire array of biological variety, not very beginnings of philosophy. The idea of species was
simply a collection of individual species, and that the variety inherent in the thinking of the Greek philosopher Plato who
that defines biodiversity exists at multiple biological levels. perceived the universe as an array of ideal forms. The actual
That is, biodiversity is not simply another name for “all of thing observed, including all material objects, even living
life.” It is a name for the heterogeneity of life, the variation of ones, was only a shadow of its true form (in Greek, its
all living things. The measurement of biodiversity is the eidos). Because each creature was a representation of a true
measurement of that variation. Thus, the conservation of or ideal form, variation among individuals within a popula-
biodiversity is not the conservation of all life, but the conser- tion and among different populations was de-emphasized and
vation of its units of variation, the conservation of the differ- emphasis was placed on the ideal that the creature imperfectly
ent kinds and forms of life (Burch-Brown and Archer 2017). manifested. Such thinking formed the basis of the typological
At the most basic level, biodiversity refers to the genetic species concept that defined species as distinct morphologi-
variety found in Earth’s living organisms. Individual genes cal types.
and their variety of forms (alleles) represent the most founda- The typological concept – subsequently combined with
tional elements at which biodiversity can be measured. In the the Greek philosopher Aristotle’s principles of logical
words of conservation biologist Andrew Dobson, genes are, divisions of organisms based on “common essence”
for biodiversity, “the smallest fundamental units of its calcu- (a unique attribute that makes the species what it is) – was
lus” (Dobson 1996:10). Genetic diversity is the basis for central to biology for many centuries. It was modified again
other units of biodiversity such as populations, subspecies, with the assertion of the English botanist John Ray that
and species. Genetic differences weigh heavily in legal and species “bred true” in producing only offspring of their own
management determinations of what is a species, whether or kind, as well as with the concept of the Swedish botanist
not a species is extinct, whether or not individual animals can Carol von Linné (Carolus Linnaeus) that species were fixed,
survive and adapt if introduced to a new area, which discrete, and natural entities created by God (Stuessy 1990).
individuals should breed with one another in a captive- But with the advent and acceptance of the theory of evolution
breeding program, and whether or not a species has the by natural selection, this typological view of species as dis-
potential to recover its genetic diversity if it increases in crete and rigid categories was replaced with the view of
numbers after a decline. British scientist Charles Darwin that species were mutable
Above the level of species, biodiversity can also be and constantly changing. Darwin stressed the evolutionary
measured as diversity in habitats, landscapes and ecosystems. integrity of species as being all individuals descended from a
In habitats, the entities of diversity are usually habitat types, common ancestor, thus providing the foundations of the
modern evolutionary species concept. However, in Darwin’s environments. . .and/or fixed chromosomal differences”
day, as in this one, it could be difficult to determine with (Frankham et al. 2012:26). The BSC fails, however, to reveal
certainty the common ancestor of every species. Biologist the basis for that isolation, which is that the two species have
Ernst Mayr proposed a more practical criterion. He offered a different numbers of chromosomes, making sperm and eggs
definition of a species as “a group of interbreeding from different species incompatible. Cope’s gray tree frog
populations that are reproductively isolated from other such has 24 chromosomes. The gray tree frog has 48.
groups” (Mayr 1969). Mayr’s definition, now known as the The tree frogs illustrate the need for a different under-
biological species concept (BSC), was useful, and therefore standing of species that goes beyond reproductive isolation.
widely employed, because it provided an operational “test” One attempt to solve the problems of the biological species
for species identity. That test was reproductive isolation. concept is the differential fitness species concept (DFSC;
By using the criterion of reproductive isolation, biologists Hausdorf 2011), which broadens the definition of species to
could identify species of plants and animals that might be be that of groups that have features that would have “negative
otherwise similar, or even identical, in physical appearance. fitness effects in other groups and cannot be regularly
To the naked eye, two tree frogs of eastern North America, exchanged between groups upon contact” (Frankham et al.
Cope’s Gray Tree frog (Hyla chrysoscelis) and Gray Tree 2012:27). In other words, not only would reproductively
frog (Hyla versicolor) are morphologically identical isolated groups be considered species, but any groups
(Fig. 2.8a), and therefore the same species at the morphologi- whose matings lead to negative fitness consequences, such
cal level. But they are reproductively isolated populations. as lower survivorship or reduced reproductive output.
Like other species of frogs and toads, males call in spring to Modern techniques of molecular genetics now permit
attract females. Their calls are different (Fig. 2.8b). Females direct examination and comparison of the DNA,
are imprinted on and attracted to the call of their own species, chromosomes, and gene loci of individual organisms and
so their songs function as a pre-mating isolating mechanism. populations, leading to what is commonly referred to today
These two species of tree frogs reveal in a single example as the phylogenetic species concept (PSC). The PSC asserts
both the strength and weakness of the BSC. The tree frogs are that species are differentiated by measuring genetic
separated as species because they are reproductively isolated. similarities, differences, and distances among populations
Reproductive isolation matters, because, as geneticist or groups of populations. The BSC could only be applied to
Richard Frankham and his colleagues noted, “. . .the devel- organisms that reproduce sexually. The PSC can be applied
opment of reproductive isolation between populations usu- to any organism that has genes – in other words, every form
ally accompanies genetic adaptation to different of life.
Fig. 2.8 (a) The Cope’s gray tree

frog (1) (Hyla chrysoscelis) and
Eastern gray tree frog (2) (Hyla
versicolor), two species identical
in physical appearance but
different in chromosomal
structure. (b) Males in each
species have a different mating
call, as shown in their respective
sonograms. The difference in calls
creates a reproductive isolating
mechanism between the species
and prevents mating events that
would combine sperm and eggs
with incompatible genetic
material. (Photos courtesy of
B. Gloriso, Amphibian Research
and Monitoring Initiative,
US Geological Survey.
Figure reprinted by permission of
Company of Biologists, Ltd.,
from McLister, D., Stevens, E.D.,
Bogart, J.P., 1995. Comparative
contractile dynamics of calling
and locomotor muscles in three
hylid frogs. The Journal of
Experimental Biology 198, 1527–
1538. # 1995 The Company of
Biologists Limited. Adapted by K.
DeVoss)
2.4 The Problem of Concept: What Is Biodiversity? 47
The power of the PSC also carries attendant difficulties. second is a problem of effective communication. If we keep
Conservation geneticist Martha Rojas has framed the the nature of these problems clear, we can focus on what kind
dilemma by asking conservationists whether they are of information we need to solve each problem.
attempting to conserve species as types or as evolutionary
units (Rojas 1995). If we view species as types, we will not be
concerned about maintaining their genetic variability. Alter- 2.4.3 The Species Concept in Conservation
natively, if we view species as evolutionary units, it is not
merely the present group of individuals that concerns us, but The phylogenetic species concept has become powerful and
the evolutionary and speciation potential of the species itself. widely used because methods to specify the genetic criteria
In this view, conservation efforts must not only preserve the on which it depends also have become powerful and widely
organism, but the organism’s ability to respond to environ- used and are now normative tools for defining species. As a
mental change; not only the individuals that make up the result, conservation biology is giving as much attention to
species, but its potential to give rise to future species. As a genetic diversity as species diversity.
discipline, conservation biology has embraced the latter Although the phylogenetic species concept offers impor-
view, and considers long-term maintenance of the evolution- tant gains in insight and precision for the work of conserva-
ary potential of organisms – that is, their genetic diversity – as tion biology, it also raises problems. Paul-Michael Agapow
essential to their survival, even though other definitions of of University College, London (UK) and his colleagues
species continue to be used or assumed in conservation conducted a literature review of 89 published studies where
practice (Table 2.2). a group of organisms was categorized under both phyloge-
Unfortunately, even experienced biologists often focus on netic and non-phylogenetic species concepts. Using
definitional aspects of the word “species” and neglect the non-phylogenetic species concepts, the studies identified
conservation of species-forming conditions. Species, as 1245–1282 species. Under phylogenetic species assessment,
geneticist Jody Hey insightfully perceived, “are the result of the same studies identified 1912–2112 species, an increase of
two processes: (1) the evolutionary processes that have 49%. Across studies, the mean number of species in a group
caused biological diversity; and (2) the human mental appa- increased an average of 121% (it more than doubled!). Fun-
ratus that recognizes and gives names to patterns of recur- gal groups increased 300%, lichens 259%, plants 146%, and
rence” (Hey 2001:328). So, when conservation biologists reptiles 137% (Agapow et al. 2004). These findings indicate
talk about saving “species” as fundamental units of biodiver- that increasing use of the phylogenetic species concept will
sity, both they and their audience must realize that they are affect conservation efforts at multiple levels. Because most
trying to solve two related but different problems. These procedures for identifying endangered species, such as those
problems are: (1) identifying the real evolutionary groups or used by the IUCN’s Red List or the US Fish and Wildlife
units we are working with, and then determining how the Service’s Endangered Species List, consider both numbers of
identification of such groups will affect our conservation individuals in a population as well as geographic range and
strategies for different, and potentially related groups and distribution, a phylogenetic species approach will effectively
(2) devising and using conceptual categories that help us to “split” more populations currently “lumped” under biological
speak to one another about such groups to others (both species and other approaches. As a result, more of these
scientists and the public) and recognize their recurring populations will fall below critical thresholds in numbers
patterns in all forms of biodiversity. The first problem is a and range size. For example, using IUCN criteria, Agapow
problem of scientific description and measurement. The et al. estimated that, regarding “Vulnerable” species, which
Table 2.2 Eight conceptual definitions of species

Concept Name Based on
Morphological Body shape and other structural features, as identified by expert opinion.
Biological Groups of actually or potentially interbreeding populations, which are reproductively isolated from other groups.
Genetic Genetic isolation of interbreeding natural populations.
Ecological A reproductive population occupying a specific ecological niche, adapted to a particular set of resources.
Paleontological Disappearances in the fossil record to understand the fossil’s past diversity.
Evolutionary A single lineage of ancestral descent which maintains its identity from other such lineages, and which has its own
evolutionary tendencies and historical fate.
Phylogenetic High degree of resemblances in many unique traits based on discriminative phenotypes. Share a common ancestor.
(Cladistic)
Biosystematic Collective group of concepts showing more than reproduction or lineage.
Designed by K. DeVoss from concepts outlined in Stuessy, T.F., 2009. Plant taxonomy: the systematic evaluation of comparative data, 2nd ed.
Columbia University Press, New York
have less than 1000 individuals, and “Endangered” species, Conservation biologists now recognize that their view of
with less than 250, a 49% increase in species number species affects the way they approach conservation issues, as
“. . .infers an average decrease in mature individuals per well as the types of problems they attempt to solve and the
species of 32.8%. If we assume the number of mature way in which they solve them. Our mental categories of what
individuals in “Vulnerable” species are distributed evenly a species is are imperfect. But whatever the limits of our
throughout the band of possible values (250–1000), a 33% understanding, we must still accurately measure biodiversity
drop will cause 11% of these species to have less than before we can conserve it. So how is biodiversity to be
250 individuals and so be reclassified as “Endangered”” measured, and what will we learn from our measurements?
(Agapow et al. 2004:169). Newly delineated phylogenetic
species also are likely to have reduced ranges, and, on the
basis of that criterion, the number of endangered species 2.5 How Do We Measure the Earth’s
would increase. Will increasing estimates of the number of Biodiversity?
extant species, endangered or otherwise, create confusion,
and eventually apathy, in the public mind? If these 2.5.1 What Biodiversity Measurements Tell Us
calculations are correct, has taxonomy become the enemy
of conservation? 2.5.1.1 Species Richness and Alpha Diversity
The idea of preserving evolutionary potential of To be meaningful to science, biodiversity must be capable of
organisms as well as the organisms themselves has been measurement as a mathematical variable. Such indices must
formalized in the concept of the “evolutionarily significant be understood for what they are, representations of features
unit,” or ESU. ESUs were originally intended to be based on of biodiversity, not biodiversity itself. But conservation
criteria associated with genetic variation within and between biologists must be able to measure and express such features
populations. In practice, the ESU concept is often replaced in ways that are objective and meaningful to others. They
with the more practical concept of the conservation manage- must clarify what feature of biodiversity is under investiga-
ment unit (MU) as a way of identifying a population or tion and what dimension of that feature is being evaluated. In
groups of populations that show evidence of genetic related- practice, the unit of biodiversity most often used is that of
ness, but also are arranged spatially in such a way that they species, which can be easily understood by the public and
can benefit from a common management strategy. For exam- clearly defined in the legal process.
ple, Claudio Ciofi and his colleagues, using both molecular Alpha diversity is the diversity of species within an eco-
genetic markers and knowledge of geographic distribution of logical community and is normally described as a measure of
populations, were able to identify genetic-based MUs for two attributes – species richness and species evenness, where
conservation of the Komodo dragon (Varanus komodoensis) “richness” is the number of species in the community and
(Fig. 2.9) on the islands that form the bulk of its range in “evenness” the relative abundance of each species in the
south-east Indonesia (Ciofi et al. 1999). community. But what is the “community”? A broad defini-
tion of a community is “all populations occupying a given
area at a particular time.” A single site is usually considered
to have multiple communities distinguished from one another
by common taxonomic levels or ecological traits. For exam-
ple, a contiguous block of tallgrass prairie might be described
in terms of its plant community, its invertebrate community,
its small mammal community, and its bird community, to
name a few. Specialized subdivisions may be used to better
identify functional relationships or specialized traits (for
example, the predator community or the detritivore
community).
Species richness is the number of species present on each
site. To compare different communities, species richness may
be expressed in numbers per unit area. Alternatively, the
species richness of different kinds of communities may be
Fig. 2.9 The Komodo dragon (Varanus komodoensis), a species compared through association measures, such as a coefficient
inhabiting islands of southeast Indonesia. Identification of molecular of community (Goodall 1973), in which two or more
genetic markers and their association with geographically distinct
communities are evaluated on the basis of the percentage of
populations of Komodo dragons have permitted conservationists to
identify genetic-based management unit (MU) populations for conser- species shared by both. If species richness is the sole measure
vation efforts aimed at conserving this species (Photo courtesy of of diversity, then the community with the larger number of
Mehgan Murphy, Smithonian’s National Zoo) species would be considered more diverse.
2.5 How Do We Measure the Earth’s Biodiversity? 49
Measures of species richness are relatively simple to col- incorporate differences in abundance into our estimates of
lect through samples or surveys. Individuals of different diversity?
species need not be counted. The only data the observer Differences in abundance can be incorporated into an
needs to record is whether the species is present or absent. estimate of alpha diversity as measures of species evenness.
The final result is a species list, consisting of the total number Our two sites, A and B, have similar numbers and kinds of
and names of species recorded on a site. A species list is easy species; however, site B not only has more species, but its
to present, interpret, and compare with other communities, species are more equally abundant. Site A not only has fewer
and the number of species present offers a useful approxima- species, it is dominated by just one species (Common Yel-
tion of the biodiversity of the area or habitat. Species lists are lowthroat, Geothlypis trichas). Therefore, site B would be
often the only biodiversity indicator available from older considered more diverse (Table 2.3). This determination has
historical data, or from remote areas where little previous implications. When a community is dominated by one or a
investigation has taken place. few species, it may be that the rarer species are at risk of
Species richness reflects a refinement of the species list, disappearing. The more common species may even be part of
standardized to reflect the number of species recorded per the problem if their behavior is detrimental to rarer species. A
sampling area or per some uniform number of observations. distribution pattern in which one species is far more abundant
Consider an example from a bird community residing in a than all others may indicate that the habitat lacks sufficient
tallgrass prairie in the state of Iowa (USA) (Table 2.3). Based diversity of structure or resources to allow many species to
on an equal number of site visits, Site A is determined to have sustainably exist within it. An examination of species even-
8 species and Site B 11 species. If species richness is the sole ness can be a first step toward generating intelligent
measure of diversity, then the site with the larger number of hypotheses about possible species interactions, leading to
species (B) is considered more diverse. greater understanding of processes that influence diversity
As a measure of diversity, species richness has positive in the community.
attributes. But using species richness alone as an index of One widely used measure of evenness, and of community
diversity has drawbacks. Species richness tells us nothing diversity, is the Shannon Index, which calculates diversity
about relative or absolute abundance of individual species. (H ) as
We do not know whether the species present are equitable in
X
numbers or whether the community is composed of a few H ¼ i pi ln ðpi Þ, ði ¼ 1, 2, 3, . . . SÞ, 0 H0 1
abundant species and many rare ones. Note for instance the
differences in species evenness in our example of the two where pi is the proportion of the total community abundance
grassland sites. The sites share many species, but the same represented by the ith species and ln ( pi) is the natural log of
species have different densities. What is a reasonable way to pi. Diversity indices like the Shannon Index incorporate
quantitative measures of species abundance in relation to
Table 2.3 The abundance of avian species from two sites in tallgrass the total abundance of all species. Thus, the value of the
prairie habitats at the DeSoto National Wildlife Refuge, Iowa, USA and Shannon Index increases as the number of species increases
the calculation of their comparative diversity. Numbers associated with and as they become more equal in abundance.
individual species represent singing males/10 hectares
Species Site A Site B 2.5.1.2 Beta Diversity
Common yellowthroat 8.24 1.21
Whereas alpha diversity measures the diversity of species
Field sparrow 2.94 2.84
within a community, beta diversity measures the diversity of
Dickcissel 1.18 2.23
species among communities. Thus, beta diversity provides a
Red-winged blackbird 0.29 0.81
measure of area diversity or regional diversity. Beta diver-
Brown-headed cowbird 2.06 1.82
American goldfinch 1.47 1.02
sity, sometimes called “beta richness,” measures the rate of
Ringneck pheasant 0.59 1.63 change in species composition in communities across a land-
Mourning dove 1.18 0.61 scape. Ecologist R. H. Whittaker is credited with the origin of
Eastern kingbird – 1.60 the term and used it to indicate the change in species compo-
Grasshopper sparrow – 4.48 sition of communities along an environmental gradient
Northern bobwhite – 2.64 (Whittaker 1975). The “gradient” is often a variable such as
Shannon diversity (H0 ) 1.64 2.25 slope, moisture, or soil pH, which can be measured in the
Designed by F. Van Dyke, based on data from Van Dyke, F., Schmeling, same way and at the same scale in all communities.
J.D., Starkenburg, S., Yoo, S.H., Stewart, P.W., 2007. Responses of Whittaker expressed beta diversity mathematically as
plant and bird communities to prescribed burning in tallgrass prairies.
Biodiversity and Conservation 16, 827–839. # 2006 Springer Science
+Business Media B.V. S=α 1,
where S is the number of species in the entire set of sites and different communities in a landscape in which important
α represents the average number of species per site (alpha environmental variables change over distance. Second, beta
diversity), with sites standardized to a common size. If every diversity provides insight into whether species in different
site has the same number of species, then S/α ¼ 1 and communities are sensitive or insensitive to changing
S/α 1 ¼ 0. In this case, the value of beta diversity would environments, and whether associations of species are inter-
be 0 when sampled sites all have the same species composi- dependent (individual species require the presence of other
tion, indicating a highly homogeneous landscape with respect species to persist) or independent (species are added or lost
to a particular environmental gradient (i.e., every site has in a more or less random fashion). Analyses of both trends
exactly the same species and there is no diversity). Such a are important in understanding biodiversity at the landscape
result would suggest that the species examined had wide level and can be useful in ranking communities in relative
tolerances for that particular environmental variable, and value with respect to conserving biodiversity. Finally, beta
probably broad ecological adaptability and wide niche over- diversity can be used to measure non-environmental
lap. In contrast, suppose the number of species on all sites gradients, and thus measure how species are gained or lost
was 100, but the average number of species at each site was relative to other factors. For example, measures of beta
10. In this case, S/α 1 ¼ (100/10) – 1 ¼ 10 – 1 ¼ 9. This diversity can be used, not only on different, adjacent sites,
result would mean every sampled site was contributing but on the same site at different times, thus determining, as
unique species. the site ages, whether the site increases or decreases in
Today the concepts and measures of beta diversity are species. Such measurement is important in evaluating the
used more broadly to reflect change in species composition effects of disturbance and time on a site’s biodiversity and
between places. Thus, while alpha diversity represents the can provide insights into ecological succession and its effects
“inventory” component of diversity (the diversity of a single on community species composition.
place), beta diversity represents the “differentiation” compo- Studies of beta diversity also can provide insights into the
nent, or how communities change over distance. Whittaker distribution of biodiversity within a geographic area or eco-
originally defined beta diversity as the differentiation of system. For Example, Van Looy et al. (2006), examined the
communities along environmental gradients, but the concept similarity of species pools of floodplain meadow plants
is applied more widely today to measurements of species between sections (“reaches”) of the River Meuse in western
compositional change in communities at any scale, regardless Europe. They determined beta diversity for every pairwise
of the mechanism that causes the change. In this broader combination of reaches (i.e., from two reaches adjacent to
perspective, beta-diversity arises through a complex array one another to two reaches at opposite ends of the river) as
of processes relating to the interaction of species traits with equal to
characteristics of the physical landscape over time
(McKnight et al. 2007). The precise measurement of beta β ¼ Σ unique (unshared) species in each reach /Σ all species
diversity in current studies is best described as a function of in both reaches.
the “distance-decay of similarity” (McKnight et al. 2007:3),
or the decrease in the similarity of composition between If each of two compared reaches had 100 unique species, and
communities with increasing geographic distance between there were no species common to both reaches, then the value
sites. For example, when beta diversity is measured as this of β would be
kind of compositional dissimilarity per unit of distance
between sites for communities of amphibians, birds, and ð100 þ 100Þ=200 ¼ 1:00:
mammals in the Western Hemisphere, it is highest along
the western coasts of both North and South America, as In contrast, if each reach had only ten unique species, but the
well as Mexico and most of Central America, but much two reaches shared 200 species in common, then the value of
lower in the Nearctic (northern Canada) and the interior of β would be
South America (McKnight et al. 2007). Beta diversity is less
well studied in ecology and conservation than its more famil- ð10 þ 10Þ=200 ¼ 0:10:
iar counterpart, alpha diversity, so more surprises are proba-
bly in store as conservationists continue their examination of Thus, the calculation of β provides an index of difference
its patterns of change. (dissimilarity) between reaches. The closer the value of β to
In general, beta diversity provides insight into three 1, the more dissimilar the two reaches are. By computing this
important characteristics of biological communities. First, value for all combinations of reaches, Van Looy et al. were
beta diversity gives a quantitative measure of the diversity able to create a “dissimilarity matrix,” (Table 2.4) and relate
of communities that experience changing environmental the occurrence of dissimilarity to geographical distances
gradients. Such a measure provides a way of comparing between reaches and to associated geographical,
2.6 Rarity and Diversity 51
Table 2.4 Dissimilarity values (S unshared species in each reach/S all area covering 8983 km2 and encompassing scores of different
species in both reaches) for floodplain meadow plants between different habitat types – as a “landscape.” Based on recent bird
reaches (sections) of the River Meuse in western Europe
surveys, it has been estimated that there are 62 species of
I II III IV V VI breeding birds within the Park’s boundaries (Smith et al.
I – – – – – –
2017). This estimate could be called the gamma diversity of
II 0.497 – – – – –
birds for this landscape. As such, gamma diversity can be
III 0.56 0.24 – – – –
thought of as a kind of “mega-alpha diversity” that
IV 0.52 0.42 0.32 – – –
encompasses both alpha diversity (sites) and beta diversity
V 0.52 0.50 0.45 0.31 – –
VI 0.45 0.54 0.52 0.39 0.34 –
(community) at larger scales.
Reprinted by permission from Wiley, from Van Looy, K., Honnay, O.,
Pedroli, B., Muller, S., 2006. Order and disorder in the river continuum:
the contribution of continuity and connectivity to floodplain meadow 2.5.2 Interrelationships of Alpha, Beta,
biodiversity. Journal of Biogeography 33, 1615–1627. # 2006 by and Gamma Diversity
Blackwell Publishing Ltd
The three types of diversity can change independently of one

management, and hydroregime characteristics of the river. another (Fig. 2.10), but in real ecosystems, they are often
They found that increasing levels of beta diversity between correlated. High levels of diversity – whether alpha, beta, or
reaches were not necessarily functions of the geographic gamma – almost always lead to some form of natural rarity
distance between reaches, but the product of differences in (Cody 1986). As species are added to a community, numbers
flow variability and surrounding valley form and soil of individuals in a given species typically decline, a phenom-
characteristics, all of which were related to the maintenance enon called alpha rarity. Beta rarity is typical of habitat
of continuity and connectivity to the river’s historic flood- specialists; abundant in one environment, rare or absent
plain meadows (Van Looy et al. 2006). from others. Gamma rarity describes species with large
populations in local communities and broad environmental
2.5.1.3 Gamma Diversity tolerances, but restricted to particular geographic areas and
Gamma diversity refers to the diversity of species across increasingly rare with increasing distance from their popula-
larger landscape levels. In more measurable terms, it is the tion centers. All these dimensions of biodiversity should be
product of the alpha diversity of a landscape’s communities measured and understood before a biologist can correctly
and the degree of beta differentiation among them. Thus, interpret the biodiversity of a given system and the processes
gamma diversity is used to express the diversity of different that produce it. If we understand the various dimensions of
kinds of communities within a landscape. The ecologist species biodiversity and the ways these dimensions can be
Martin Cody defined gamma diversity as “the rate at which measured, we can begin to understand what makes some
additional species are encountered as geographical species rare and more at risk of extinction.
replacements within a habitat type in different localities”
(Cody 1986). Gamma diversity is, then, “a species turnover
rate with distance between sites of similar habitat, or with 2.6 Rarity and Diversity
expanding geographic areas” (Cody 1986). Unlike beta
diversity, gamma diversity is independent of habitat and 2.6.1 It is Common to be Rare
calculated as
As Charles Darwin remarked over 150 years ago, “Rarity
dS=dD½ðg þ lÞ=2, precedes extinction” (Darwin 1859). That insight remains
true today, and rarity remains an essential concern in the
the rate of change of species composition with distance. In study of conservation biology. Diversity and rarity are
this formula, D is the distance over which species turnover positively correlated. Alpha, beta, and gamma diversity
occurs, and g and l are rates of species gain and loss. Cody’s will increase from community to landscape to region if the
value is difficult to calculate directly, as it requires the use of number of species and their proportional evenness of abun-
calculus to determine the derivatives (dS and dD) from a dance increase, which usually means that densities of most
species-area curve. Today gamma diversity is more often species may become lower as diversity becomes higher.
defined as the total number of species recorded for the To maximize biological diversity, conservationists must
group of sites or communities that make up a landscape, manage rarity.
although the definition of a “landscape” can be both difficult The archetypal rare species is one characterized by small
and arbitrary. But suppose we were to use the example of populations, specialized habitat requirements, and restricted
Yellowstone National Park in the US state of Wyoming – an geographic range. Some species, especially the most
Fig. 2.10 Relationships among alpha, beta, and gamma diversity at a inventory of all the species in one community at one site (alpha diver-
landscape scale. Such relationships can be understood and measured sity), the change in (differentiation of) different communities on differ-
quantitatively on a three-dimensional axis (a) interrelating numbers of ent sites (beta diversity), or the diversity you might encounter among all
species (alpha diversity), changes in environmental conditions (beta the communities you would find in a region (gamma diversity).
diversity) or distance across a larger landscape (gamma diversity). (Designed by K. DeVoss from concepts in Cody 1986; Hillside photo
Conceptually, you can visualize (b) each kind of diversity as an modified from M.A. Ibarra Neri, under CC4.0 BY-SA)
We can visualize what makes the Kirtland’s warbler rare

by examining a “typology of rarity” developed by
Rabinowitz et al. (1986) (Fig. 2.12). The typology is based
on eight categories created by three dichotomies: (1) Is the
population dense or sparse? (2) Does the species use many
different habitats or only one or a few habitats (3) Does the
species have a wide geographic distribution or a narrow
geographic distribution? Rabinowitz and her colleagues sur-
veyed botanical experts regarding the abundance and distri-
bution of 177 species of wildflowers in the United Kingdom
for which data on abundance and distribution were available.
Using the data and surveys, they placed each species in one of
the categories. Of these, only about a third (36%) were
“common” in all respects, having large populations, wide
distributions, and a broad range of habitats. Only 2% were
Fig. 2.11 A male Kirtland’s warbler (Setophaga kirtlandii), a formerly floral equivalents of the Kirtland’s warbler, rare in every
endangered species which combines all components of rarity: small
populations, limited geographic range, and extreme habitat specializa- dimension, having small populations, specialized habitat
tion. (Photo courtesy of Ron Austing) preferences, and restricted geographic ranges. The majority
(62%) were “rare” in one of two aspects of abundance. Each
critically imperiled, meet all of these criteria. For example, element of rarity needs attention to understand its effect on
the formerly endangered Kirtland’s warbler (Setophaga biodiversity.
kirtlandii) has an estimated world population of fewer than
6000 adult individuals (Fig. 2.11). It is an extreme habitat
specialist, breeding only in homogeneous, young, even-aged 2.6.2 Habitat Generalists Versus Habitat
stands of jack pine (Pinus banksiana). It has a highly Specialists
restricted geographic distribution, with more than 60% of
its breeding effort concentrated in five counties in the US Habitat generalists are species that can exploit a variety of
state of Michigan. habitats in a given geographic range. Within that range, they
Fig. 2.12 Eight categories of

species abundance in British
wildflowers based on geographic
range, habitat use, and relative
population size. Note that only
one category (broad habitat
specificity, wide geographic
distribution, and large local
population) can truly be
considered “common.” Species in
all seven other categories are rare
in one or more dimensions.
(Original figure design by M. J.
Bigelow, modified by K. DeVoss,
from concepts in Rabinowitz et al.
1986)
are resistant to extinction through habitat loss or changes in small subset of the vegetation communities endemic to this
land use because, if one habitat is degraded or destroyed, they region, including three species on the IUCN Red List (Grant
move to another habitat or adapt to the altered habitat. The and Samways 2007). For habitat specialists like these
Sulawesi Tonkean macaque (Macaca tonkeana), an dragonflies, loss of preferred habitat is catastrophic and
Indonesia species of monkey that traditionally used mini- leads directly to endangered status.
mally disturbed tropical forests, has shown ability to thrive
in forests converted to agricultural plantations for production
of coffee and cacao. The macaques adapt by changing their 2.6.3 Large Populations Versus Small
food habits, activity patterns, and group size in the plantation Populations
habitat (Riley 2007).
In contrast, habitat specialists are successful and competi- Some species almost always occur in large numbers and high
tive in one or a few types of habitat, but unable to use others. densities. Their abundance may be the result of natural his-
For example, several species of dragonflies (Order Odonata) tory traits such as high reproductive rates, high rates of
found in the Cape Floristic Region of South Africa use only a juvenile and adult survival, or strong competitive abilities
Fig. 2.13 The mountain lion (Puma concolor), an example of a “rare”

species with extensive geographic range and wide habitat tolerance
but uniformly low population density. The mountain lion historically
had the widest distribution of any American mammal other than Homo
sapiens, but local densities were almost always less than 1 individual /
20 km2. (Photo courtesy of US National Park Service)
that allow them to dominate other species. In contrast, other

species, even if widespread geographically, are never abun- Fig. 2.14 The Haleakala silversword (Argyroxiphium macrocephalum),
dant anywhere. Natural history traits that contribute to low a “rare” species with a dense population of individuals (50,000) confined
abundance include low reproductive and survival rates, to a single site, the crater of Haleakala, a Hawaiian volcano. (Photo
courtesy of A. Rullison and US National Park Service)
specialized diets (especially among carnivores), and need
for large areas to find food or complete their life cycle.
The mountain lion (Puma concolor) (Fig. 2.13) histori- Wisconsin, and Michigan from which they were formerly
cally possessed the largest distribution of any American extirpated. If a species has a more restricted distribution, it
mammal, ranging from the southern tip of the Arctic circle is more susceptible to changes associated with local or
in North America to southern Chile and Argentina in South regional land use, human population growth, or climate
America, and from the Atlantic to the Pacific coasts in both change.
hemispheres. As a large, territorial carnivore, one resident An extreme example of a restricted distribution is the
lion requires a minimum of 16–20 km2 and have been known Haleakala silversword (Argyroxiphium sandwicense)
to use areas of up to 600 km2 (Hempker 1982). As a result, no (Fig. 2.14), a striking plant covered with fine, silvery hair
single local area has ever had large numbers of mountain and producing a tall flower stalk at maturity, after which the
lions. When small populations become isolated, prey avail- plant dies. Fifty thousand individuals live on the Hawaiian
ability is reduced, and habitat is restricted. This has been the island of Maui, but all in a single location, the crater of the
case in the subspecies of mountain lion known as the Florida volcano Haleakala.
panther (Puma concolor coryi) in which the population has The silversword, in its extremely confined range,
become imperiled and risk of extinction has increased. illustrates a problem that leads directly to endangerment in
many species. This is the phenomenon of endemism in which
a species is restricted to a particular area or region. Many
2.6.4 Widespread Distribution Versus tropical species are highly endemic, although there are
Restricted Distribution examples of endemic temperate and polar species as well.
In some areas, endemism is characteristic of nearly all spe-
If a species possesses a widespread distribution, like the cies. For example, 90% of Hawaiian plants (including the
mountain lion, the probability of persistence at regional or Haleakala silversword) and 100% of Hawaiian land birds are
global levels can remain high even if individual populations endemic. In the Fynbos region of southern Africa, known for
are threatened, or even exterminated, in particular areas. If its unusual and unique plant communities, 70% of all plant
such species have dispersal abilities sufficient to allow species are endemic. On a continental scale, 74% of
individuals to move among populations, they can periodi- Australian mammals are found only in Australia (Pimm
cally re-colonize sites suffering local extinction, something et al. 1995).
mountain lions have begun to do in the United States, The restricted range of an endemic species makes it espe-
re-establishing populations in states such as South Dakota, cially vulnerable to extinction because local changes in land-
use patterns or climate affect all individuals. If a species is correlations among different taxa were weaker (Table 2.6).
concentrated in a single area, there is no “reserve” of Thus, protecting endemic species of one taxon would not
individuals in another location that might be used to replenish ensure that endemic species in other taxa would be protected
the species if it suffers extinction. Kerr (1997) tested the in the same area, and protecting areas with high species
hypothesized relationship between endemism and biodiver- richness in one taxon would not protect species richness of
sity empirically in North America at four different taxonomic other taxa in the same areas. Kerr examined this correlation in
levels: a class (mammals), a family of butterflies mammals, a group sometimes considered an “umbrella” that
(swallowtails, Papilionidae), a subfamily of moths protects species in other taxa. But Kerr’s data suggests that
(Plusiinae) and a genus of bees (Lasioglossum). He first the “umbrella” provided by mammal diversity is “leaky”
divided the area into quadrats of 2.5 2.5 , then estimated when it comes to conservation of other groups, such as
endemism for each group by counting the number of quadrats invertebrates (Table 2.6).
in which the species occurred, taking its inverse, and sum- Conservationists have always wanted to believe that ende-
ming the total for each quadrat through the formula. mism and biodiversity are correlates, distributions of
endemic species and threatened species are correlates, and
X
S distributions of endemic and threatened species in one group
Endemism ¼ Q1 will correlate with the distribution of endemic and threatened
i¼1
species in other groups. These correlations would make the
conservation of biodiversity so much easier. Unfortunately,
where S is the total number of species in the taxon being
ideas are not true because we wish them to be, even for the
measured and Q is the number of quadrats within the species’
best of reasons. The realities, revealed by more thorough
range. By structuring the formula in this way, species with
analysis of species distribution patterns, suggest that not all
narrow ranges would receive high scores for endemism. Kerr
the correlations conservationists hope for really exist. For
then determined the correlation between a quadrat’s rank in
example, studies by John Lamoreux and his colleagues
endemism for each taxon and its rank in species richness in
have shown that endemism and richness in the four terrestrial
the same taxon.
classes of vertebrates (mammals, birds, reptiles and
Correlations with endemism were high in all cases in these
amphibians) were, when considered separately and by
taxa (Table 2.5), indicating that areas that protect the endemic
ecoregions, highly correlated between and among classes.
species of a taxon are likely to also protect high levels of the
biodiversity in that taxon. However, Kerr found that
Table 2.6 Pearson correlations measuring strength of relationship of

Table 2.5 Pearson correlations measuring strength of relationship
rank in species richness or endemism between different taxa in the
between species rank in richness and endemism within 10 taxa in the
conterminous United States and Canadaa
conterminous United States and Canada
Mammal Lasioglossum Papilionidae
Species richness Correlation with endemisma
Species richness
All Mammalia 0.807
Lasioglossum 0.833 – –
Artiodactyla 0.807
Papilionidae 0.831 0.676 –
Carnivora 0.384
Plusiinae 0.514 0.610 0.376
Chiroptera 0.814
Endemism
Insectivora 0.523
Lasioglossum 0.805 – –
Lagomorpha 0.665
Papilionidae 0.594 0.341 –
Rodentia 0.773
Plusiinae 0.459 0.516 0.238
Lasioglossum 0.851
Papilionidae 0.703 Reprinted by permission from Wiley, from Kerr, J.T., 1997. Species
richness, endemism, and the choice of areas for conservation. Conserva-
Plusiinae 0.772
tion Biology 11, 1094–1100. # 2003 Society for Conservation Biology
Reprinted by permission from Wiley, from Kerr, J.T., 1997. Species a
All correlations shown are significant at P < 0.001. Values of endemism
richness, endemism, and the choice of areas for conservation. Conserva- are log (x + 1) transformed. For more details on Pearson correlations, see
tion Biology 11, 1094–1100. # 2003 Society for Conservation Biology Table 2.5
Values of Pearson correlations can range from 1 to 1. A value of Note that, although correlations are still significant, there is much less
1 indicates that there is a perfect linear positive linear relationship correlation (values are smaller) between levels of endemism or species
between the two variables (they change together by the same amount richness in two unrelated taxa (e.g. endemism in mammals and ende-
and in the same direction). A value of 1 indicates perfect linear mism in Plusiinae) than between the values of endemism and species
negative linear relationship (i.e. the variables are perfectly but inversely richness within the same taxonomic group. Correlation values indicate
correlated; when one increases by x, the other decreases by x). A value of that endemism is strongly associated with species richness, and vice
0 indicates that two variables have no linear correlation versa, when dealing with the same kinds of organisms, but endemism or
a
All correlations are significant at p < 0.001, and endemism is log (x + 1) species richness in one group is not necessarily associated with ende-
transformed mism or species richness in other groups
For example, variation in species richness in mammals well as in all classes combined (Table 2.8). Although corre-
explained nearly two-thirds of the variation in richness in lation between endemism and richness was low at global
the three other vertebrate classes. Similarly, variation in scales, aggregate regions selected for high levels of ende-
endemism in birds explained over 61% of the variation in mism captured more species than expected by chance. A
endemism in other vertebrate classes (Table 2.7) (Lamoreux selection of only 10% of the Earth’s terrestrial surface area
et al. 2006). However, correlations between endemism and could potentially capture 56.5% of terrestrial vertebrate
richness were low, with one variable explaining less than endemic species and 61.6% of all vertebrate species
10% of the variation in the other in any vertebrate class, as (Lamoreux et al. 2006).
Table 2.7 Pearson correlation coefficients of terrestrial vertebrate diversity measures

Amphibians Reptiles Birds Mammals Four Classes
Richnessa 0.591 0.380 0.715 0.668
Endemismb 0.503 0.587 0.612 0.490
Richness x Endemismc 0.096 0.085 0.068 0.099 0.025
Reprinted by permission from Springer Nature, from Lamoreux, J.F., Morrison, J.C., Ricketts, T.H., Olson, D.M., Dinerstein, E., McKnight, M.W.,
Shugart, H.H., 2006. Global tests of biodiversity concordance and the importance of endemism. Nature 440, 212–214. # 2006 Nature Publishing
Group
For richness and endemism, values reflect correlation of that variable in the given class with a counterpart index of richness or endemism in the three
other classes. Values for richness x endemism indicate the correlation between endemism and richness within a class and of the four classes
combined
P < 0.05; P < 0.01
a
Correlation between class richness and a richness index of the three remaining classes
b
Correlation between class endemism and an endemism index of the three remaining classes
c
Correlation between richness and endemism within each class, and of the four classes combined
Table 2.8 Patterns of cross-taxon surrogacy across birds, mammals, and amphibians
Target groups (% of species represented in set)a
Richness index Surrogate group(s) Nspp. Ncells Birds Mammals Amphibians
Total Birds 9626 421 – 79.4 0.3 55.5 0.7
Mammals 4104 509 91.7 0.1 – 61.1 0.5
Amphibians 5619 831 90.9 0.2 86.2 0.2 –
Birds & Mammals 13,730 714 – – 68.4 0.5
Birds & Amphibians 15,245 1028 – 89.8 0.2 –
Mammals & Amphibians 9723 1077 95.0 0.1 – –
All three groups 19,349 1223 – – –
Rarity Birds 2424 380 – 43.3 1.1 22.5 1.3
Mammals 1026 432 68.3 0.4 – 27.0 0.7
Amphibians 1405 560 63.7 0.5 51.6 0.6 –
Birds & Mammals 3450 656 – – 35.7 0.9
Birds & Amphibians 3829 808 – 63.1 0.6 –
All three groups 4855 1033 – – –
Threat Birds 1096 282 – 51.7 0.9 31.2 1.4
Mammals 1033 357 60.7 0.6 – 39.7 0.9
Amphibians 1856 454 62.7 0.4 59.7 0.4 –
Birds & Mammals 2129 518 – – 49.2 0.6
Birds & Amphibians 2952 627 – 67.2 0.5 –
All three groups 3985 821 – – –
Reprinted by permission from Springer Nature, from Grenyer, R., Orme, C.D.L., Jackson, S.F., Thomas, G.H., Davies, R.G., Davies, T.J., Jones, K.
E., Olson, V.A., Ridgely, R.S., Rasmussen, P.C., Ding, T.-S., Bennett, P.M., Blackburn, T.M., Gaston, K.J., Gittleman, J.L., Owens, I.P.F., 2006.
Global distribution and conservation of rare and threatened vertebrates. Nature 444, 93–96. # 2006 Nature Publishing Group
a
Values represent the percentage of species in the target group represented in complementary sets of grid cells designed to contain all members of the
surrogate group (mean +/ standard deviation). Nspp is the total number of species in the surrogate group. Ncells is the number of cells in the optimal
complementarity set
2.7 The Problem of Distribution: Where Is Biodiversity Found? 57
Although the findings of Lamoreux et al. offer some 1.4% of the Earth’s terrestrial surface, contain 44% of the
encouragement to conservation biologists who have long world’s known terrestrial plant species and 35% of known
hoped, and often assumed, that different taxonomic groups terrestrial vertebrates (Malcolm et al. 2006). More than half
have congruent geographical patterns of diversity, the prior- of all threatened plants and terrestrial vertebrates are endemic
ity targets of conservation efforts are not all species but to hotspots (Brooks et al. 2002). But hotspots today are not
endangered and threatened species. If the correlations only centers of global biodiversity, they are also centers of
identified by Lamoreux et al. also applied to threatened and global extinction, a product of their high endemicity in areas
endangered species, this would mean that protecting experiencing accelerating rates of habitat degradation.
threatened species in one group would protect threatened Conservation of the world’s biodiversity will require more
species in other groups. To evaluate this possibility, Richard than identifying, or even effectively protecting, global biodi-
Grenyer and his colleagues conducted a more focused analy- versity hotspots. To understand how to formulate more com-
sis on rare and endangered species of high conservation prehensive, and more effective, conservation strategies, we
priority (Grenyer et al. 2006). They found that, although must examine more closely the global patterns of biodiversity
there was correlation in patterns of species richness among distribution.
birds, mammals, and amphibians, congruence in the distribu-
tion of rare and threatened species in these groups was low,
especially for the rarest species. Using high resolution
Can you envision any ways that precise measurement
databases with grid cells that could be adjusted to varying
of the quantity of biodiversity in an area might help a
sizes, Grenyer et al. found, for example, that, if amphibians
manager make more coherent public statements and
were treated as the conservation target, 55.5% of amphibian
policies about the value of diversity in the same area?
species would be present in a complementary set of grid cells
If so, how, and what form would this take?
selected to contain all members of a surrogate group, such as
birds. But the same relationship dropped to 22.5% for rare
(endemic) species of amphibians and birds, and 31% for
threatened species (Table 2.8) (Grenyer et al. 2006). In the
rarest species (those with the smallest 10% of ranges),
pairwise correlations between groups were negative. This 2.7 The Problem of Distribution: Where Is
means, as Grenyer et al. put it, “. . .the very rarest birds, Biodiversity Found?
mammals, and amphibians live in different places from one
another” (Grenyer et al. 2006:94). Grenyer and his colleagues 2.7.1 Global Patterns of Biodiversity
concluded “. . .even among terrestrial vertebrates, the extent Distribution
to which rare and threatened species from one group can act
as a surrogate for corresponding species in other groups is All vertebrates, as well as plants, show marked increases in
severely limited, especially at the finer scales most relevant to the number of species as one moves from temperate to
conservation.” (Grenyer et al. 2006:95). tropical latitudes (Fig. 2.16) (Reid and Miller 1989; Huston
1994). However, there are exceptions in which diversity
increases in temperate, or even polar areas. Groups that
2.6.5 Conserving Endemic Species show this pattern include sea birds, lichens, marine benthic
organisms, parasitic wasps, and soil nematodes (Huston
Recall Lamoreux et al.’s finding that a careful selection of 1994).
only 10% of the Earth’s terrestrial surface area could contain As with latitude, biological diversity follows an inverse
56.5% of terrestrial vertebrate endemic species and 61.6% of relationship to altitude. This pattern is not surprising given
all vertebrate species (Lamoreux et al. 2006). The that increases in altitude produce environmental and climate
non-random distribution of global biodiversity, specifically effects similar to those of latitude. In plants, for example,
its disproportionate concentration in a small portion of the diversity tends to be highest at low to middle latitudes and
Earth’s surface, is the basis for the “hotspot” approach to lowest at high latitudes and altitudes. Within these broad
ranking areas for conservation, one example of a geographic- patterns, important regional and habitat trends in diversity
based approach to conservation. Today the conservation exist. In marine environments, diversity is highest in coral
community recognizes 25 biodiversity hotspots (Fig. 2.15), reef habitats, coastal zones and estuaries, and tropical marine
each containing at least 1500 endemic plant species (Brooks ecosystems compared to temperate marine ecosystems. In
et al. 2002). In every hotspot, the number of endemic species terrestrial habitats, diversity of tropical rainforests is higher
is high. Even higher is the ratio of endemics to the area in than all other habitats; it is, in fact, legendary. Investigators
which they occur. These hotspot areas, today covering only have found up to 300 different tree species per hectare (2.47
Fig. 2.15 The world’s biodiversity hotspots, centers of global biodi- Conservation International. Eleven hotspots have been added since then,
versity containing only 1.4% of the Earth’s terrestrial surface yet with nine of them shown here in blue. The two newest biodiversity
containing 44% of the world’s known terrestrial plant species, 35% of hotspots (not featured on the map) include Forests of East Australia and
known terrestrial vertebrates, and more than half of the world’s the North American Coastal Plain. (Base map adapted under CC BY-SA
threatened plants and terrestrial vertebrates. The first twenty-five 3.0)
hotspots (green) were identified by Myers et al. 2000 and adopted by
acres) in study plots in Peru (Wilson and Peter 1989). Simi- contrast, an area of the same size in a North American
larly, Ghillean Prance, former Director of Britain’s Royal temperate forest might contain only 15 to 20 tree species
Botanic Gardens and one of the world’s foremost authorities (Van Dyke et al. 1988). As many ant species (43) and genera
on tropical plant diversity, determined that numbers of tree (26) were collected from one tropical rainforest tree in Peru
species ranged from 87 to 473 on sites no larger than 1 ha in as are present in all of the British Isles (Wilson 1989a, b). In
tropical forests in Brazil, Ecuador, Peru, and Bolivia temperate areas, freshwater wetlands contain disproportion-
(Table 2.9) (Prance 1994), with most species coming from ately high levels of species diversity at the landscape level
different genera, and many from different families. In and may be particularly important in systems of low diversity
2.7 The Problem of Distribution: Where Is Biodiversity Found? 59
(for example, boreal swamp forests contribute high biological scale in time frames that can assist current conservation
diversity to otherwise low-diversity boreal forest ecosystems) decisions. Even in established protected areas and reserves,
(Hornberg et al. 1998). Island floras and faunas tend to have information on biodiversity may be spotty and variable by
high rates of endemism, and so make large contributions to site. A recent examination of the best sources of information
world biodiversity. In streams and rivers, recent research has on biodiversity in Biosphere Reserves in Spain, for example,
demonstrated that headwaters (streams near the origin of found that information from any one source was invariably
river systems) contribute disproportionately to the biodiver- partial and incomplete. Only about three-quarters (76%) of
sity of the river systems they generate, especially in their vertebrates in the reserves were named in all three of the most
contributions to beta (landscape scale) diversity, with indi- authoritative sources, and multiple sources had to be
vidual localities in headwater areas each accounting for large consulted to create a more complete inventory of actual
proportions of regional-scale biodiversity associated with the biodiversity in the reserve (Pino-del-Carpio et al. 2014).
stream (Finn et al. 2011). The Spanish example shows the difference between accuracy
and completeness when assessing biodiversity on a site- and
area-specific basis, and the need to consult multiple sources.
2.7.2 Identifying Key Biodiversity Areas – If existing sources are not entirely reliable, there is always
Conservation with Incomplete Data the alternative of getting information yourself – through
direct on-site surveys – but such an approach is time- and
There is not enough money, labor, and expertise to identify, effort-intensive, and often expensive. Thus, conservation
count, and map the distribution of every species at a global biologists have been engaged for some time in attempting
to find non-census indicator methods that can rapidly identify
areas with disproportionately high levels of biodiversity.
Initial efforts to accomplish this goal relied on the previously
described umbrella species concept for biodiversity preser-
vation, a strategy which assumes that areas that are species-
rich for one taxon will be species-rich for others. Unfortu-
nately, empirical data provide little support for this assump-
tion. As noted earlier, most studies have found that
taxonomic distributions of organisms are independent. No
one group predicts distributions of others, and rare species
often do not occur in areas that are otherwise species rich
(Kerr 1997; Grenyer et al. 2006; Lamoreaux et al. 2006).
Other recent studies have attempted to gain a more com-
prehensive picture of regional biodiversity by combining
knowledge of ranges of traditional “species” with informa-
tion on phylogenetic lineages based on genetic analysis. For
example, Leslie Rissler and her colleagues compared spatial
patterns of endemism and conservation value of 22 species of
reptiles and amphibians in California (USA) with the 75 phy-
Fig. 2.16 Latitudinal patterns in species richness from tropical to
temperate regions. In most taxa the number of species increases from logenetic lineages they contained, as determined from previ-
temperate to tropical regions. (Figure based on data from Reid and ously published genetic studies (Rissler et al. 2006). In this
Miller 1989) case, phylogenetic analysis revealed several areas of high
Table 2.9 Occurrence of tree taxa at different levels found on 1 ha inventory areas in neotropical forests
locality area/ha min dbh indiv. species gen. fam. reference
Belém, Brazil 1 10 423 87 65 31 Black and Pavan (1950)
Manaus, Brazil 1 5 1561 473 187 54 Valencia et al. (1994)
Cuyabeno, Ecuador 1 10 693 307 138 46 Valencia et al. (1994)
Añangu, Ecuador 1 10 728 228 126 53 Balslev et al. (1987)
Mishana, Peru 1 10 842 275 – 50 Gentry (1988)
Yanamono, Peru 1 10 580 300 – 58 Gentry (1988)
Alto Ivon, Bolivia 1 10 649 94 61 28 Boom (1986)
Reprinted by permission of the Royal Society, from Prance, G.T., 1994. A Comparison of the Efficacy of Higher Taxa and Species Numbers in the
Assessment of Biodiversity in the Neotropics. Philosophical Transactions: Biological Sciences 345, 89–99. # 1994 The Royal Society
Note, min dbh ¼ minimum tree diameter at breast height in meters; indiv. ¼ total number of individuals
Fig. 2.17 Differences between

species distributions (shaded
areas) and distributions based on
genetic lineage (unique symbols)
of one species of amphibian, the
California newt (Taricha torosa),
and one species of reptile, the
California kingsnake
(Lampropeltis zonata). In each
species, phylogenetic analyses
reveal distributions of unique
evolutionary lineages that would
have been overlooked in an
analysis of species’ distribution
alone. (Reprinted by permission
from University of Chicago Press
Journals, from Rissler, L.J.,
Hijmans, R.J., Graham, C.H.,
Moritz, C., Wake, D.B., 2006.
Phylogeographic lineages and
species comparisons in
conservation analyses: a case
study of California herpetofauna.
The American Naturalist 167,
655–666. # 2006 The University
of Chicago)
conservation value and unsuspected areas of phylogenetic anything, and no species in the Red List’s Category One
endemism that would not have been identified in a species (Extinct) can now be saved. For species that remain, protec-
level analysis (Fig. 2.17). Such analysis also aided in detection must begin by identifying where threatened species live,
tion of sites that could be described as “phylogenetically determining what measures must be taken to ensure their
irreplaceable.” Traditionally, managers have designated irre- survival, and then finding people with the will, influence,
placeable sites as those protected to ensure persistence of resources, and authorization to carry out the plan.
species. Using a phylogenetic approach, irreplaceable sites Conservation efforts historically began with legislation
also would include sites protected to ensure persistence of intended to prevent overharvest of game species and curtail
unique genetic lineages. the trade in wildlife products that promoted such harvests. US
But, as noted earlier, phylogenetic approaches, while dis- conservation laws like the Lacey Act did just that. Originally
covering otherwise hidden lineages of high conservation passed in 1900, the Lacey Act became the first US federal law
value, also increase numbers of endemic, vulnerable, and protecting wildlife by creating and enforcing civil and crimi-
endangered species needing protection, straining already lim- nal penalties for illegal trade of animals and plants. Under the
ited resources for conservation. And there may be insufficient Lacey Act, it is unlawful to import, export, sell, acquire, or
information available to conduct the multi-level, complemen- purchase fish, wildlife or plants that are taken, possessed,
tary species and lineage assessments of biodiversity needed to transported, or sold in violation of US or Indian (Native
discover all significant phylogenetic lineages that are present. American) law, or in interstate or foreign commerce involv-
ing any fish, wildlife, or plants taken possessed or sold in
violation of State or foreign law. Amended by the US Con-
2.8 Preserving and Managing Biodiversity gress in 2008, the Lacey Act now includes a wider variety of
prohibited plants and plant products, including products
2.8.1 Past Approaches to Conservation made from illegally logged woods from any part of the
Management: Conservation Legislation world, for import. Properly applied, the Lacey Act can be a
powerful tool to protect biodiversity by cutting off the profit
Global conservation and scientific information resources, motive from illegal trade in plant and animal products. Using
although admittedly imperfect, can help locate and define a similar approach, the Convention on International Trade in
which species are in need of special protection. Foremost Endangered Species of Wild Fauna and Flora (CITES)
among these is the previously described IUCN “Red List” regulates international trade in plant and wildlife products
(www.iucnredlist.org) which places listed species in different and aims to prohibit trade in endangered species altogether.
categories of endangerment. However, no list ever protected However, a nation requires strength of intention as well as a
2.8 Preserving and Managing Biodiversity 61
high level of enforcement capacity to make conservation laws Such examples demonstrate that it sometimes takes more
matter. Many nations do not possess these attributes. As a than laws and boundaries to conserve species. Despite the
result, violations of CITES are plentiful and predictable violence that sometimes accompanied conservation efforts in
worldwide. these contexts, national parks and other protected areas were
vigorously developed in the US and Canada from the late
nineteenth century on and subsequently imitated throughout
2.8.2 Protected Areas the world. In different contexts, however, they had mixed,
and often less, success. In addition to the injustices that the
Concurrent with development of conservation laws to protect establishment of national parks and refuges commonly
species, conservation efforts also have historically foisted upon human residents of areas designated for protec-
concentrated on setting aside protected areas, a strategy tion, planned locations of parks and refuges were often deter-
intended to preserve habitats and ecosystems where charis- mined by political consideration rather than scientific
matic endangered species lived, or where the habitat or eco- assessment – or human compassion – with predictable
system itself was considered sufficiently unique to warrant shortcomings in their conservation value. Preserves often
special protection. Past motivations and strategies in this failed to contain functionally complete ecosystems or ranges
effort, like most endeavors in conservation, were not always big enough for populations of larger mammals. Many were
well conceived or effectively executed. Protected areas were placed in areas of low biodiversity containing few unique or
sometimes established to save species from market hunting, endemic species. In countries with weak governments, the
as was, in part, the case in the first US national park, “protection” afforded was minimal, hindered by lax enforce-
Yellowstone, established in 1872, where the last remaining ment which made the areas vulnerable to poaching motivated
population of wild bison (Bison bison) remained. With bison by market-driven exploitation of the preserve’s plants and
and other park species being devastated by poachers, the US animals. Their fixed boundaries also made them susceptible
Army was called in to protect the animals, and in a short time to deterioration associated with climate change, as well as to
had removed poachers from Yellowstone, sometimes with detrimental effects of activities of human communities bor-
deadly force. The first US “wildlife refuge,” Pelican Island, dering protected areas. In the face of these problems,
was established by President Theodore Roosevelt in 1903 to conservationists began to consider other strategies, not to
protect water birds, especially those in the US state of replace protected areas, but to complement them.
Florida, from market hunters who were making large profits In the current effort of global biodiversity conservation,
selling feathers of these species to meet demand arising from the IUCN World Commission on Protected Areas (WCPA)
then-current fashions in ladies’ hats. This reserve also arose provides the conservation community with the world’s pre-
in response to lawlessness and violence perpetrated by mar- mier network of protected area expertise, assisting
ket hunters, not only against birds, but people. In 1901, the governments and conservation organizations to plan for and
American Ornithologist’s Union and Florida Audubon Soci- create protected conservation areas. The WCPA has
ety (FAS) led a successful campaign to pass legislation in contributed to the systematic identification and methods of
Florida for the protection of non-game birds. Four wardens effective protection of areas of high biodiversity through its
were hired by the FAS to protect birds from market hunters, development of criteria and methodology for identifying Key
but the hunters were motivated, determined, and ruthless, Biodiversity Areas (KBAs) as sites that could contribute
murdering two wardens in the line of duty. Realizing that disproportionately to global persistence of biodiversity
the birds (and the wardens) needed greater protection, Florida (IUCN 2016a). The WCPA’s development of an international
conservationists persuaded Roosevelt to establish the Pelican standard for identifying and protecting identified areas of
Island preserve. On March 14, 1903 by Executive Order, it high biodiversity, consolidated and published in 2016 as A
became the first US federal bird reservation. Many African Global Standard for the Identification of Protected Areas
governments have responded more recently, and more vio- (IUCN 2016a), provides explicit definitions for KBA criteria
lently, to the same kinds of problems. In the 1980s and 1990s, and threshold conditions for sites to be considered for classi-
military force was directed against similarly ruthless fication as a protected area.
poachers to protect black rhinoceros (Diceros bicornis),
elephants (Loxodonta africana and L. cyclotis) and game
wardens in national parks and other protected areas through 2.8.3 Biodiversity Conservation, Landscape
programs such as Operation Stronghold (Zimbabwe) and Conservation, and Human Development
Operation Uhai (Tanzania). Likewise, authorization was
given to wardens, police, and military personnel to “shoot Although the strategy of conservation through protected
on sight” any poachers in parks and protected areas in the areas has a long history and some success, it has often
Central African Republic, Kenya, and Malawi (Sarkar 2012). dichotomized natural landscapes and human culture instead
hotspot areas for tropical forest plants that were vulnerable to

species loss because of high rates of endemism coupled with
high rates of deforestation (Table 2.10) (Myers 1988). In each
of these areas, the number of endemic species was high, and
the ratio of endemics to the area of the forest in which they
occurred even higher. Unfortunately, the boundaries of these
and subsequently designated biodiversity hotspots were too
coarse for setting practical and specific conservation goals.
Additionally, these and other hotspots have lost most of their
original habitat. Managers now must concentrate on what
remains, which often makes conservation efforts highly
site-specific (Harris et al. 2005) and re-introduces many of
the problems of traditional protected areas that hotspots were
intended to solve.
Fig. 2.18 The Sidamo lark (Heteromirafra sidamoensis), an In a radical departure from the “hotspot” approach to
endangered species that depends upon modified human landscapes in
Ethiopia. (Photograph courtesy of Paul Donald, Birdlife) biodiversity conservation, conservationists Peter Karieva
and Michelle Marvier are among those now advocating for
giving priority to so-called “biodiversity coldspots,” placing
of integrating them. Over-reliance on protected areas for increased emphasis on protection of areas providing high
conservation can end up separating humans from nature, benefit to human welfare. They point out that, worldwide,
causing declines in species dependent on human modified economically poor human communities are heavily depen-
landscapes, like the blue crane (Anthropoides paradiseus) in dent on direct use and harvesting of biodiversity resources.
South Africa, the Sidamo lark (Heteromirafra sidamoensis) But, in these cases, the biodiversity resources come from
(Fig. 2.18) in Ethiopia, and remaining populations of giraffes common and abundant species in landscapes where overall
in West Africa (Giraffa camelopardis peralta) (Trimble and biodiversity may be low (coldspots) (Kareiva and Marvier
van Aarde 2014). In northern India, a human-constructed 2007). Nevertheless, such areas, although possessing lower
wetland created by the maharajahs of Bharatpur in the nine- species diversity, provide more ecosystem services needed by
teenth century was used as a private hunting preserve by people, and important areas for many animal species as
these rulers, but also served as a grazing area for local village migratory stops, seasonal homes, or nesting sites (Kareiva
livestock (primarily buffalo, Bubalus bubalis) and a source of and Marvier 2003, 2007). Such considerations of both human
irrigation for village agriculture. Used by hundreds of species need and biodiversity conservation must be combined into
of resident and migratory birds, it was the last location in global conservation strategies if the fabric of biodiversity is to
south Asia where the migratory Siberian crane (Grus be sustained worldwide.
leucogeranus) was recorded. Set aside as a protected area An even more systemic version of Kareiva and Marvier’s
for conservation after India’s independence in 1947, it was “coldspot” strategy is landscape-level conservation, which
designated as Keoladeo Ghana National Park in 1981. has emerged as an alternative to biodiversity conservation
Grazing was banned shortly thereafter. Removal of buffalo strategies relying solely on protected areas. Although
and other grazers permitted increase of Paspalum grass protected areas can be effective as sites which preserve bio-
(Paspalum spp.) and other grasses formerly controlled by diversity as a primary objective, individual protected areas
domestic livestock. The grasses choked shallow waterways are often too small to adequately capture necessary
and reduced bird populations, nesting activity, and fish abun- components of ecological processes or meet needs of viable
dance (Sarkar 2012). populations of animals with large home ranges. In contrast,
Chastened by these and other negative experiences, and landscape-level conservation efforts focus on landscape
better informed by global patterns of biodiversity distribu- integrity and connectivity, including privately owned lands,
tion, many conservationists have begun to advocate alterna- urban areas, and agricultural areas, taking into account
tive approaches. The previously discussed “biodiversity human dependence on resources (Sharma et al. 2010). Some-
hotspots” strategy arose, in part, out of such concerns, not times described as Integrated Development Projects (IDPs) or
only as a first approximation of the location of priority areas Community-based Natural Resource Management
for biodiversity conservation, but as an intentional strategy to (CNRMs), such approaches consider economic as well as
persuade policy makers to put legislation and resources for scientific assessments of biodiversity value, especially
conservation protection where they would have the greatest human dependence on resources needed for subsistence by
impact on small populations of endemic species. In the local residents. IDPs and CNRMs are characterized by
1980s, conservationist Norman Myers identified four such investment of power and decision-making in local,
Table 2.10 Areas of disproportionately high plant biodiversity (“hotspots”) in tropical forests
Extent of forests (km2) Number of
endemics in
Plant species original Original endemics as Present forest area as
Present in original forests proportion of Earth’s plants proportion of Earth’s land
Area Original (primary)a forests (percentage) total (percent) surface (percent)
Madagascar 62,000 10,000 6000 4900 (82) 1.96 0.00675
Atlantic coast 1,000,000 20,000 10,000 5000 (50) 2.00 0.0135
Brazil
Western 27,000 2500 10,000 2500 (25) 1.00 0.0017
Ecuador
Columbian 100,000 72,000 10,000 2500 (25) 1.00 0.0486
Chocó
Uplands of 100,000 35,000 20,000 5000 (25) 2.00 0.0236
Western
Amazonia
Eastern 340,000 53,000 9000 3500 (39) 1.40 0.0358
Himalayas
Peninsular 120,000 26,000 8500 2400 (28) 0.96 0.0175
Malaysia
Northern 190,000 64,000 9000 3500 (39) 1.40 0.0400
Borneo
Philippines 250,000 8000 8500 3700 (44) 1.48 0.0054
New 15,000 1500 1580 1400 (89) 0.56 0.001
Caledonia
Totals 2,204,000 292,000 Xb 34,400 13.8 0.2
For Comparison:
Hawaii 14,000 6000 825 745 (88) 0.30 0.004
Queensland 13,000 6300 1165 435 (37) 0.17 0.004
Reprinted by permission from Springer Nature, from Myers, N., 1988. Threatened biotas: “Hot spots” in tropical forests. Environmentalist 8, 187–
208. # 1988 Springer Publishing Company
a
Some, though not many, primary forests species can survive in degraded forests
b
It is unrealistic to sum total these figures for plant species, on the grounds that there is some overlap between adjacent regions, e.g., some plants
occur in Peninsular Malaysia, Northern Borneo and the Philippines
Note: There is a great range of accuracy in these figures. At one end of the range, some figures are thoroughly well documented and can be generally
considered accurate to within 5 percent or better. At the other extreme, they are little more than “informed assessments” or even “educated
guestimates.” The rest fall in between, with more of them clustered towards the “accurate” end of the range than the “imprecise” end
community-based institutions, integration of social and eco- were detrimental to a local economy dependent on livestock-
logical values, and participatory interaction with mutual based farming systems and polycrop agriculture practiced on
learning among all decision-making parties (Trimble and relatively small (1–2 ha) family-operated holdings (Nautiyal
van Aarde 2014). and Nidamanuri 2012), all practices that promoted native
Results of landscape-level conservation and integrated biodiversity. The Park’s new, more restrictive policies not
development efforts have been mixed, but some have only degraded local biodiversity by changing traditional
demonstrated tangible advantages over top-down conserva- farming practices but created personal and political hostility
tion strategies imposed by national governments. In India’s to initiatives designed to preserve biodiversity in the
Rajiv Gandhi National Park, for example, in the mountains of protected area. These conditions not only had an adverse
the Western Ghats, one of the world’s recognized biodiver- effect on the local economy and household incomes of local
sity hotspots, 94% of residents living within this recently families but contributed to a breakdown of traditionally posi-
established park have been opposed to park conservation tive relationships between nature and local human
policies (Nautiyal and Nidamanuri 2012). This was communities, provoking increased hostility toward
understandable, given that park policies and practices government-initiated conservation efforts. Similar outcomes
included a ban on agriculture in and around the park, resulting from the establishment of national parks have
restrictions on raising and grazing livestock, a ban on become increasingly common in recent years throughout
collecting non-timber products from forest, and exclusion of the Majority World.
local and indigenous communities from decision-making in The advantages of community-based, landscape-level
conservation programs and development of tourism activities conservation strategies have been exemplified in two notable
(Fig. 2.19) (Nautiyal and Nidamanuri 2012). Park policies successes represented in the Kailish Sacred Landscape and
Fig. 2.19 Percentage of people expressing negative attitudes toward dependent upon and accustomed to subsistence agriculture were at the
park policies and effects (y axis) and reason given for negative attitudes heart of many objections, as was a lack of opportunity for local citizens
(x axis) inside and outside Rajiv Gandhi National Park (RGNP), India. to participate in decision-making on park policies or benefit from park-
Government restrictions on rearing livestock and grazing, selective bans related tourism. (Reprinted by permission from Springer Nature, from
on agriculture and harvesting non-timber forest products (NTFP), dam- Nautiyal, S., Nidamanuri, R.R., 2012. Ecological and socioeconomic
age to crops from park wildlife, and failure to involve local citizens in impacts of conservation policies in biodiversity hotspots: a case study
activities and decision-making associated with conservation and tourism from Rajiv Gandhi National Park, India. Journal of Environmental
created negative and even adversarial relationships between local Studies and Sciences 2, 165–177. # 2012 AESS
citizens and government. Imposed restrictions on a human culture
the Bramhaputra-Salween Landscape, large areas including preservation. That view is changing, partly in response to
portions of India, Nepal, and China. These efforts shifted the the inexorable global trend in urbanization, but also in the
ratio of land-use rights from the government to individuals, light of recent studies demonstrating that urban areas can
increasing private and community-based ownership of lands, contribute substantially to regional biodiversity in some
such as forests, with high levels of biodiversity, and circumstances. For example, recent research has
strengthening engagement in the management of such forests demonstrated that urban centers can have higher populations
by local communities (Sharma et al. 2010). At a global level, of wild animals than rural areas of equivalent size and can be
the United Nations program of Biosphere Reserves, which centers of high species richness and local and regional biodi-
incorporates greater roles of subsistence use and continued versity (Marzluff 2001; Marzluff and Rodewald 2008; Nilon
residence of human communities within and adjacent to areas 2011). Although urban residents usually have less interaction
designated for biodiversity protection, has followed a similar with and dependence upon local biodiversity for their
approach toward landscape-level conservation and livelihoods than residents of rural communities, they can
sociologically-informed management policies. Such efforts develop positive attitudes toward wildlife and biodiversity
continue to develop and strengthen emphases on participa- through education and provision of positive contacts with
tory approaches, decentralized management of resources, and nature, such as in urban parks, if such contacts can occur
greater investment of decision-making power in local close to where they live (Nilon 2011). Further, managers of
communities. urban biodiversity can effectively engage urban populations
in conservation by making specific conservation practices
and activities understandable and accessible to urban
2.8.4 Urban Biodiversity Conservation populations, as was accomplished in a program for bird
conservation in the US city of Columbia, Missouri
In traditional views of conservation, urban centers were never (Table 2.11) (Marzluff 2001; Marzluff and Rodewald 2008;
seriously considered as contributors to biodiversity Nilon 2011).
Table 2.11 Practices fostering urban bird conservation with potential and actual activities supporting grassland and shrubland bird conservation in
a mid-sized US city, Columbia, Missouri
Recommended practice Potential activities Actual activities
Protect natural areas and matrix Identify habitats Limited/no
Plan explicitly for open space in new subdivisions Planning for grassland/shrubland habitats No
Management of built and natural spaces in developed areas Zoning and guidelines/ordinances Zoning in place/no guidelines
Enhance existing habitat in Research-based Yes
Natural areas/open spaces Management guidelines –
Improve management of matrix habitats Guidelines for property owners/managers Limited and not specific
Celebrate urban biodiversity Information to and engagement of residents Limited by income and education
List of practices developed by Marzluff and Rodewald (2008). Potential and actual activities reprinted by permission from Springer Nature, from
Nilon, C.H., 2011. Urban biodiversity and the importance of management and conservation. Landscape and Ecological Engineering 7, 45–52.
# 2011 International Consortium of Landscape and Ecological Engineering and Springer
Fig. 2.20 Diversity and species richness in urban, agricultural and humans (bird feeders). Native species persist in remnant habitat patches
wildland settings. (a) Processes of community formation in urban but are vulnerable to longer-term decline. (3) At high levels of urbani-
birds. (1) Food resource abundance increases with urbanization, zation, coexistence mechanisms between native and invader species
supporting higher overall bird densities. (2) In wildland areas, a greater collapse (region c) as predation pressure declines and food resources
variety of more specialized food resources are captured by native, non- become more abundant and homogeneous, reducing or eliminating
human-dependent species. In urban areas, introduction of human- niche separation based on foraging differences, permitting invader spe-
provided foods supports greater numbers of invader species than native cies to become the dominant foragers. Native species become locally
species. (3) Predation mediates community composition in wildland extirpated. Dotted line indicates changes in level of overall bird diversity
areas, but has less effect in urban areas where native predators are less along a wildland to urban gradient. (Reprinted by permission from
abundant. (b) Patterns of change in urban bird communities. (1) In Oxford University Press, from Shochat, E., Lerman, S.B., Anderies, J.
wildlands, native species (blue line) outcompete human-dependent M., Warren, P.S., Faeth, S.H., Nilon, C.H., 2010. Invasion, Competi-
invader species (red line, region a). (2) At intermediate levels of urbani- tion, and Biodiversity Loss in Urban Ecosystems. Bioscience 60, 199–
zation, invader species begin to out-compete native species (region b) by 208. # 2010 American Institute of Biological Sciences)
exploiting more stable and homogenized food resources supplied by
Urban settings also can create conditions that reduce bio- urban biodiversity conservation in the following chapter
diversity compared to rural areas or wildlands. For example, (Chap. 3).
wildlife food supplementation, such as bird feeding stations,
are important mechanisms for increases in urban bird
densities, as is lower predation pressure (Marzluff and 2.8.5 Biodiversity Technology: Finding Areas
Rodewald 2008). Under these conditions, bird diversity of Conservation Value Using Remotely
declines because communities become dominated by a Sensed Data
small number of invasive bird species more efficient at
using available resources, leaving less available food for The development of remote sensing (RS) technology and the
less efficient foragers (Fig. 2.20) (Shochat et al. 2010). We data it collects, much of which is now available at no cost to
will examine the problems and opportunities associated with the user, offers expanded opportunity for determining and
monitoring conservation value of areas worldwide. As RS refine conservation priorities to one habitat type and selected
expert Gary Geller and his colleagues have noted, “Every eight sites predicted to be rich in threatened species. This
remotely sensed image of Earth can be considered a approach was cost-effective because habitat losses within
biological dataset” (Geller et al. 2017). As problems of hotspots require discriminating choices of individual sites
accessibility and cost are being overcome, the potential for as a prerequisite to effective conservation. Using this method,
using RS in monitoring changes in biodiversity and guiding Harris et al. were able to identify areas small enough for cost-
conservation efforts is increasing. effective preservation, yet rich enough in biodiversity to
RS works fundamentally by measuring the energy make a real difference in saving species. In retrospect, the
reflected and emitted from the surface of the Earth, authors concluded, “Many conservationists wish to preserve
differentiating different kinds of surfaces based on the spec- the entire land area of biological hotspots, which are rich in
trum of wavelengths reflected or emitted. When calibrated species but low in habitat, such as the Atlantic Forest Region
using on-site measurements, RS data can identify not only of Brazil. Although conservation wants the lot, funding,
general types of surface conditions, but also vegetation types politics, and the amount and spatial extent of remaining
and even individual plant and animal species (Fig. 2.21). An forests complicate this goal.. . . Refining hotspot conservation
increasingly common use of RS in biodiversity conservation means identifying specific locations (individual habitat
is to use on-site measurements of landscape, vegetation, or patches) of realistic size and scale for managers to protect
other environmental conditions where a species has been and politicians to support” (Harris et al. 2005:1967).
seen or detected, then develop a model that permits some or This example of using remotely-sensed data in conserving
all of those conditions to be identified by RS, a technique biodiversity is embedded in an increasingly systemic
known as species distribution modeling (SDM). For example, approach to the worldwide monitoring of biodiversity. The
SDM has been used to describe or predict how climate Group on Earth Observations (GEO) has designed and now
change has or could affect species distributions by using operates the Global Earth Observing System of Systems
current or forecast climate data in the area where a species (GEOSS). GEO itself is a voluntary partnership of
occurs (Geller et al. 2017). Although there are a variety of governments and international organizations which provides
specific applications than can be used to aid biodiversity a framework for international cooperation on many fronts,
conservation, RS is finding its greatest current applications including but not limited to biodiversity conservation
in initial biodiversity assessment and long-term monitoring. (Muchoney and Williams 2010). GEO is now involved in
For initial assessment, RS can be used at global scales, survey monitoring biodiversity loss, with particular attention to the
large land areas, and provide initial assessment of remote stated requirements of the Convention on Biological Diver-
locations. For monitoring, RS can make repeat images of sity (CBD). The biodiversity element of GEO, the Group on
the same site essential for evaluating site-specific change Earth Observations Biodiversity Observation Network (GEO
over time. BON) provides a global, scientifically robust framework for
RS is particularly valuable, indeed essential, in places observations designed to detect changes in biodiversity by
where rapid rates of habitat destruction sometimes make coordinating gathering and delivery of data related to such
historical sources of obsolete. For example, in Brazil’s Atlan- change. GEO BON stores, organizes, and provides access to
tic Forest Region, an ecosystem which has suffered extensive datasets needed for biodiversity conservation, including pro-
reduction in area and degradation in habitat quality, vision of global land survey data (in partnership with the
biologists Grant Harris, Clinton Jenkins, and Stuart Pimm United States Geological Survey (USGS) and the US
noted that “[older] maps of historical species richness National Aeronautics and Space Administration (NASA).
identified priority areas that are no longer forested, with little The global land cover maps produced can characterize extent
or no biodiversity left to conserve” (Harris et al. 2005). To and condition of natural and managed vegetation, phenology
solve this problem, they combined on-site investigation with (temporal and seasonal changes associated with biological
RS technology to develop a three-step method for identifying communities), and roads and human settlements, as well as
areas of conservation priority. First, they mapped an area of monitor data from specific sites used in longer-term ecologi-
interest (in this case, a remnant of Brazil’s Atlantic Forest) cal research.
using satellite imagery designed for regional analysis Although the technology is impressive, the most important
(AVHRR, SPOT VGT, and MODIS). Then, using contribution of GEO BON may be its creation of a platform
predictions of areas and locations of remaining intact forests for the human community to think globally and, perhaps, act
generated from the mapper and supplemented with field- cooperatively with regards to biodiversity loss and conserva-
collected data on bird species’ ranges and elevations, they tion, informed by accurate and relevant information that is
identified a subregion with the highest density of threatened accessible and transparent to governments and organizations
birds. Lastly, they used the results of their first two steps to with vested interests in biodiversity conservation.
Fig. 2.21 Three scale-dependent views of the Peruvian Andes–Ama- Red colors indicate higher N + P and lower LMA relative to yellow and
zon region derived from airborne imaging spectroscopy. (a) Peru-wide blue colors. (c) Individual species detections within the zoom box of
map shows the distribution of functionally-distinct forests. Different panel (b), derived using species-specific canopy spectra. (Reprinted by
colors indicate varying combinations of remotely sensed canopy data permission from Elsevier, from Asner, G.P., Martin, R.E., 2016.
indicators such as foliar nitrogen (N), phosphorus (P), and leaf mass per Spectranomics: Emerging science and conservation opportunities at
area (LMA). (b) Zoom image from the Peru-wide map indicates major the interface of biodiversity and remote sensing. Global Ecology and
changes in canopy N, P and LMA within a lowland Amazonian forest. Conservation 8, 212–219. # 2016 The Authors)
Fig. 2.22 Structure of the

International Union for
Conservation of Nature (IUCN)
Categories of Endangerment
(Reprinted by permission from
the IUCN)
2.8.6 Should Management of Biodiversity The aforementioned IUCN Red List of Ecosystem
be Species-Based or Ecosystem-Based? Categories provides an objective framework for identifying
and categorizing ecosystem categories relative to their poten-
Today there is vigorous debate within the conservation com- tial for biodiversity conservation (Fig. 2.22). The criteria
munity regarding the proper foundational approach to biodi- associated with the Red List of Ecosystem Categories is
versity conservation. Traditionally, conservation efforts have becoming an increasingly used assessment tool for
focused on individual species. More recently, many conserva- integrating ecosystem and biodiversity conservation to pro-
tionists have begun to advocate an ecosystem-based approach, vide a common standard for coordinated global conservation
with preservation of rare species viewed as an “added bonus” efforts (IUCN 2016b).
(Thompson 2010). Some support for this change is advocated
by scientists desiring a more science-based paradigm for
conservation. Other support comes from conservation 2.9 Better Indicators for Biodiversity
managers and bureaucrats who feel overwhelmed and Conservation
exasperated by the seemingly insurmountable tasks of design-
ing species-specific conservation plans for an ever-increasing 2.9.1 The Value of Taxon-specific Surrogates
number of threatened species (Likens and Lindenmayer
2012). A third group, dominated by conservation planners We have already seen earlier the problems associated with
and policy makers, advocates integrated, multi-faceted using diversity levels of one indicator or surrogate species or
approaches to prevent conservation strategies from becoming species groups to represent diversity of other groups, as well as
overly narrow and failing to incorporate the full range of “best the problems of attempting to find correlations between diver-
practices” in conservation management. For example, Likens sity and endemicity, diversity and rarity, and diversity and
and Lindenmayer have shown that the recovery of northern conservation value (Kerr 1997; Lamoreux et al. 2006; Grenyer
deciduous forests from clearcutting, a process involving and et al. 2006). But can the diversity found in one taxonomic level
affecting the entire forest ecosystem, is dependent on a rela- (for example, the number of families in the group) provide an
tively small number of species-specific responses to such indicator of biodiversity at other taxonomic levels in the same
disturbance which could be better managed with an ecosystem group? Likewise, could changes in the status of the surrogate or
approach to conservation (Likens and Lindenmayer 2012). “indicator” taxon be accurate indicators of changes in the
Similarly, research on Australia’s threatened Leadbeater’s biodiversity of other taxonomic levels in that group? In other
Possum (Gymnobelideus leadbeateri) has revealed that the words, are there taxon-based indicators, or “taxon surrogates”
causes of its decline have been primarily related to ecosystem whose presence, absence, or richness is correlated with changes
(forest) structure. As a result, habitat requirements for the in biodiversity in a particular taxonomic group. For example,
possum have become guidelines for logging practices that could an estimate of species richness of a designate group, such
will not only protect this species but be used to identify as reptiles, be predicted by using a surrogate variable more
ecological reserves that need protection from logging (Likens easily estimated or known from previous studies, such as reptile
and Lindenmayer 2012). family richness?
2.9 Better Indicators for Biodiversity Conservation 69
A taxon-based indicator approach has potential for wider organism groups; plants, amphibians, reptiles, and mammals.
applicability than a single indicator species approach Their choice of these groups was based on: (1) the need to
because, instead of relying on a particular species, with an limit the number of groups to keep the index from becoming
equally particular range and distribution, it relies on a rela- unmanageable; (2) popular appeal and established efforts to
tionship between richness in the same group of organisms at conserve species in these groups; and (3) availability of
lower and higher taxa (for example, a positive correlation reliable regional information for these groups.
between species richness and family richness in the same Williams et al. divided the world into grid cells of
group), and therefore could be used in any locality. Although 611,000 km2 at intervals of 10 longitude, and then, using a
an intuitively appealing concept, success of this approach in variety of existing data sets, calculated three measures of
real investigations has been mixed. Let us look at one “family richness” for each cell. Absolute family richness
example. was obtained by summing local family richness counts (num-
Mexico is one of the world’s 17 “megadiversity” countries ber of families in the grid cell) for each of the four groups
which contain 66–77% of the world’s known species (Gaston according to the formula
and Spicer 2004:89). In Mexico, virtually all of the world’s
biomes are represented, and floristic richness is estimated to Absolute family richness ¼ f p,1 þ f a,1 þ f r,1 þ f m,1
be between 22,000 and 30,000 species. Here plant taxono-
mist Jose Villaseñor and his colleagues applied a surrogate where f is the number of families in each group, designated
approach by using existing inventories of local and regional by subscript (i.e., p for plants, a for amphibians, r for reptiles,
floral to see if richness in higher taxa could predict species and m for mammals). Proportional family richness is deter-
richness in vascular plants (Villaseñor et al. 2005). They mined by summing the local proportion of family richness in
found that predictive power depended on both taxon and the different major groups through the formula
vegetation type. When all families of vascular plants were
considered together, variation in plant genera richness Proportional family richness ¼ f p,1 =F p þ f a,1 =F a

explained 85% of variation in species richness. Explanatory þ f r,1 =F r þ f m,1 =F m :
power increased if analysis was restricted to a particular
vegetation type, in which case genera richness could explain The new term in this formula, F, represents the total world-
up to 95% of variation in species richness! Unfortunately, wide number of families in each group. Thus, it effectively
many existing inventories and other databases do not specify equalizes the contributions of groups with different numbers
to the level of genus, but only to family, and family richness of families and therefore weights the index to favor areas that
was a less reliable indicator, explaining only 64% of variation have a greater proportion of the families in each group, not
in species richness. Villaseñor et al. believe that, based on simply the areas that have the greatest number of families. A
previous studies in Mexico, about 77% of plant families in a third measure, proportional family richness weighted for
newly-studied area could be identified within 2 years. Such species richness, is calculated as.
identification could potentially be accomplished in even less
time if the inventory was focused on identifying plants only Sp( fp, 1/Fp) + Sa( fa, 1/Fa) + Sr( fr, 1/Fr) + Sm( fm, 1/Fm).
to family level from the start. Regarding the ability of this
approach to predict species richness in new areas, they The new term, S, is the total number of species in each group
asserted that “higher taxon/species function may be highly (Williams et al. 1997).
predictive, particularly when analyses are restricted to Using this method, Williams et al. found a pattern of
ecologically homogeneous regions” (Villaseñor et al. increasing diversity with decreasing latitude as well as close
2005:237). correlation of the three indices with one another (0.949–0.991
in Spearman’s rank correlation coefficients). Given the corre-
lation of results, it is not surprising that the methods made
2.9.2 Can Taxon Surrogates Analyze Global similar identifications of regions with high biodiversity. Cen-
Patterns of Biodiversity? tral and southern Columbia, Nicaragua, southern Mexico and
southern peninsular Malaysia were all identified as biodiver-
If taxonomic surrogates can work for one group of organisms sity “hotspots” in at least two of the three methods.
(vascular plants), in one country (Mexico), would it be possi- The approach used by Williams et al. has weaknesses, the
ble to combine family indices of different kinds of organisms most obvious being the exclusion of groups more species rich
to produce an aggregate biodiversity index for regions that than those chosen. Williams et al. excluded, among other
could be mapped worldwide, an approach that might avoid groups, insects, fungi, and bacteria, each of which contains
the problems associated with the use of individual surrogate more species than all vertebrate groups combined. And the
species? Williams et al. (1997) attacked the problem in a study also does not tell us if patterns of biodiversity at family
classic analysis by combining family richness in four major levels are similar at higher levels, like orders or classes. But
the indices do provide estimates of diversity that cover a have been the first systematic attempt to define an algorithm
broad range of groups of conservation concern, and the of selection criteria for prioritizing selection of conservation
method can be applied to other groups as reliability of data reserves (Pressey 2002). From this first effort and those that
improves and conservation efforts expand to include followed, systematic conservation planning (SCP) has
other taxa. developed into a recognized approach for conservation prior-
itization that is widely used today. One of its outcomes has
been the development of a framework for identifying conser-
vation priorities in reserve design by integrating the relative
It is not possible to assess, manage, or preserve all
threat of loss of an area or system with its irreplaceability
biodiversity in a system or management unit. Based
(ecological uniqueness). We could display these criteria visu-
on available techniques and approaches, what do you
ally in two dimensions (Fig. 2.23), with the highest priority
think would be the “best” way for a manager to evalu-
for conservation assigned to areas that rank high in both
ate biodiversity, and what criteria would you provide
criteria (the upper right portion of the graph). But such
the manager to determine when “enough” diversity has
areas might not always be immediately available for protec-
been preserved?
tion, so conservationists following an SCP approach should
also recognize opportunism as a third criterion, a strategy of
acquiring areas that are available for protection even if they
rank lower regarding threat or irreplaceability. Such informed
opportunism takes advantage of opportunities to acquire pro-
2.10 How Do We Prioritize Areas tection for available areas that rank highest in threat and
for Biodiversity? irreplaceability criteria, whereas less desirable uninformed
opportunism “protects” areas low in threat or irreplaceability,
2.10.1 Current Global Prioritization Strategies i.e., areas that may not need protection, and which might
constitute a waste of limited financial and other resources
Jamie Kilpatrick’s landmark publication, “An Iterative needed to create protection.
Method for Establishing Priorities for Implementing Conser- Today there are identifiably nine major global biodiversity
vation Action” (Kilpatrick 1983), is considered by many to conservation strategies, or, more precisely, templates for
Fig. 2.23 A framework for identifying conservation priorities for threat and irreplaceability. “Uniformed opportunism” “protects” areas
protected areas based on threat and irreplaceability. Highest priority with low ranks in these criteria, and so reflects potentially poor use of
areas would be located in the upper right portion of the graph due to conservation resources and effort. (Reprinted by permission from
high rankings in both threat and irreplaceability. The region of Wiley, from Pressey, R.L., Bottrill, M.C., 2008. Opportunism, Threats,
“informed opportunism” (central portion of graph) reflects intentional and the Evolution of Systematic Conservation Planning. Conservation
departure from highest priority areas to take advantage of opportunity to Biology 22, 1340–1345. # 2008 Society for Conservation Biology)
acquire areas currently available which have moderate to high ranks in
2.10 How Do We Prioritize Areas for Biodiversity? 71
identifying and selecting land areas to conserve for biodiver-

sity, and all of them employ some variation of threat and
irreplaceability criteria. These are: (1) the Crisis Ecoregion
(CE) strategy, which prioritizes conservation of ecosystems
facing the highest threats of destruction and degradation;
(2) Biodiversity Hotspots (BH), which selects landscapes
with the highest species diversity per unit area; (3) Endemic
Bird Areas (EBA), which prioritizes areas with the highest
densities of endemic bird species; (4) Centers of Plant Diver-
sity (CPD), which targets areas with exceptional plant diver-
sity per unit area; (5) Megadiversity Countries, which
identifies nations with the highest levels of biodiversity, and Fig. 2.24 Global biodiversity conservation priority templates placed
designs conservation plans sensitive to national interests and within the conceptual framework of irreplaceability and vulnerability.
boundaries; (6) Global 200 Ecoregions (G200), a science- (a) Reactive approaches focus on the protection of areas of high vulner-
based global ranking of the Earth’s most biologically out- ability and immediate threat. Proactive approaches focus on areas with
high biodiversity still relatively unaffected by human influence. (b) Four
standing terrestrial, freshwater and marine habitats; (7) High approaches that do not incorporate vulnerability as a criterion but
Biodiversity Wilderness Areas (HBWA), identifying areas consider only irreplaceability (uniqueness) in conservation.
that combine high levels of biodiversity, relatively low Abbreviations explained in text. (Reprinted by permission from
human population density, and high landscape connectivity; AAAS, from Brooks, T.M., Mittermeier, R.A., da Fonseca, G.A.B.,
Gerlach, J., Hoffmann, M., Lamoreux, J.F., Mittermeier, C.G., Pilgrim,
(8) Frontier Forests (FF), a strategy targeting the world’s J.D., and Rodrigues, A.S.L., 2006. Global Biodiversity Conservation
remaining large intact natural forest ecosystems; and (9) Last Priorities. Science 313, 58–61. # 2006 American Association for the
of the Wild (LW), a strategy placing conservation priority on Advacement of Science)
areas representing the largest and relatively “wildest” (lowest
human population and environmental impact) places in each
of their biomes (Brooks et al. 2006). prioritizes wilderness areas is amenable to landscape-level
Each strategy uses a different approach to biodiversity management practices at large spatial scales that can be
conservation, with unique, and, sometimes, complementary applied over fairly long time periods, while a strategy that
strengths and weaknesses, but all can be understood in terms prioritizes irreplaceability and threat must often act quickly at
of the two criteria foundational to SCP. In using these criteria, specific sites.
such templates can be classed as proactive or reactive in their Global conservation priority strategies that arise from SCP
relative emphasis on irreplaceability (rarity and uniqueness) are effective at raising money because they offer easily
or threat (vulnerability to species in the area or to the area identifiable targets that attract support of large donors. How-
itself) (Fig. 2.24). ever, the same strategies reveal the limitations of such
Reactive strategies set targets based on levels of threat templates, for they have often failed to inform or affect actual
(risk of loss). For example, CE is a highly reactive strategy, conservation management and implementation because they
prioritizing conservation areas according to immediate risk. are not designed to address it. Areas of high biodiversity are
Proactive strategies target areas which are still relatively also often areas with high monetary values of the ecosystem
pristine, thus hoping to save intact systems and avoid the services they provide (Turner et al. 2007), but not all of these
effort and expense of trying to restore degraded ones. HBWA strategies protect biodiversity equally well, nor do they con-
is an example of a proactive strategy, attempting to conserve serve the same amount of ecosystem services (Table 2.12,
areas that are still relatively unaffected by negative human Fig. 2.26) (Turner et al. 2007). Recall that biodiversity con-
influences. In contrast, conservation templates that focus on servation and safeguarding ecosystem services have histori-
endemic species, such as EBA, use irreplaceability (endemic cally been the two major objectives of conservation biology,
birds occur in only one place or region) as the guiding so differences in effectiveness on both points matter. This
criterion. Some strategies, such as those that focus on multiplicity of strategies does a good job hitting a variety of
endemic species or biodiversity alone, consider only irre- conservation targets, but exposes one of the most important
placeability, not vulnerability. Templates that prioritize the problems in conservation: poorly defined goals and, in many
same thing tend to place priority on many of the same areas, cases, inefficient strategies for achieving them. This problem
while those that prioritize different things have much less is not necessarily a criticism of SCP but is often a failing of
overlap in their selection of priority conservation areas. those who apply it without sufficient insight or foresight
(Fig. 2.25). Seventy-nine percent of all land on Earth is regarding its consequences. In the case of these nine specific
prioritized in at least one of these systems, but the different global conservation strategies, it is precisely because of their
templates require different approaches. A strategy that specificity and, often, narrowness of focus, that no one
Fig. 2.25 Maps of nine global biodiversity conservation priority Pilgrim, J.D., and Rodrigues, A.S.L., 2006. Global Biodiversity Con-
templates. Abbreviations explained in text. (Reprinted by permission servation Priorities. Science 313, 58–61. # 2006 American Association
from AAAS, from Brooks, T.M., Mittermeier, R.A., da Fonseca, G.A. for the Advacement of Science)
B., Gerlach, J., Hoffmann, M., Lamoreux, J.F., Mittermeier, C.G.,
Table 2.12 Estimated ecosystem service value (ESV) within templates (Fig. 2.26) for global biodiversity conservation
Area Total ESV (billion US$ per year) Percentage Concordance
(million Mean ESV (US$ above indexd
Global template km2) per km2per year) Observed Randoma Maximumb randomc (percentage)
High-biodiversity wilderness 11.5 200,720 2314 701 4708 230 40.3
areas (Mittermeier et al. 2003)
Frontier forests (Bryant et al. 13.2 188,224 2387 803 5151 210 38.7
1997)
Most proactive 7.6 217,356 1659 464 3681 257 37.1
Global 200 ecoregions (Olson 53.8 86,857 4671 3270 7466 43 33.4
and Dinnerstein 1998)
Last of the wild (Sanderson 35.0 98,356 3440 2127 6838 62 27.9
et al. 2002)
Megadiversity countries 49.8 77,457 3860 3031 7340 27 19.2
(Mittermeier et al. 1997)
Endemic bird areas 13.8 88,710 1222 838 5301 46 8.6
(Stattersfield et al. 1998)
Centers of plant diversity 12.2 83,779 1023 743 4888 38 6.8
(WWF and IUCN 1994–1997)
Most reactive 12.1 76,057 917 734 4849 25 4.5
Biodiversity hotspots 23.0 69,071 1588 1398 6289 14 3.9
(Mittermeier et al. 2004)
Random terrestrial km2 – 60,813 – – – – 0.0
Crisis ecoregions (Hoekstra 42.7 46,038 1967 2598 7118 24 14.0
et al. 2005)
Reprinted by permission from Oxford University Press, from Turner, W.R., Brandon, K., Brooks, T.M., Costanza, R., da Fonseca, G.A.B., Portela,
R., 2007. Global conservation of biodiversity and ecosystem services. Bioscience 57, 868–873. # 2007 American Institute of Biological Sciences
Note: All monetary values are in 2005 US dollars
a
ESV in randomly selected 1-km2 cells, with the total area equivalent to that of each template
b
Maximum ESV attainable for the total area equivalent to that of each template
c
Significance of percentage deviation from random is evaluated with a randomization test (N ¼ 10,000, p < 0.001 in all cases)
d
Percentage of ESV represented beyond that expected at random, relative to the maximum attainable
Fig. 2.26 Spatial concordance of global biodiversity priorities and values for both. (Reprinted by permission from Oxford University Press,
ecosystem service value (ESV). Increasing intensities of green and red from Turner, W.R., Brandon, K., Brooks, T.M., Costanza, R., da
represent, respectively, increasing rank ESV and increasing rank con- Fonseca, G.A.B., Portela, R., 2007. Global conservation of biodiversity
sensus biodiversity priority. White corresponds to low values for both and ecosystem services. Bioscience 57, 868–873. # 2007 American
variables, black to high values for both, and shades of gray to covarying Institute of Biological Sciences)
strategy provides a unifying approach to prevent the global ongoing and increasing effects of global climate change
ecosystem from experiencing unsustainable degradation and (ClimateWise), which, if not accounted for, will render con-
biodiversity loss or to conserve the entire array of ecosystem servation strategies ineffective due to changing conditions in
services on which human beings depend. the very areas targeted for preservation. A ClimateWise strat-
egy would prioritize conservation targets where the ecosys-
tem chosen for conservation could be maintained under
2.10.2 Developing More Advanced Integrated expected future climate change (Freudenberger et al. 2013).
Global Conservation Strategies The pathways to actually generate a scheme of conservation
priorities require further information and assessment, but
In a pointed critique of these strategies, Lisa Freudenberger such information is available, and, from it, intelligent assess-
of the University of Sustainable Development Eberswalde ment can be made. When Freudenberger and her colleagues
(Germany) and her colleagues proposed a revised method employed their criteria and integrated all three considerations
of prioritizing conservation goals addressing three essential on a global scale, they generated highest values for the
components for global effectiveness. They argue that any Amazon region and tropical Andes of South America, parts
model for achieving reduction in worldwide biodiversity of western North America, and the Appalachian region of
loss must be eco-functionally wise (EcoWise) in taking into eastern North America (Fig. 2.27). Freudenberger’s team
account, from science, the way that ecosystems supporting noted that their choices, based on this more holistic approach,
biodiversity actually function and work. Further, setting represented “those ecosystems that exhibit strong ecological
priorities in conservation must be socioeconomically wise and environmental functionality, and this provides an impor-
(SocioWise) in addressing human economic and social tant global service in mitigating against the impacts of cli-
motivations that will sustain the conservation effort. Eco- mate change” (Freudenberger et al. 2013:1273).
nomic resources for conservation are limited and setting Although conservationists might disagree regarding areas
conservation priorities must account for social and economic to be prioritized for protection, developing and employing a
conditions of areas considered for conservation. Priority, more globally integrated strategy of biodiversity conserva-
therefore, should be given to areas where economic, social, tion is an essential and urgent task in conservation biology.
and political conditions are such that conservation efforts Failure to develop more systematic and effective systems of
would actually work under existing conditions. Finally, the prioritizing areas for conservation, such as the one developed
model for making conservation decisions must address by Freudenberger and her colleagues, will almost certainly
Fig. 2.27 Conservation priority-setting indices combining EcoWise, (Reprinted by permission from Springer Nature, from Freudenberger, L.,
SocioWise, and ClimateWise priorities with equal weighting of all Hobson, P., Schluck, M., Kreft, S., Vohland, K., Sommer, H., Reichle,
indicators (a), SocioWise and ClimateWise priorities (b), EcoWise and S., Nowicki, C., Barthlott, W., Ibisch, P.L., 2013. Nature conservation:
ClimateWise priorities (c), and EcoWise and SocioWise priorities priority-setting needs a global change. Biodiversity and Conservation 22,
(d), Blue to yellow areas indicate transitions from low to high values. 1255–1281. # 2013 Springer Science+Business Media Dordrecht)
result, not only in loss of biodiversity and essential global diversity at a single site (alpha diversity), choose a site
ecosystem services for the world, but loss of credibility for with high species richness. But what if the management
the global conservation community. objective is to maximize the differentiation of biodiversity
Conservationists must understand these and other in the larger landscape (beta diversity)? In that case, a
integrated global approaches so that they can determine strategy of simply reserving all sites with high levels of
which ones to employ for particular conservation problems. species richness will have little value if different areas con-
But it is also essential that they move beyond generalized tain the same species. Most sites are dominated by generalist
global strategies to landscape- and site-specific management species, even sites with high levels of species richness.
plans that conserve biodiversity. Ecological specialists may occur at sites with relatively
low species richness. At landscape levels, the key is not
to select the sites with highest richness, but to select sites
2.10.3 Management Approaches to Biodiversity in which species compositions are most dissimilar to one
at Landscape Levels another. By protecting areas of less biological similarity,
protection of regional endemic species and ecological
2.10.3.1 Gathering Appropriate Background Data specialist species is enhanced, and these species may
A first step in conserving biodiversity is to appreciate contribute most to biodiversity at regional and landscape
differences in spatial scale. If the goal is to maximize local levels.
2.10.3.2 Maintaining Ecological and Evolutionary conservation areas (FCAs) are identified as “a geographic
Processes Promoting Biodiversity domain that maintains focal ecosystems, species, and
Although accurate inventories of diversity are important, supporting ecological and evolutionary processes within
biodiversity is neither maintained nor managed simply by their natural range of variability” (Poiani et al. 2000). FCAs
counting biological “things.” The processes that shape biodi- are delineated as sites, landscapes, and networks. Functional
versity, although they may operate over long time spans, sites conserve one or more endangered species or rare
should also be subjects of conservation management. The ecosystems, typically at a local scale. Functional landscapes
most important among these are maintenance of gene flow, encompass full terrestrial and aquatic habitat gradients and a
creating resources for habitat and niche specialization, and diversity of ecological and evolutionary processes needed to
maintaining effective population sizes. maintain those gradients and species within them. Functional
Gene flow between population subunits is maintained by networks provide spatial context, configuration, and connec-
connectivity between subunits. In cases where physical tivity for regional-scale species conservation (Poiani et al.
connections, such as habitat corridors, can enhance connec- 2000). A similar approach developed by Jonathan Higgins
tivity (movement and interchange of individuals), managers and his colleagues has been used for aquatic systems. Here
may be able to increase gene flow at varying spatial scales. classification starts with determination of an aquatic zoogeo-
More specific considerations of how this is done will be graphic unit (AZU) (a watershed), which serves as the overall
considered in subsequent chapters on genetics (Chap. 5), planning unit. Within an AZU are one or more ecological
populations (Chap. 6), and habitat and ecosystem conserva- drainage units (EDUs). Within a single EDU are multiple
tion (Chaps. 7, 8, and 9). aquatic ecological systems (AESs). Within a single AES,
Creating resources for habitat and niche specialization macrohabitats can be managed for individual species or spe-
requires a manager to have knowledge of species-specific cies groups (Higgins et al. 2005). Using GIS, Higgins et al.
needs, and the skills to meet them. Various forms of habitat created an inventory of mapped and classified units used to
alteration, including prescribed fire; permanent, periodic, or identify and differentiate spatial patterns of aquatic
seasonal flooding; alteration and removal of vegetation; or ecosystems. Conservation planning and priorities are then
addition of key resources (for example, nest site structures) developed to preserve a diversity of system types, rather
can, when intelligently applied, increase resources for niche than simply individual populations.
specialization, contributing to an environment in which more The Nature Conservancy (TNC) is now developing a more
species, especially those with more specialized needs and systemic approach to the use of FCAs, creating and con-
preferences, can persist. Particular techniques, and case stantly updating a portfolio of such areas within and across
histories that illustrate them, will be examined in greater ecoregions. Through this portfolio approach, TNC works
detail in Chaps. 7, 8, and 9 in our examination of habitat, with partners to conserve a full array of ecosystems and
landscape and ecosystem conservation. viable native species. Individual ecoregional portfolios are
Maintaining effective population sizes permits the end products of ecoregional conservation planning, with
populations to retain genetic diversity and environmental each portfolio containing a selected set of places representing
adaptability. Making accurate and precise estimates of what the full distribution and diversity of native species, natural
constitutes an effective population size requires knowledge communities and ecosystems in an ecoregion (The Nature
of a species mating system as well as the number of Conservancy 2015).
individuals present. The methods for making such estimates
will be an important focus of our examination of conservation
genetics in Chap. 5.
Many US conservation laws are either species-based
(e.g., Endangered Species Act, Marine Mammal Protec-
tion Act) or impact-based (e.g., National Environmental
2.10.4 Regional Biodiversity – Defining
Policy Act, Clean Water Act). None are designed to
Functional Conservation Areas
protect biodiversity per se. What would be the value of a
conservation statute written explicitly to protect biodi-
Because distribution of global biodiversity is complex,
versity, and why might it require “functional conserva-
mapping and protecting key areas requires taxon-specific
tion areas” as management units to be effective?
approaches, careful conceptual methods, and sophisticated
technologies for spatial problem solving. Poiani et al.
(2000) addressed this problem through the development of
“functional conservation areas.” By examining biodiversity Regional and landscape conservation criteria, no matter
at different spatial scales, Poiani et al. (2000) defined how well-conceived, are even more valuable if
ecosystems and species at four different levels: local, inter- complemented with reliable information on species, habitat
mediate, coarse, and regional. Within these levels, functional and ecosystem distributions at multiple scales. Thus,
conservationists working to preserve biodiversity increas-

ingly employ Geographic Information Systems (GIS) and it. Biodiversity is a concept that requires further refine-
the related Gap Analysis Program (GAP), which relates the ment, and it cannot stand alone apart from other con-
level of conservation protection of a land unit to the amount servation priorities. Current measurements of
of biodiversity and number of threatened species it contains biodiversity, with their emphasis on species richness
(Chap. 7) to supply information necessary for intelligent and evenness, do not always reveal correlations
management and conservation decisions. Scientific between diversity and conservation value. New indices
approaches utilizing GIS can involve organizing and that address taxonomic uniqueness and ecological
overlaying thematic data, such as soils, vegetation, hydrol- importance must be developed and used in conjunction
ogy, and dominant vegetation, within a defined area, and then with traditional measures of diversity if biodiversity is
relating such data, along with land use patterns and locations to provide meaningful information about the relative
of existing nature reserves, with the distribution of value of different community and landscape
endangered or endemic species. Using such a technique, assemblages. However, the choice of method(s) to
conservation biologists and land use planners can determine: quantify biodiversity must be justified by the priorities
(1) what proportion of an area’s biodiversity is protected in an of the conservation goals.
existing distribution of nature preserves and under existing We will increasingly see the concept of biodiversity
land use practices; (2) whether such protection can be incorporated into overall strategies of international
expected to permit the persistence of endangered or endemic conservation organizations and into the management
species and (3) the best location and arrangement of new plans and assessments of government conservation
nature preserves or the best areas to attempt to change current agencies. The challenge will be to design such
land use patterns. Only when conservation biologists effec- strategies and plans with care and insight so that they
tively integrate global patterns of biodiversity, taxon- and truly assess the condition of biodiversity at appropriate
site-specific variations, well-organized conceptual conservation and management scales, and not merely
frameworks for biodiversity protection, and technologically pay lip service to the concept of biodiversity while
advanced data analysis and land use planning toward the goal ignoring, or even harming, the substance of it.
of protecting biodiversity at multiple scales is there reason-
able hope that biological diversity will persist in any area or
ecosystem.
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The Anthropocene: Conservation
in a Human-Dominated Nature 3
If we are to have an accurate picture of the world, even in its present diseased condition, we must interpose
between the unused landscape and the misused landscape a landscape that humans have used well.
Wendell Berry (1995:64)
Keywords Stratigraphy (ICS), released the results of its vote to designate

Anthropocene · Invasive species · Novel ecosystems · a new geologic epoch – the Anthropocene – to officially mark
Urban ecological systems · Urban conservation · Anthro- the manifold and planetary effects that human beings are now
pogenic biomes · Anthromes · Urban biodiversity · creating on the ecology, natural processes, and even geologi-
Neoprotectionism · Convivial conservation cal records of planet Earth (Subcommission on Quaternary
Stratigraphy 2019) (Fig. 3.1). Among the 33 members pres-
ent for the vote, 29 were in favor and four opposed.
The vote of a scientific committee (or, in this case, sub-
Overview commission), is not normally newsworthy. But this vote is
In this chapter, you will learn: different. The ballots that were cast were representative of the
increasing recognition that humans have become the domi-
1. Why human actions have become dominant ecolog- nant ecological force on earth, shaping and reshaping the
ical forces in Earth systems. interactions of all other species. This transition in relationship
2. How such actions are leading to the development of is global and marked by a variety of processes, even
novel ecosystems and biomes. influencing the geologic record on a planetary scale, and
3. How non-native species moved by humans affect ushering in what may now be called the Anthropocene
natural and novel ecosystems. Epoch. And this transition forces us to confront and to ques-
4. How the growth of urban areas and populations tion many cherished assumptions of our traditional
affects global biodiversity. approaches to conservation. The purpose of this chapter is
5. How conservation strategies must adapt to a human- to determine if, indeed, conservation can continue in a
dominated world. human-dominated world. And if it can, what kind of conser-
vation must it be?
These questions will not wait. The AWG now plans to
submit a formal proposal for designating the new epoch to the
ICG, which oversees the official geologic time chart, by
2021. To receive formal designation, geological time periods
3.1 Dawn of the Anthropocene: Human must be defined by their lower boundary, that is, their begin-
Impacts Define a Geologic Epoch ning at a globally synchronous “marker” or “signal” which
identifies the start of the new epoch (Lewis and Maslin 2015).
3.1.1 Scientists Cast a Vote
The global marker must signal an event in stratigraphic
material, such as rock, sediment, or glacier ice, known as a
In 2019, the Anthropocene Working Group (AWG), a
Global Stratotype Section and Point (GSSP) that indicates
34-member working group under the Subcommission on
changes to the Earth system (Lewis and Maslin 2015).
Quaternary Stratigraphy of the International Commission on

82 3 The Anthropocene: Conservation in a Human-Dominated Nature
fifteenth century (Table 3.1, Lewis and Maslin 2015), as all of

these mark historic transitions in human interaction with the
global environment. Geologists Simon Lewis and Mark
Maslin are precise and specific on the arguments for making
14
C the marker of choice. “The global 14C peak provides an
unambiguously global change in a number of stratigraphic
deposits. . .We propose that the GSSP marker should be the
14
C (radioactive carbon 14) peak, at 1964, within dated
annual rings of a pine tree (Pinus sylvestris) from King
Castle, Niepołomice, 25 km east of Krako, Poland” (Lewis
and Maslin 2015:177). But this radioactive marker, although
scientifically and officially important to make the argument
for official designation of a new geologic epoch, is not the
only sign that humans now have an unprecedented role in
affecting and changing the Earth. Others include recent
deposits of new minerals and rock types that include alumi-
num, concrete, and plastics. Fossil fuel combustion by
humans has now resulted in the formation of black carbon,
inorganic ash spheres, and spherical carbonaceous particles
Fig. 3.1 A sediment core from west Greenland displaying an abrupt worldwide, with a near-synchronous global increase in these
stratigraphic transition from glacial sediments to nonglacial organic materials around 1950. Recent stratigraphic deposits show
matter, one of numerous “markers” of global warming and other anthro- that sedimentary fluxes in the Earth’s strata have intensified,
pogenic effects that are now evident in geologic strata. (Photo reprinted
by permission from J. Briner)
as well as enhanced erosion caused by human deforestation
and road construction. There are additional geochemical
signatures of elevated levels of polyaromatic hydrocarbons,
As Colin Waters of the British Geological Survey and his polychlorinated biphenyls, pesticide residues, concrete
colleagues expressed it in the pages of Science in 2016, “Any residues, elevated carbon dioxide and methane levels, as
formal recognition of an Anthropocene epoch in the geologi- well as increased presence of lead in sediments coinciding
cal time scale hinges on whether humans have changed the with the use of leaded gasoline between ~1945 and 1950
Earth system sufficiently to produce a stratigraphic signature (Fig. 3.2) (Waters et al. 2016). Soil nitrogen and phosphorus
in sediments and ice that is distinct from that of the Holocene concentrations have doubled in the past century because of
epoch. . .” (Waters et al. 2016:137). Regarding the existence increased fertilizer use, and these have generated widespread
of such a stratigraphic signature or GSSP, the AWG reported signatures in lake strata and nitrate levels in Greenland ice
“The sharpest and most globally synchronous of these higher than any time during the last 100,000 years (Fig. 3.2).
signals, that may form a primary marker, is made by the Compared to 14C deposits, these activities have not left
artificial radionuclides spread worldwide by the thermonu- stratigraphic markers as precise or unambiguous in the
clear bomb tests from the early 1950s”, specifically a sudden Earth’s sediments, and they are not as globally synchronous,
spike in the presence of a radioactive isotope of carbon with but all add support to the argument that delineation of a new
an atomic weight (the sum of the protons and neutrons in one geologic epoch is warranted.
atom of the element) of 14 (14C, “normal,” nonradioactive Geologic time is divided into a hierarchical series of units
carbon has an atomic weight of 12 (12C) (Subcommission on of smaller and smaller time spans (Fig. 3.3). The present
Quaternary Stratigraphy 2019). period, according to The Geologic Time Scale 2018, is the
Holocene Epoch (Greek for ‘entirely recent’) which began
11,650 years before present within the Quaternary Period
3.1.2 The Broader Debate (which started 2.588 million years ago). The concept of a
geologic age defined by human activity and effect is not new,
In recent years, some scientists have argued for designating as humans count newness. The Holocene (“Recent Whole” or
the beginning of the Anthropocene to coincide with recent “Entirely Recent”) Epoch had already been designated for the
global temperature anomalies (climate warming), the expan- post-glacial period of the past ten to twelve thousand years by
sion of agriculture and livestock cultivation that started over the International Geological Congress in Bologna in 1855
5000 years ago, the surge in mining activity that began more (Italy) (Crutzen and Stoermer 2000), although the designa-
than 3000 years ago (Monastersky 2015), or the clash tion did not become official until 2008 with the official
between Old and New World people that began in the late recognition by the ICS (Monastersky 2015). In 1854, 1 year
3.1 Dawn of the Anthropocene: Human Impacts Define a Geologic Epoch 83
Table 3.1 Potential start dates for a formal Anthropocene Epoch

Primary
stratigraphic
Event Date Geographical extent marker Potential GSSP datea Potential auxiliary stratotypes
Megafauna 50,000–10,000 year Near-global Fossil None, diachronous Charcoal in lacustrine deposits
extinction BP megafauna over ~40,000 year
Origin of ~11,000 year BP Southwest Asia, Fossil pollen None, diachronous Fossil crop pollen, phytoliths,
farming becoming global or phytoliths over ~5000 year charcoal
Extensive ~8000 year BP to Eurasian event, global CO2 inflection None, inflection too Fossil crop pollen, phytoliths,
farming present impact in glacier ice diffuse charcoal, ceramic minerals
Rice 6500 year BP to Southeast Asian event, CH4 inflection 5020 year BP CH4 Stone axes, fossil
production present global impact in glacier ice minima domesticated ruminant
remains
Anthropogenic ~3000–500 year BP Local event, local Dark high None, diachronous, Fossil crop pollen
soils impact, but organic matter not well preserved
widespread soil
New–Old 1492–1800 Eurasian–Americas Low point of 1610 CO2 minima Fossil pollen, phytoliths,
World collision event, global impact CO2 in glacier charcoal, CH4, speleothem
ice δ18O, tephra ℑ
14 15
Industrial 1760 to present Northwest Europe Fly ash from ~1900; diachronous N: N ratio and diatom
revolution event, local impact, coal burning over ~200 year composition in lake sediments
becoming global
Nuclear 1945 to present Local events, global Radionuclides 1964 14C peakb 240
Pu: 239Pu ratio, compounds
weapon impact (14C) in tree- from cement, plastic, lead and
detonation rings other metals
Persistent ~1950 to present Local events, global For example, Peaks often very Compounds from cement,
industrial impact SF6 peak in recent so difficult to plastic, lead and other metals
chemicals glacier ice accurately dateb
Reprinted by permission from Springer Nature, from Lewis, S.L., Maslin, M.A., 2015. Defining the Anthropocene. Nature 519, 171–180.
# 2015 Macmillan Publishers Limited
Note: For compliance with a Global Stratotype Section and Point (GSSP) definition, a clearly dated global marker is required, backed by correlated
auxiliary markers that collectively indicate global and other widespread and long-term changes to the Earth system. BP, before present, where
present is defined as calendar date 1950
a
Requires a specific date for a GSSP primary marker. ℑ From Huaynaputina eruption in 1600
b
Peak, rather than earliest date of detection selected, because earliest dates reflect available detection technology, are more likely influenced by
natural background geochemical levels, and will be more affected by the future decay of the signal, than peak values
prior to the Bologna designation, Welsh geologist and pro- biodiversity (Chap. 2). Human activity has brought Earth’s
fessor of theology, Thomas Jenkyn, may have been the first plant and animal species to the brink of the Earth’s sixth great
to publish the idea of an evidence-based human geologic time extinction event, increasing the species extinction rate by one
unit (Jenkyn 1854a, b, Lewis and Maslin 2015). He describes thousand- to ten thousand fold in tropical rain forests alone
the then present day as “the human epoch” based on the likely (Crutzen and Stoermer 2000). Humans contribute twice the
future fossil record (Jenkyn 1854a, cited in Lewis and Maslin atmospheric sulfur dioxide emissions as natural processes,
2015). Jenkyn wrote, “All the recent rocks, called in our last apply more nitrogen to the Earth’s surface and move more
lesson Post-Pleistocene, might have been called soil on its surface than all natural processes combined
Anthropozoic, that is, human-life rocks” (Jenkyn 1854b, (Crutzen and Stoermer 2000, Ellis and Ramankutty 2008).
cited in Lewis and Maslin 2015:172). By 1864, US author They capture 25–35% of the ocean’s primary production,
George Perkins Marsh had published Man and Nature, 40% of primary surface production, and more than 50% of
detailing the global extent of modifications of the Earth by all accessible freshwater (Crutzen and Stoermer 2000). More
human action. In 1873, humanity’s increasing environmental than 75% of the Earth’s ice-free surface has now been in
influence was even more explicitly stated by the Italian some way altered by human residence or activity. The less
geologist Antonio Stoppani whose described human effects than 25% that remain as “wildlands” support only 11% of the
as a “new telluric force [a force relating to the Earth] which in Earth’s present net primary production (Ellis and
power and universality may be compared to the greater forces Ramankutty 2008). We have come to the point in human
of earth,” referring to the “anthropozoic era” (Crutzen and history in which the traditional world maps of the Earth’s
Stoermer 2000:17). biomes are becoming outdated and inaccurate. The world
Not all of the effects of humans on the Earth are buried today is dominated by what some scientists are now calling
beneath the surface. Many are plain to see, including many anthropogenic biomes or “anthromes” (Ellis and
that we have already noted in our examination of global Ramankutty 2008).
Fig. 3.2 Summary of the

magnitude of key markers of
anthropogenic change that are
indicative of the Anthropocene.
(a) Novel markers, such as
concrete, plastics, global black
carbon, and plutonium
(Pu) fallout, shown with
radiocarbon (14C) concentration.
(b) Long-ranging signals such as
nitrates (NO3–), CO2, CH4, and
global temperatures, which
remained at relatively low values
before 1950, rapidly rose during
the mid-twentieth century and, by
the late twentieth century,
exceeded Holocene ranges.
(Reprinted by permission from the
American Association for the
Advancement of Science,
from Waters, C.N., Zalasiewicz,
J., Summerhayes, C., Barnosky,
A.D., Poirier, C., Galuszka, A.,
Cearreta, A., Edgeworth, M.,
Ellis, E.C., Ellis, M., Jeandel, C.,
Leinfelder, R., McNeill, J.R.,
Richter, D. d., Steffen, W.,
Syvitski, J., Vidas, D., Wagreich,
M., Williams, M., Zhisheng, A.,
Grinevald, J., Odada, E., Oreskes,
N., and Wolfe, A.P., 2016. The
Anthropocene is functionally and
stratigraphically distinct from the
Holocene. Science 351, aad2622–
aad2622. # 2016 AAAS)
3.1.3 The Anthropogenic Biome biological communities, even at regional and continental
scales. The signatures are a product of, as Waters et al., put
Biomes reflect a concept originally developed to describe it “three linked force multipliers: accelerated technological
large areas of regionally similar vegetation which generally development, rapid growth of the human population, and
occur in predictable patterns of latitude and climate (Fig. 3.4). increased consumption of resources. These have combined
Biomes came to be the most basic units ecologists would use to result in increased use of metals and minerals, fossil fuels,
to describe overall global patterns of ecosystem form, process and agricultural fertilizers and increased transformation of
and biodiversity (Ellis and Ramankutty 2008). The term was land and nearshore marine ecosystems for human use”
first suggested by the US ecologist Frederic Clements in the (Waters et al. 2016:2–3). The net effect of technological
early twentieth century, which he used as semi-acronym for development, population growth, and resource consumption
“biotic community of Möbius” (because Clements drew on has been a conversion of natural biomes to agriculture, cities,
the ideas of Karl Möbius, a nineteenth century German biol- roads, and other human constructs, and to the replacement of
ogist who was a pioneer in the development of ecology, wild animals and plants by domesticated species to meet
especially ecological distribution of communities). World growing demands for food. These problems have become
classification of terrestrial biomes began with eight (Allee worse since the beginning of the so-called “Great Accelera-
1949) or nine (Kendeigh 1961) biome types, but with time tion,” a period of dramatically increased economic activity
became more complex. and resource consumption that intensified these and other
What has appeared as anthropogenic signatures in geolog- geologic signals of the human effect on nature (Lewis and
ical sediments are indicators of changes that humans have Maslin 2015).
made in the surface and ecological processes of the Earth At the beginning of the eighteenth century (1700), nearly
itself, and which are now affecting the composition of half of the terrestrial biosphere was “wild.” These wild areas
GSA GEOLOGIC TIME SCALE v. 5.0

CENOZOIC MESOZOIC PALEOZOIC PRECAMBRIAN
MAGNETIC MAGNETIC BDY.
AGE POLARITY PICKS AGE POLARITY PICKS AGE PICKS AGE
PERIOD EPOCH AGE PERIOD EPOCH AGE PERIOD EPOCH AGE EON ERA PERIOD AGES
CHRON.
CHRON.
ANOM.
ANOM.
(Ma) (Ma) (Ma) (Ma) (Ma) (Ma)

HIST.
(Ma)
HIST
(Ma)
HOLOCENE 66.0
1 C1 QUATER- 0.012 30 C30 541
CALABRIAN
NARY PLEISTOCENE* 1.8 31 C31 MAASTRICHTIAN 251.90 EDIACARAN
2 C2 GELASIAN
2.58 70 Lopin- CHANGHSINGIAN 254.14
2A C2A PIACENZIAN 32 C32 72.1 635
PLIOCENE 3.600 gian WUCHIAPINGIAN
259.1 CRYOGENIAN
ZANCLEAN 33 260 720
PERMIAN
3 CAMPANIAN CAPITANIAN 750 NEOPRO-
CRETACEOUS
5 C3
5.333 C33 Guada- 265.1
80 WORDIAN 268.8 TEROZOIC
3A C3A MESSINIAN LATE lupian ROADIAN TONIAN
7.246 83.6 272.95
4
SANTONIAN KUNGURIAN
C4 86.3 ~283.5
NEOGENE
CONIACIAN 280
4A Cisura-
PROTEROZOIC
C4A TORTONIAN 90 89.8 ARTINSKIAN 1000 1000
10 5 TURONIAN lian
C5 93.9 290.1
MIOCENE
11.63 CENOMANIAN SAKMARIAN 295.0

STENIAN
SERRAVALLIAN 34 C34 ASSELIAN 298.9
5A 100 100.5 300 GZHELIAN 1200
PENNSYL-
C5A 303.7
CARBONIFEROUS
13.82 LATE
VANIAN
KASIMOVIAN 307.0 1250 MESOPRO-
15 5B C5B LANGHIAN ALBIAN MOSCOVIAN ECTASIAN
MIDDLE TEROZOIC
5C C5C
15.97 110 315.2
1400
5D C5D ~113 320 EARLY BASHKIRIAN
5E C5E BURDIGALIAN 323.2
6 C6 LATE SERPUKHOVIAN 1500 CALYMMIAN
120 APTIAN 330.9
SIPPIAN
20
MISSIS-
20.44 EARLY 1600
6A C6A
M0r
6B C6B AQUITANIAN M1 ~125 340 MIDDLE VISEAN
23.03 M3 BARREMIAN STATHERIAN
6C C6C 130 M5 ~129.4 346.7 1750
HAUTERIVIAN EARLY TOURNAISIAN
OLIGOCENE
7 C7 ~132.9 1800
25 7A C7A M10 358.9
8 C8 CHATTIAN VALANGINIAN
TERTIARY
M12 360
140 M14 ~139.8 FAMENNIAN OROSIRIAN
9 C9 M16
BERRIASIAN 2000
DEVONIAN
10 C10 27.82 M18
LATE PALEOPRO-
M20 ~145.0 ~372.2 2050
30 11 C11 TEROZOIC
150 M22 TITHONIAN FRASNIAN
12 380
C12
RUPELIAN LATE ~152.1 ~382.7 RHYACIAN
M25
KIMMERIDGIAN GIVETIAN
JURASSIC
MIDDLE ~387.7 2250
M29 ~157.3 EIFELIAN 2300
13 C13 33.9 160 ~393.3
OXFORDIAN
35 15 C15 ~163.5 400 EMSIAN SIDERIAN
CALLOVIAN
RAPID POLARITY CHANGES
16 C16 PRIABONIAN ~166.1

BATHONIAN EARLY ~407.6
17 MIDDLE ~168.3 PRAGIAN ~410.8 2500 2500
C17
37.8 170 BAJOCIAN ~170.3
PALEOGENE
AALENIAN LOCHKOVIAN
18 ~174.1 ~419.2
BARTONIAN 420 PRIDOLI
ORDOVICIAN SILURIAN
40 C18 ~423.0 NEOARCHEAN
19 LUDLOW LUDFORDIAN ~425.6
C19 41.2 180 TOARCIAN GORSTIAN
HOMERIAN
~427.4
WENLOCK ~430.5 2750
EOCENE
SHEINWOODIAN
20 ~182.7 ~433.4 2800
TELYCHIAN
LLANDO- ~438.5
C20 PLIENSBACHIAN 440 AERONIAN ~440.8
ARCHEAN
LUTETIAN EARLY VERY RHUDDANIAN
45 190 ~443.8
~190.8 HIRNANTIAN
~445.2
KATIAN MESO-
21
SINEMURIAN LATE ~453.0 3000
C21 47.8 SANDBIAN ARCHEAN
200 ~199.3 ~458.4
HETTANGIAN ~201.3 460
DARRIWILIAN
22 C22 MIDDLE ~467.3
50 RHAETIAN DAPINGIAN 3200
~470.0 3250
~208.5
TRIASSIC
23 C23
YPRESIAN 210 FLOIAN
EARLY ~477.7
480 TREMADOCIAN
24 PALEO-
LATE
~485.4
C24
NORIAN FURON- AGE 10 ARCHEAN
55 220 ~489.5
GIAN JIANGSHANIAN 3500
56.0 ~494
PAIBIAN
PALEOCENE
25 ~497
CAMBRIAN
C25
THANETIAN 500 GUZHANGIAN ~500.5 3600
26 ~227 Epoch 3 DRUMIAN
230 ~504.5
59.2 AGE 5
C26 ~509
60 SELANDIAN CARNIAN AGE 4
3750
~514
~237
Epoch 2
AGE 3 EOARCHEAN
27 61.6 520
C27 240 LADINIAN ~521
MIDDLE ~242
28 C28 AGE 2
DANIAN ANISIAN TERRE- ~529 4000 4000
29 247.2 NEUVIAN
65 C29 OLENEKIAN FORTUNIAN
66.0 250 EARLY 251.2 HADEAN
30 C30 INDUAN 251.90 540 541.0
Walker, J.D., Geissman, J.W., Bowring, S.A., and Babcock, L.E., compilers, 2018, Geologic Time Scale v. 5.0: Geological Society of America, https://doi.org/10.1130/2018.CTS005R3C. ©2018 The Geological Society of America
*The Pleistocene is divided into four ages, but only two are shown here. What is shown as Calabrian is actually three ages—Calabrian from 1.80 to 0.781 Ma, Middle from 0.781 to 0.126 Ma, and Late from 0.126 to 0.0117 Ma.
The Cenozoic, Mesozoic, and Paleozoic are the Eras of the Phanerozoic Eon. Names of units and age boundaries usually follow the Gradstein et al. (2012), Cohen et al. (2012) , and Cohen et al. (2013, updated) compilations. Numerical age estimates
and picks of boundaries usually follow the Cohen et al. (2013, updated) compilation. The numbered epochs and ages of the Cambrian are provisional. A “~” before a numerical age estimate typically indicates an associated error of ±0.4 to over 1.6 Ma.
REFERENCES CITED
Cohen, K.M., Finney, S., and Gibbard, P.L., 2012, International Chronostratigraphic Chart: International Commission on Stratigraphy, www.stratigraphy.org (accessed May 2012). (Chart reproduced for the 34th International Geological Congress, Brisbane,
Australia, 5–10 August 2012.)
Cohen, K.M., Finney, S.C., Gibbard, P.L., and Fan, J.-X., 2013, The ICS International Chronostratigraphic Chart: Episodes v. 36, no. 3, p. 199–204 (updated 2017, v. 2, http://www.stratigraphy.org/index.php/ics-chart-timescale; accessed May 2018).
Gradstein, F.M, Ogg, J.G., Schmitz, M.D., et al., 2012, The Geologic Time Scale 2012: Boston, USA, Elsevier, https://doi.org/10.1016/B978-0-444-59425-9.00004-4.
Previous versions of the time scale and previously published papers about the time scale and its evolution are posted to http://www.geosociety.org/timescale.
Fig. 3.3 Geologic Time Scale v 5.0. (Reprinted by permission from GSA, from Walker, J.D., Geissman, J.W., Bowring, S.A., and Babcock, L.E.,
compilers, 2018, Geologic Time Scale v. 5.0: Geological Society of America # 2018 The Geological Society of America)
were without human settlements or substantial land use, and surface of the Earth, but all life that inhabits it and all systems
most (45%) of the rest of the planet was classified as that support it.
“semiwild,” having only limited use as areas for settlement These changes alter the fundamental way we look at
and agriculture (Ellis et al. 2010). But as environmental geographical ecology, as well as how we attempt to succeed
scientist Erle Ellis and his colleagues report, “By 2000, the at conservation. Given the global effects of human settlement
opposite was true, with the majority of the biosphere in and activity, Ellis and others have developed a new system of
agricultural and settled anthromes [biomes influenced or biome classification consisting of five human-dominated or
modified by human use or residence], less than 20% seminat- human-influenced biomes, or “anthromes,” and a sixth cate-
ural and only a quarter left wild. . .At present, and ever more gory, wildlands, comprising everything else (Ellis and
in the future, the form and process of terrestrial ecosystems in Ramankutty 2008) (Fig. 3.6). As Ellis and colleague Navin
most biomes will be predominantly anthropogenic, the prod- Ramankutty have noted, “. . .unlike classic biomes, which
uct of land use and other direct human interactions with attempt to represent fairly homogeneous forms of vegetation,
ecosystems” (Ellis et al. 2010:589). Today all major ecosys- anthromes represent complex mixtures of different land uses
tem drivers (habitat change, climate change, invasive species, and land covers that are far harder to characterize in simple
over-exploitation of species, and pollution) are driven by terms” (Ellis and Ramankutty 2008). This approach is trans-
human influence (Fig. 3.5, Millennium Ecosystem Assess- formationally radical in the way we would view the Earth’s
ment 2005), and these influences have changed not only the biological communities. It represents “an alternate view of
Fig. 3.4 Basic representation of

major world biomes as a function
of temperature and moisture.
(Reprinted under CC0
1.0 Universal Public Domain)
the terrestrial biosphere, based on an empirical analysis of and crops, but was used far more intensively, leading to more
global patterns of sustained direct human interaction with rapid ecological change and loss of biodiversity (Ellis et al.
ecosystems, yielding a global map of ‘anthropogenic 2010).
biomes’” (Ellis and Ramankutty 2008:439). Ellis and To better understand these changes sequentially and
Ramankutty hypothesize that anthropogenic biomes will dif- conceptually, follow the flow chart shown in Fig. 3.8 from
fer substantially from “natural” biomes in basic ecosystem top to bottom. A growing human population intensifies
processes and biodiversity and that these differences “will be resource use and diversifies in human enterprises that require
at least as great as those between the conventional biomes resource use. Land is transformed and its biotic components
when observed using equivalent methods at the same spatial are changed in the process. Biogeochemical cycles begin to
scale” (Ellis and Ramankutty 2008:445). They further be altered, first locally, then regionally, and finally globally.
hypothesize that “these differences will be driven by Together, changes in these cycles and modification and inten-
differences in population density and land use between the sification of land use create changes in global climate
biomes. . ., a trend already evident in the general tendency (Chap. 4) and loss of biodiversity (Chap. 2).
toward increasing cropped area, irrigation, and rice produc- The new anthrome classification system is neither perfect
tion with increasing population density” (Ellis and nor complete. It does not replace the traditional biome
Ramankutty 2008:445). Finally, they believe that anthropo- classifications appropriate to natural systems still present
genic biomes will increasingly depict global patterns of eco- around the world, but its designers advocate for more inten-
system processes and biodiversity over time as human tional and in-depth studies of these relatively new anthropo-
influence on ecosystems increases (Ellis and Ramankutty genic biomes to “encourage a richer view of human–
2008:445). ecosystem interactions across the terrestrial biosphere, and
Over the eighteenth and nineteenth centuries this shift was that this will, in turn, guide our investigation, understanding,
relatively slow and gradual, but accelerated in the twentieth and management of ecosystem processes and their changes at
century (Fig. 3.7, Table 3.2) such that “rapid intensification global and regional scales” (Ellis and Ramankutty 2008:446).
of land use in the twentieth century finally pushed the bio- Human effects on the Earth have not only changed the distri-
sphere into its present anthropogenic state” (Ellis et al. bution of traditional regional associations of plants and
2010:602). The primary drivers of change were a sixfold animals, they have created associations of plants, animals,
increase in the amount of pasturelands with accompanying and biogeochemical processes never encountered before.
expansion of croplands into what were formerly wild grass- Each one of these new creations belong to an entirely differ-
land, shrublands and woodlands (Ellis et al. 2010). Changes ent category for conservation. That is the category of the
occurred not only because more land was used for livestock novel ecosystem.
Fig. 3.5 Principal drivers of change in the Earth’s biodiversity and indicate continuation of the current level of impact; diagonal and vertical
ecosystems. Cell color indicates impact of each driver on biodiversity in arrows indicate progressively increasing trends in impact. (Reprinted by
each type of ecosystem over the past 50–100 years. “High impact” permission from the World Resources Institute, from Millennium Eco-
means the driver has significantly altered biodiversity in that biome; system Assessment, 2005. Ecosystems and human well-being: synthe-
“low impact” indicates that it has had little influence on biodiversity in sis. Island Press, Washington, DC.# 2005 World Resources Institute)
the biome. Arrows indicate trend in the driver. Horizontal arrows
Fig. 3.6 Anthropogenic biomes: world map and regional areas. Biomes and Ramankutty, N., 2008. Putting people in the map: anthropogenic
are arranged into groups (see Table 3.2) and sorted in order of human biomes of the world. Frontiers in Ecology and the Environment
population density. (Reprinted by permission of Wiley, from Ellis, E.C., 6, 439–447. # 2008 The Ecological Society of America)
ecosystem from (formerly) treeless tundra to a new and

3.2 Understanding and Managing the Novel formerly non-existent system that the authors described as
Ecosystem “grassland-steppe.” The authors’ grasped, if perhaps imper-
fectly, the significance of their findings in the words of their
3.2.1 Origins of Novel Ecological Associations conclusions, “Perhaps the most unexpected result of our
simulations is the conclusion that climatic warming, in the
The first use of the term “novel ecosystem” in peer-reviewed absence of an increase in precipitation, will lead to the wide-
scientific literature is generally agreed to be found in the spread distribution of grassland-steppe or other novel
landmark paper, “Time lags and novel ecosystems in drought-resistant vegetation type that is currently rare or
response to climate change in Arctic Alaska,” by F. Stuart absent in the landscape” (Chapin and Starfield
Chapin III and Anthony M. Starfield, published in 1997 in the 1997:457–458).
journal, Climate Change (Chapin and Starfield 1997). In this The Chapin and Starfield study proved to be only the first
paper, Chapin and Starfield described an ecological simula- of many that documented the change in ecosystem types and
tion model that they created to predict the future progress of properties, not from one traditional ecosystem category to
an already ongoing ecological process – the invasion of the another, but from an historic “natural” ecosystem to one with
Arctic tundra by trees (especially aspen, Populus spp.) as a unique characteristics not previously encountered. The phe-
result of climate warming. This invasion was not simply a nomenon became so frequent and widespread in ecological
change in local species composition, but a change in studies, and resulting publications of them, that it naturally
3.2 Understanding and Managing the Novel Ecosystem 89
Fig. 3.7 Conceptual model and characteristics of contemporary anthro- primary production (NPP). (Reprinted by permission of Wiley, from
pogenic biomes (“anthromes”) in relation to (a) changes in human Ellis, E.C., and Ramankutty, N., 2008. Putting people in the map:
population density and land use; which form patterns of (b) ecosystem anthropogenic biomes of the world. Frontiers in Ecology and the Envi-
structure, processes, and biodiversity; which characteristics structures ronment 6, 439–447. # 2008 The Ecological Society of America)
and levels of (c) human population density, land use and cover, and net
prompted the development of new terms and definitions. ecosystems. . .have species compositions and relative
Ecologist Richard Hobbs and his colleagues offered an abundances that have not occurred previously within a
early definition and criteria for novel ecosystems. “Novel given biome. The key characteristics are (1) novelty: new
Table 3.2 Descriptions of anthropogenic biomes (anthromes)

Group Biome Description
Dense settlements Dense settlements with substantial urban area
11 Urban Dense built environments with very high populations
12 Dense settlements Dense mix of rural and urban populations, including both suburbs and villages
Villages Dense agricultural settlements
21 Rice villages Villages dominated by paddy rice
22 Irrigated villages Villages dominated by irrigated crops
23 Cropped and pastoral villages Villages with a mix of crops and pasture
24 Pastoral villages Villages dominated by rangeland
25 Rainfed villages Villages dominated by rainfed agriculture
26 Rainfed mosaic villages Villages with a mix of trees and crops
Croplands Annual crops mixed with other land uses and land covers
31 Residential irrigated cropland Irrigated cropland with substantial human populations
32 Residential rainfed mosaic Mix of trees and rainfed cropland with substantial human populations
33 Populated irrigated cropland Irrigated cropland with minor human populations
34 Populated rainfed cropland Rainfed cropland with minor human populations
35 Remote croplands Cropland with inconsequential human populations
Rangeland Livestock grazing; minimal crops and forests
41 Residential rangelands Rangelands with substantial human populations
42 Populated rangelands Rangelands with minor human populations
43 Remote rangelands Rangelands with inconsequential human populations
Forested Forests with human populations and agriculture
51 Populated forests Forests with minor human populations
52 Remote forests Forests with inconsequential human populations
Wildlands Land without human populations or agriculture
61 Wild forests High tree cover, mostly boreal and tropical forests
62 Sparse trees Low tree cover, mostly cold and arid lands
63 Barren No tree cover, mostly deserts and frozen land
Reprinted by permission of Wiley, from Ellis, E.C., and Ramankutty, N., 2008. Putting people in the map: anthropogenic biomes of the world.
Frontiers in Ecology and the Environment 6, 439–447. # 2008 The Ecological Society of America
species combinations, with the potential for changes in eco- combinations of plant and animal species not previously
system function; and (2) human agency: ecosystems that are encountered in natural ecosystems (Hobbs et al. 2006:2).
the result of deliberate or inadvertent human action, but do The previously described tundra – to – grassland steppe
not depend on continued human response or human interven- transition modeled by Chapin and Starfield was initially
tion for their maintenance” (Hobbs et al. 2006:2). More altered by climate change (a change in environmental
directly, we might say that novel ecosystems are “new conditions) which permitted the entry of new species that
combinations of species under new abiotic conditions” formerly could not cope with tundra conditions. Once
(Seastadt et al. 2008:547). Novel ecosystems are generated established, such new species changed conditions further
by ecological processes affected by human activity and inter- and made it difficult for traditional tundra species to
vention. They arise primarily from the degradation of wild re-enter the system.
systems (to some form of intensive human use, like agricul- Morse et al. (2014) subsequently reframed the definition
ture) or subsequent abandonment of intensively used systems of a novel ecosystem as “a unique assemblage of biota and
(like a cropland) (Fig. 3.9). environmental conditions that is the direct result of inten-
Hobbs and others offer a more detailed definition of novel tional or unintentional alteration by humans (i.e., human
ecosystems as “a system of abiotic, biotic and social agency) sufficient to cross an ecological threshold that
components (and their interactions) that, by virtue of human facilitates a new ecosystem trajectory and inhibits its return
influence, differ from those that prevailed historically, having to a previous trajectory regardless of additional human inter-
a tendency to self-organize and manifest novel qualities with- vention. The resulting ecosystem must also be self-sustaining
out intensive human management” (Hobbs et al. 2013:58). in terms of species composition, structure, biogeochemistry,
Novel ecosystems result from “biotic response to human- and ecosystem services. A defining characteristic of a novel
induced abiotic conditions and/or novel biotic elements ecosystem is a change in species composition relative to
(e.g., land degradation, enrichment of soil fertility, introduc- ecosystems present in the same biome prior to crossing a
tion of invasive species)” and, as a result, produce new threshold” (Morse et al. 2014:1).
Fig. 3.8 A conceptual model

illustrating humanity’s direct and
indirect effects on the Earth
system. (Reprinted by permission
from the American Association
for the Advancement of Science,
from Vitousek, P.M., Mooney,
H.A., Lubchenco, J., and Melillo,
J.M., 1997. Human Domination
of Earth’s Ecosystems. Science
277, 494–499. # 1997 AAAS)
Novel ecosystems are formed in response to environmen-

tal stress, either from changes in abiotic factors (which is
? ?
common in ecosystem degradation) or through colonization
Novel by new species (which may occur when an intensively used
ecosystems system is abandoned), which may make it difficult for native
‘Wild’ Intensive
species to re-enter the abandoned system. Although the new
agriculture ecosystem may return to its historic norms under some
conditions, it is now vulnerable, in its altered state, to internal
restructuring of processes and species interaction which can
Degradation Abandonment lead to an entirely new system.
Invasion The fact that definitions in science inevitably get longer
does not guarantee that they also inevitably get better, but the
Fig. 3.9 Novel ecosystems arise from degradation and invasion
of “wild” or natural systems or from abandonment of intensively Morse et al. definition does improve the precision of the
managed systems for uses such as agriculture. (Reprinted by permission concept of a novel ecosystem, and provides some test criteria
from Wiley, from Hobbs, R.J., Arico, S., Aronson, J., Baron, J.S., for determining if a given system is indeed ecologically
Bridgewater, P., Cramer, V.A., Epstein, P.R., Ewel, J.J., Klink, C.A., novel. Specifically, their definition is in part dependent on
Lugo, A.E., Norton, D., Ojima, D., Richardson, D.M., Sanderson, E.W.,
Valladares, F., Vilà, M., Zamora, R., Zobel, M., 2006. Novel origin. Was the ecosystem a product of alteration of the
ecosystems: theoretical and management aspects of the new ecological environment by humans? A second criterion is whether or
world order. Global Ecology and Biogeography 15, 1–7. not the system has crossed an “ecological threshold.” Now,
# 2006 Blackwell Publishing Ltd) what does that mean? As Morse et al. note, a threshold can be
inferred if the ecosystem sustains itself in its new post-
threshold state without human assistance (Hobbs et al. 2009, degradation (degrading the ecological features of a landscape
Morse et al. 2014) and “positive feedbacks inherent in a in order to create human benefit, such as converting the
novel ecosystem are one means by which a threshold can complexity of species in a native prairie to the monoculture
prevent the return to a previous state. . .To verify that an of a corn field) (Arrow 2). Humans may also generate a novel
ecosystem has crossed a threshold, a measurable difference ecosystem by manipulating the natural system to maximize
must exist between the ecosystem’s previous and current human gains from ecosystem services in a designed system,
state. We propose that the occurrence of ecological such as damming a river to generate more water for irrigation
parameters outside their historical ranges of variabili- in a local area than the flowing river would have provided
ty. . .could be indicative of a threshold crossing. . .” (Morse (Arrow 3). With further manipulation of an already impacted
et al. 2014:2–3). That is, a novel ecosystem is one that has system, humans may create a designed system which
changed enough that it will not return to the former “natural” remediates a degraded system to provide ecosystem services
state without human intervention. A third criterion is species that had been lost, such as planting trees on a vacated oil
composition. Is the system’s species composition a unique, drilling pad in a forest to restore wildlife habitat and mitigate
“we’ve never seen this before” arrangement of species par- climate warming (Arrow 4). Finally, the altered system may
ticular to the new system? Finally, is the new system self- transition to a natural state (Arrow 5), either because the
sustaining? Will it remain and persist in its present condition system still maintains the capacity to do so (no “thresholds”
without (or in spite of) further human intervention? To judge were crossed) or because humans actively restore the site
whether this criterion has been met, longer time spans through additional effort, such as planting grasses and
(at least 10–100 years) are necessary to determine if the wildflowers in an abandoned corn field to re-create a prairie
system is persistent in the long term, not only in species ecosystem (Morse et al. 2014).
composition (which changes over time in all ecosystems),
but in the ecological processes that characterize it and the
environmental conditions they influence (Morse et al. 2014). 3.2.2 Are Novel Ecosystems Good or Bad?
Human action can create novel ecosystems in four differ-
ent ways (Fig. 3.10). Perhaps the most common is uninten- It is a near-instinctual reaction, especially among traditional
tional alteration (humans engaged in activities of which they ecologists and conservationists, to immediately attach a neg-
did not understand the environmental effects) or intentional ative connotation to a term like “novel ecosystem,” as well as
Fig. 3.10 Role of human agency in creation and maintenance of altered maintenance. Arrows 7 and 8 designate pathways from designed
ecosystems. Arrow 1 designates natural ecosystem processes. Arrow (restored) ecosystems (Arrow 7) or impacted ecosystems (Arrow 8) to
2 represents unintentional alteration or intentional degradation by that of a novel ecosystem. (Reprinted by permission from N. Morse,
humans. Arrow 3 depicts alteration of ecosystem services for human from Morse, N.B., Pellissier, P.A., Cianciola, E.N., Brereton, R.L.,
benefit. Arrow 4 indicates environmental remediation (such as restora- Sullivan, M.M., Shonka, N.K., Wheeler, T.B., McDowell, W.H.,
tion of ecosystem services without actual restoration of the original 2014. Novel ecosystems in the Anthropocene: a revision of the novel
ecosystem). Arrow 5 represents full ecosystem restoration (processes, ecosystem concept for pragmatic applications. Ecology and Society 19
structure, and components). Arrow 6 shows degradation of a designed (2):12. # The Authors)
(restored) ecosystem to an impacted ecosystem state through lack of
to the reality it describes. In fact, it sounds rather like “inva- even after dramatic losses of native species diversity if simple
sive species” at the ecosystem level. And, indeed, the two functional roles are provided by introduced species”
may be closely connected, with novel ecosystems often (Mascaro et al. 2012:221). The increase in species richness
formed by invasion and spread of nonnative plants and was the result of a greater richness of introduced tree species
animals. Further, the novel ecosystem lacks a “history,” or as well as a lack of significant decline in richness of native
at least one distinguished by any natural or evolutionary tree species. Novel and native forests had about the same
pedigree. We are drawn, both as tourists and as scientists, levels of aboveground litterfall – a key factor in subsequent
to systems of extended lineage. If we come to a fork on a levels of organic matter in forest soils – as well as above-
hiking trail where one arrow points to something called the ground biomass (Mascaro et al. 2012).
“restored forest” and the other to the “old growth forest,” The authors found support for their hypothesis that “eco-
what true nature lover would not spurn the first and head for system function in novel forests would meet or exceed levels
the second? found in native forests in terms of aboveground biomass and
There is warrant to what might appear at first to be mere productivity. . ., nutrient turnover and below ground carbon
prejudice. For one of us (Van Dyke) who has worked in storage. . .All significant changes in ecosystem functional
efforts associated with tallgrass prairie restoration, there is properties increased with the increase in tree species richness
the real experience that even an intense and well-crafted and diversity in novel forests” (Mascaro et al.
restoration effort may, over its first 3–5 years, contain per- 2012:229–230). The work of Mascaro et al. not only suggests
haps 60–80 plant species. The Konza Prairie in Kansas that novel ecosystems will proliferate, but that many will
(USA), one of the last natural and extensive (3487 ha) perform their ecological functions at least as well as the
tallgrass prairies in the United States, contains over 600 spe- traditional systems they might replace. If such ecosystems
cies, an order of magnitude greater. But there is also need to can perform ecological functions as well as or better than
put natural prejudice aside, recognize that novel ecosystems native systems, they should also be as amenable to principles
and restored ecosystems are not necessarily the same, and and practices of ecosystem management (Chap. 9) as those
examine how novel systems might actually function in native systems, but might require different techniques and
providing ecosystem services and contributing to approaches. If this might be true, what methods would be
biodiversity. employed in the management of novel ecosystems, and to
Recall from our examination of biodiversity in the previ- what ends would we direct them?
ous chapter that a greater diversity of species with different
sensitivities to a range of environmental conditions usually
leads to greater stability of ecosystem properties and a greater
If novel ecosystems can sometimes perform and pro-
number of functional ecosystem processes (Tilman et al.
vide ecosystem services as well or better than native
1997). Biologist Joseph Mascaro and his colleagues put this
ecosystems, should managers intentionally remove
assumption to the test in their examination of ecosystem
native ecosystems and establish novel ones to enhance
function and species diversity in novel forests on Hawai’i
benefits to humans?
Island, part of the US state of Hawaii, a group of volcanic
islands in the Pacific Ocean. “Hawaii,” noted Mascaro et al.,
“. . .has much higher regional plant richness that it did histor- 3.2.3 Managing Anthropogenic Biomes
ically, due to >1000 plant species introductions and only ~71 and Novel Ecosystems
known plant extinctions, resulting in 100% increase in rich-
ness” (Mascaro et al. 2012:221). As such, Hawaii is an ideal Near the city of Boulder (estimated human population of
incubator for the breeding of novel ecosystems, and many of 104,000 in 2019) in the US state of Colorado, gravel had
these manifest themselves as novel associations of non-native been mined from a 100 ha bottomland site originally covered
forest trees. Examining these, Mascaro et al. found that by tallgrass prairie, a natural but now rare ecosystem through-
“novel forests had significantly higher tree species richness out the United States. After gravel mining ceased in the early
and higher diversity of dominant tree species. We further 1990s, native grasses were seeded onto the site in 1998.
found that above ground biomass, productivity, nutrient turn- Following higher than normal precipitation in the seeding
over (as measured by soil-available and litter-cycled nitrogen year, the site experienced three consecutive years of below
and phosphorus) and below ground carbon storage either did normal precipitation (69–90% of average), an increasingly
not differ significantly or were significantly greater in common manifestation of anthropogenic climate change in
novel forests relative to native ones. We found that the the western and intermountain US. When gravel mining was
addition of introduced N2-fixing tree species on N-limited initiated, soils removed from the site were stockpiled with the
substrates had the strongest effect on ecosystem function, a intention of replacing them when mining was completed.
pattern found by previous empirical tests. Our However, soil stockpiled in this way, especially for long
results. . .suggest basic ecosystem processes will continue periods of time, will be different in chemistry and structure
when it is placed back on the site compared to what it was Nevertheless, some general principles may be (cautiously)
when removed. In this case, the “returned” soil was lower in drawn from this example. The first is the application of an
organic matter, moisture, and organic nitrogen than that adaptive management approach (Chap. 9), a key feature of
characteristic of a typical tallgrass prairie and more like sound ecosystem management generally and management of
soils associated with arid steppes (Seastedt et al. 2008). novel ecosystems particularly. In adaptive management,
Seeds sown on the restored site were all native species, but management actions are designed as experiments so that the
managers intentionally added some species with wider envi- results of the action will effectively inform subsequent man-
ronmental tolerances and greater drought resistance than agement actions. Although this case history lacked compari-
species uniquely associated with tallgrass prairie. The plant son or control sites to see what would have happened if the
community that ultimately emerged consisted of more seed mixture had been restricted to only native tallgrass
drought-resistant species, and particularly dominated by one prairie plants, its concurrent monitoring of species emergence
warm-season grass, the salt sacaton (Sporobulus airoides) and coverage in the face of environmental (weather)
(Seastedt et al. 2008) (Fig. 3.11). Also known as alkali conditions and its comparison of the emerging plant commu-
sacaton, this grass’s first name reflects its ability to thrive in nity to basic knowledge of historic tallgrass prairie
dry, saline (salt-laden) soils, and the second its ability to communities provided insight and understanding as to why
tolerate a wide range of acidic conditions (pH) in soil, includ- a different kind of plant community was ultimately successful
ing high (alkaline) pH levels. Salt sacaton and other warm on this site. Additional monitoring that demonstrated that the
season grasses more typical of more arid mixed-grass and new community was largely devoid of invasive species was
shortgrass steppes, rather than tallgrass prairie, eventually also a key finding, suggesting that this novel ecosystem
contributed over 90% of the site’s plant coverage, and largely might meet the “stability” criterion for a genuine novel eco-
resisted invasions by other species, both native and system as proposed by Morse et al. (2014) and persist in its
non-native (Cherwin et al. 2009). As Seastedt et al. noted in alternative stable state, although no plant community is per-
retrospect, “This successful revegetation effort resulted from manent, and all change in response to variation in climate
the selection of a seed source containing representatives of (e.g., Huntley and Webb 1989). Also contributing to this
short-, mixed-, and tallgrass species. . .The diverse seed mix successful restoration effort was the recognition by managers
used by the revegetation team interacted with climate that traditional management approaches would not be likely
conditions and unusual soil characteristics to select the com- to achieve management goals (Seastedt et al. 2008). The
munity type and produced an impressive, albeit novel, grass- authors themselves recognized these limitations, but also
land community” (Seastedt et al. 2008:551). appreciated key elements that could contribute to success in
This is but one case history of managing a novel ecosys- managing other novel systems. “A search for general rules
tem, in this instance one that emerged in a restoration effort. that can be used to manage novel ecosystems,” they noted, “is
Fig. 3.11 What was originally a

tallgrass prairie in northern
Colorado (USA) until the 1950s
was excavated to extract gravel
through the 1990s. Its restoration
was achieved, not by replicating
the original species composition
of tallgrass prairie plants, but by
adding grass species more
resistant to increasingly dry
conditions associated with climate
change, resulting in a novel
ecosystem. (Reprinted by
permission from Wiley,
from Seastedt, T.R., Hobbs, R.J.,
Suding, K.N., 2008. Management
of novel ecosystems: are novel
approaches required? Frontiers in
Ecology and the Environment
6, 547–553. # 2008 The
Ecological Society of America)
3.3 The Ecology of Non-native and Invasive Species 95
likely to be a long and possibly unproductive exercise. A 2008:446). Classification of anthropogenic biomes is not
logical approach would be to maximize genetic, species, and intended and should not be used to replace existing biome
functional diversity wherever possible, to increase the viabil- systems whose categories are based on real differences in
ity of communities and ecosystems under certain climate climate, terrain, and geology, but an anthropogenic biome
regimes. Monitoring responses to any action, or lack of action, system can encourage a richer view of human–ecosystem
remains the key activity. . . management actions produced interactions throughout the world, because such understand-
what were perceived as desirable outcomes by manipulating ing and interaction can “guide our investigation, understand-
mechanisms that enhanced desirable system components, ing, and management of ecosystem processes and their
rather than by removing or suppressing undesirable changes at global and regional scales” (Ellis and Ramankutty
species. . .attention to a rigorous ‘experimental’ design, 2008:446).
including reference areas whenever possible, is [also] appro- Today 40% of all ice-free land is used for agriculture or in
priate if not essential to a defensible, informative, and publicly urban environments. Almost as much (37%) of ice-free land
acceptable management program” (Seastedt et al. 2008:552). not used for these purposes is embedded within anthromes
Management is more than method. If effective, it is driven having these uses. It is the ecology of these ‘unused lands’
first by clearly defined objectives. The growing prevalence of that now deserves the attention of conservationists to main-
novel ecosystems requires conservationists to change their tain, enhance and restore ecological functions in these
thinking about what might be worth conserving. Again remnants, because anthropogenic landscapes have become
Seastedt et al. offer insight, affirming that conservation man- the most abundant terrestrial ecosystems on Earth (Hobbs
agement of novel ecosystems “must explicitly acknowledge et al. 2006). The novel ecosystems they contain must also
the current status and predict human conditions of these be targets of conservation. What do we know, and what do
systems. Old styles of management, which focused on we have to learn, about the ecology of species that help create
removing undesirable species or conditions from ecosystems novel ecosystems, their patterns of movement, and their
to return them to a prior condition, may no longer be suffi- mechanisms of invasion?
cient. We need to consider, and experiment with, novel
approaches to ecosystem management that focus on desired
outcomes or trajectories, rather than simply taking preventa- 3.3 The Ecology of Non-native and Invasive
tive or therapeutic measures” (Seastedt et al. 2008:547). Species
Our recognition of the need for intentional management of
novel ecosystems must also translate upward to the level of 3.3.1 How Do Invasive Species Affect Existing
managing anthromes. Specialists on anthrome characteristics, Ecosystems and Create New Ones?
like Ellis and Ramankutty (2008), declare that conserva-
tionists cannot ignore the importance of understanding The letter “I” in “HIPPO,” the “acronym of extinction” that
interactions between managed and natural systems. Avoid- helps conservationists recite the five great threats to biodiver-
ance and ignorance of these relationships is no longer an sity (Chap. 2), stands for “invasive species,” the second most
option. Instead, “anthropogenic biomes offer a framework commonly listed factor contributing to endangerment after
for incorporating humans directly into global ecosystem habitat loss and degradation. More particularly, invasive spe-
models” (Ellis and Ramankutty 2008), a framework that has cies are documented to be the second leading cause of extinc-
been increasingly used for more complete and complex tion in the United States and cause damages in excess of US
analyses of global vegetation change over the same periods $120 million (Crowl et al. 2008). The spread of non-native
(e.g., Hurtt et al. 2011). species from one part of the world to another by human
Such incorporation requires change in perspective. Ellis beings, sometimes intentionally and oft-times unintention-
and Ramankutty, among others, are calling for ecologists and ally, is no longer just another song of lament for the conser-
conservationists to “come home” and invest their effort vation community to sing with sorrow and regret. It must
where humans live and are most directly affecting the Earth now be, and increasingly is, understood as an important
and its biodiversity. As they put it bluntly, “Ecologists have manifestation of the Anthropocene Epoch, the role of humans
long been known as the scientists who travel to uninhabited as a primary, and often destructive, agent of disruption of
lands to do their work. As a result, our understanding of biological communities through our transport of species to
anthropogenic ecosystems remains poor when compared new areas thousands of miles from their native environments.
with the rich literature on “natural” ecosystems. . .Building Although anthropogenically-induced climatological, geolog-
ecological science and education on a foundation of anthro- ical, and land use factors contribute to the formation of novel
pogenic biomes will help scientists and society take owner- ecosystems, one of the things that makes them “novel” is that
ship of a biosphere that we have already altered irreversibly, they are composed of new, previously unencountered
and moves us toward understanding how best to manage the arrangements of organisms. And the most important cause
anthropogenic biosphere we live in” (Ellis and Ramankutty of this effect is the spread of invasive species.
The first major work on invasive species biology, The

Ecology of Invasions by Plants and Animals, was published
by British ecologist Charles Elton in 1958. By this point in
his distinguished career, Elton had realized the significance
of biological invasions and the dangers of scientific igno-
rance. “We must,” he said, “make no mistake: we are seeing
one of the great historical convulsions in the world’s fauna
and flora” (Elton 1958:32). International commerce has aided
and increased the movement of species globally and across
taxonomic groups. Only a fraction of transported species
become established, and only about 1% become pests, but
additions and effects of all non-native species, including
invasive pest species, are cumulative over time. Today there
are as many alien established plant species in New Zealand as
native species. The US states of Arizona and Montana, which
historically had no fish species in common, now share
33 (Mooney and Cleland 2001). These changes in native Fig. 3.12 A typical sequence of invasion for most non-native species
biota have occurred because humans transport species for begins with movement via a transport mechanism from the species’
human benefit, and invasive species use human means of native range to a novel environment. Upon arrival the new species
may (1) not survive, (2) persist but fail to spread, or (3) begin to
transport for their benefit. A typical invasion sequence
reproduce and expand their local distribution. If the population has
follows a pattern of (1) transport from native range to a sufficient resources and insufficient mortality for competitors, predators
novel environment, (2) local establishment in the new envi- or parasites to control its numbers, it may begin to expand its distribution
ronment (which often takes many years and multiple regionally and continentally, resulting in multiple, discrete, and
expanding populations. (Reprinted by permission from Wiley,
“invasions”) and (3) range expansion leading to establish-
from Crowl, T.A., Crist, T.O., Parmenter, R.R., Belovsky, G., Lugo,
ment and naturalization at regional or continental scales, at A.E., 2008. The spread of invasive species and infectious disease as
which point there is no hope of eradicating the invasive drivers of ecosystem change. Frontiers in Ecology and the Environment
species (Fig. 3.12). 6, 238–246. # The Ecological Society of America)
Invasive species do not simply create change by reducing
or replacing populations of native species. They can also
change ecosystems and ecological structure. The invasion
of the zebra mussel (Dreissena polymorpha) (Fig. 3.13)
into the North American Great Lakes and its subsequent
spread to major river systems of the US Midwest altered
abiotic factors including water transparency, nutrient cycling,
and benthic habitat structure, as well as biotic factors of food-
web structure, bioaccumulation of contaminants and the
diversity of native freshwater mussels (Strayer et al. 1999;
Crowl et al. 2008). The zebra mussel also provided introduc-
tion of its own roundworm parasite (Bucephalus
polymorphus), which caused reductions in populations of
cyprinid freshwater fish, the round worm’s intermediate Fig. 3.13 The zebra mussel (Dreissena polymorpha), a native of the
host (Crowl et al. 2008). Cheatgrass (Bromus tectorum), a Black and Caspian Seas of Eurasia, which has now expanded its range
native of Eurasia and the Mediterranean Basin, has become throughout North America. Zebra mussels can attach to any hard sub-
strate, and to other living creatures, in the latter capable of causing direct
resident in all 50 US states and established large populations
mortality. An exceptionally efficient filter-feeder, the zebra mussel can
in arid and semi-arid shrublands and grasslands of the US alter ecosystem structure by consuming large amounts of phytoplankton,
Intermountain West. An annual rather than perennial grass, often the most important component of primary production in many
long-term accumulation of dead cheatgrass shortens the Fire aquatic systems. (Photo courtesy of C. Rees/USGS, public domain)
Return Interval (FRI), promoting more frequent fire events,
which favors further invasion, exclusion of native plants, and Kowarik offers a precise way of evaluating such damage.
loss of carbon (C) to the atmosphere (D’Antonio and Damage from an alien species can be said to have occurred
Vitousek 1992; Young and Allen 1997; Crowl et al. 2008). when there is “a significant adverse effect on a biotic or
Although native species are often described as “damag- abiotic conservation resource that has an impact on (a) the
ing” to the native environment, exactly what we mean by value of the conservation resource, (b) the conservation
“damage” should be carefully defined. Ecologist Ivan resource as an ecosystem component, or (c) the sustainable
use of the conservation resource” (Bartz et al. 2010, cited in negative impacts on other species or resources with special
Kowarik 2011:1980). Effects of novel species should not be conservation interest” (Kowarik 2011:1980) (Information
called detrimental “when it is not associated with significant Box 3.1).
Information Box 3.1: Beavers in Tierra del Fuego Information Box 3.1 (continued)
Beaver (Castor canadensis) (Information Box Fig. 3.1), Anderson et al. found that “Ecosystem engineering
a rodent native to North America but introduced to many by invasive beavers transformed between 20 and 43%
parts of the world, are what we call an “ecosystem of the studied hydrological networks [connected
engineer” due to their practice of building dams across streams and ponds]. . .beavers significantly reduced
flowing streams to create upstream impoundments substrate diversity compared with flowing streams,
(“beaver ponds”). Christopher Anderson and his but displayed very similar parameters to the natural
colleagues chose to study the effects of beavers in one conditions recorded for peat bogs and lakes. . .”
of their non-native habitats, the islands comprising both (Anderson et al. 2014:217). Impoundments created by
the Argentinian and Chilean portions of the Tierra del beavers caused BOM [benthic organic matter] standing
Fuego Archipelago at the southern tip of South America. crop to significantly increase, and benthic
Rather than investigate direct effects of competition with macroinvertebrate diversity and richness to decrease,
other species, Anderson et al. focused on “the indirect compared with natural stream habitats.
and broader-scale impacts of invasive species [beaver] Overall, Anderson et al. noted, “At the landscape
on ecosystems and landscapes” (Anderson et al. level, we did not find a difference in benthic diversity
2014:215). Specifically, Anderson and his colleagues or richness in landscape scenarios that included only
compared patch- and landscape-level effects of invasive natural and those that had both natural patches and
beavers on biodiversity and ecosystem functions in beaver-impacted areas. . .In contrast. . .the beaver’s
freshwater streams and ponds. Their hypothesis was influence on the ecosystem function associated with
that if patch-level habitat conditions of beaver streams carbon standing crop increased by an average of
affected by beaver were the same as in natural streams, 71.6% +/ 17.1% (ranging from 43.6% to 111.3%)
beavers would not alter stream biodiversity at the land- in the studied watersheds” (Anderson et al. 2014:218).
scape scale. They also hypothesized that beaver influ- Anderson et al. concluded “. . .while they [beaver]
ence on broader ecological processes, such as carbon reduced the diversity of benthos as compared to that
retention and standing biomass crop, would be large and found in natural lotic [stream] patches, they only
reflective of overall magnitude and extent of the generated biotic assemblages in their engineered lentic
beaver’s engineered ecosystems (ponds formed by [pond] patches that were very similar to those found in
building dams in streams) in the landscape (Anderson natural lentic habitats. . .invasive beavers ultimately
et al. 2014). had no measurable influence on stream landscape-
level biodiversity, but they did change the ecosystem
function related to carbon cycling. . .enhancing second-
ary production of benthos by an order of
magnitude. . .and transitioning benthic production and
BOM decomposition rates in sub-Antarctic streams to
values more similar to temperate zones. . .” (Anderson
et al. 2014:219–220). So, even such an intentional
ecosystem engineer as the beaver does not necessarily
degrade the system to which it is introduced, but can
change it in important ways.
Information Box Fig. 3.1 Beaver (Castor canadensis), North 3.3.2 How Humans Move Invasive Species
America’s largest rodent and one of its most effective ecological
engineers due to their practice of building dams across flowing
streams to create upstream impoundments (“beaver ponds”). Effects, both good and bad, associated with invasive species
(Photo courtesy of Erin Poor, US Geological Survey, public will increase in response to increasing levels of global trade
domain) (Levine and D’Antonio 2003) and climate change (Chap. 4).
Regrettably, the human role in spreading such species has
(continued)
often been deliberate in its intent while ignorant of its at that time was projected at US$120 billion per year
consequences. In the nineteenth century, the Naturalization (Pimentel et al. 2005) (Table 3.3).
Society in New Zealand attempted, and for the most part Most introduced species fail to establish persistent
succeeded, in re-creating the ambience of an English coun- populations, and most of the successful ones that do live
tryside in some parts of New Zealand by releasing common inconspicuous lives among the natives. Some, including
British songbirds, often to the detriment of the native birds of many kinds of crop plants, are beneficial. But a small number
New Zealand (Godfray and Crawley 1998). But even more experience population growth and range expansion that can
introductions are accomplished without any goal at all, as have devastating ecological and economic effects on the
people, along with the material they transport, serve as native communities.
conduits for plant and animal stowaways carried unnoticed
throughout the world. Generally, invasive species reach new
environments either through human commerce in living 3.3.3 Patterns and Characteristics of Successful
organisms (what might be called “intentional introductions”) Invasions and Invaders
or unintended transportation as “stowaways” in planes,
trains, and automobiles or in the packages or ballast they Although details and specifics vary on a case by case basis,
are carrying, or, especially in aquatic invasions, through we can generalize a predictable pattern of successful invasion
human-created artifacts like canals or pipelines that connect characterized by seven distinct stages (Fig. 3.15): introduc-
formerly disconnected areas (Fig. 3.14). In aquatic invasions, tion (intentional or accidental), colonization (sustained resi-
whole non-native communities of creatures attached to the dence on at least one new site), establishment (positive
bottom of ships or living in the ballast water of such vessels population growth on one or more new sites), dispersal,
may be transferred at once to new environments (Ruiz et al. spatially distributed populations, invasive spread, and adap-
1997). Many species of insects may move throughout the tation to the new environment. Whether benign or pestilent,
world in processed or unprocessed wood products, while such invasions cause changes in community composition,
others that feed on vegetables and fruits, such as the Mediter- structure and function. Indeed, it is impossible to understand
ranean fruit fly or “medfly” (Ceratitis capitata) often move existing communities worldwide without an understanding of
worldwide as adults or larvae in produce shipments (Carey species’ introduction and invasion. Thus, Godfray and
1996). Plants may be dispersed long distances as spores or Crawley asserted that “. . .the composition of many (perhaps
seeds, or actively collected and planted under cultivation, even most) communities is determined by the history of
only to later escape to the wild. And humans, including introduction” (Godfray and Crawley 1998).
conservation managers, still introduce non-native plants into Introduced species reduce biodiversity not only through
public and private lands, preserves and wildlands in some competition but also predation. Predation is often most
areas in efforts to revegetate sites affected by fire, erosion, or devastating to prey species when an introduced predator
overgrazing, or to enhance forage production for wild or encounters native prey species that have evolved few or no
domestic herbivores. defenses against it. One example of extinctions caused solely
It is difficult to assess the total number of non-native by an introduced predator is the case of the brown tree snake
species worldwide or their rates of invasion, as estimates (Boiga irregularis) (Fig. 3.16), which reached the Pacific
vary according to phylogenetic group and reference time island of Guam in 1967. The snake’s spread coincided with
frame, but by the 1990s, some scientists were making com- the disappearance and extinction of three species of birds
prehensive investigations and inventories that made quantita- native to the island. Research confirmed that the snake – the
tive estimates possible. From their own research and reviews only predator unique to Guam – was indeed the cause.
of other sources, forest scientist Pekka Niemelä and insect Such narratives of non-native species invasions and their
ecologist Willliam J. Mattson concluded that nearly 2000 effects make for fascinating reading (e.g., Elton 1958;
species of insects and 2000 species of weedy plants had Pimentel 2002; Mooney et al. 2005; Sax et al. 2005). And it
entered North America in the last 500 years (Niemelä and is easy to recite case histories of invasive species “horror
Mattson 1996). Godfray and Crawley (1998) estimated that at stories” recounting how particular invaders devastated native
least 20,000 non-native plant species have been introduced biodiversity in specific places. But invasive species biology
into Great Britain, nearly 1200 of which have become and management has been increasingly criticized for its lack
naturalized. And, among aquatic organisms, Carlton and of a comprehensive theory of “invasion biology,” the
Geller (1993) listed 46 species of non-native species biological patterns and processes that determine their proba-
introduced around the world from ballast-water discharges bility of successful invasion and subsequent spread. Such
since the 1970s. David Pimentel and his colleagues estimated condemnation is just. Only recently has the conservation
in 2005 that 50,000 non-native species had been introduced community begun to shift away from descriptions of and
in the United States alone, and their economic cost to society prescriptions for invasive species on a case-by-case basis.
Fig. 3.14 Major pathways through which non-native species enter new by permission from Wiley, from Lodge, D.M., Williams, S., MacIsaac,
areas, countries, or regions and are transported within them. The right- H.J., Hayes, K.R., Leung, B., Reichard, S., Mack, R.N., Moyle, P.B.,
hand pathway of commerce in living organisms also assumes the left- Smith, M., Andow, D.A., Carlton, J.T., McMichael, A., 2006.
hand pathway of “stowaway introductions” because commerce in Biological Invasions: Recommendations for US Policy and Manage-
selected species entails the possibility of other species “hitchhiking” ment. Ecological Applications 16, 2035–2054. # 2006 Ecological
with the selected species, as well as the possibility of misidentification Society of America)
of selected species at the point of origin or delivery, or both. (Reprinted
The current emphasis is to explicate patterns, principles and life cycle as well or better than native species, or they can
theories regarding such invasions in general that we may exploit a niche that lacks a native species.
better understand individual case histories in particular, and Many invaders are especially adept in category one. For
thus better respond to and manage non-native populations example, in a classic study of the invasive characteristics of
such that native species are conserved. 24 species of pine (Pinus spp.) by Rejmánek and Richardson
(1996), 12 species were classified as non-invasive (planted on
at least three continents but never reported as spreading) and
3.3.4 Understanding Invasive Processes 12 as invasive (spreading on at least two continents). After
evaluating ten life history traits in both groups via discrimi-
Successful invaders are rare. They must be species that pos- nant analysis, Rejmánek and Richardson found only three
sess traits predisposing them to transport by humans and that were significant in predicting invasiveness. These were
survival of the conditions and selection regimes encountered the square root of mean seed mass, the square root of the
during transport, introduction, establishment and spread. For minimum juvenile period, and the mean interval between
all their variety, successful invaders typically possess at least large seed crops. Invasive species had low mass of individual
one of three characteristics: (1) they can deliver seeds, breed- seeds, short juvenile periods, and short intervals between
ing individuals, or other types of propagules at a high rate at large seed crops. The second and third traits allowed invaders
an opportune moment for invasion and at a high density to an to achieve early and consistent reproduction once established,
opportune site or sites; (2) they are able to persist for while the first contributed to higher numbers of widely dis-
extended periods at low densities under unfavorable persed seeds. Interestingly, invasive species were all
conditions until favorable conditions permit their populations concentrated in the same subgenus of pines (Diploxydon),
to grow to higher densities; and (3) they are a good “ecologic while non-invasive species were all members of a different
match” for the environment, and are able to exploit local subgenus (Strobus). This dichotomy suggested that, at least
conditions and abiotic factors that favor completion of their in pines, membership in a subgenus might be the first
Table 3.3 Estimated annual costs associated with some alien species introduction in the United States
Category Nonindigenous species Losses and damages Control costs Total
Total
Plants 25,000
Purple loosestrife – – 45
Aquatic weeds 10 100 110
Mealeuca tree NA 3–6 3–6
Crop weeds 24,000 3000 27,000
Weeds in pastures 1000 5000 6000
Weeds in lawns, gardens, golf courses NA 1500 1500
Mammals 20
Wild horses and burros 5 NA 5
Feral Pigs 800 0.5 800.5
Mongooses 50 NA 50
Rats 19,000 NA 19,000
Cats 17,000 NA 17,000
Dogs 620 NA 620
Birds 97
Pigeons 1100 NA 1100
Starlings 800 NA 800
Reptiles and Amphibians 53
Brown Tree Snake 1 11 12
Fish 138 5400 NA 5400
Arthropods 4500
Imported fire ant 600 400 1000
Formosan termite 1000 NA 1000
Green crab 44 NA 44
Gypsy moth NA 11 11
Crop pests 13,900 500 14,400
Pests in lawns, gardens, golf courses NA 1500 1500
Forest pests 2100 NA 2100
Mollusks 88
Zebra mussel – – 1000
Asian clam 1000 NA 1000
Shipworm 205 NA 205
Microbes 20,000
Crop plant pathogens 21,000 500 21,500
Plant pathogens, in lawns, gardens, golf courses NA 2000 2000
Forest plant pathogens 2100 NA 2100
Dutch elm disease NA 100 100
Livestock diseases 14,000 NA 14,000
Human diseases NA 7500 7500
Total 120,105
Numbers millions of US dollars. NA - estimate not available
Reprinted by permission from Elsevier, from Pimentel, D., Zuniga, R., Morrison, D., 2005. Update on the environmental and economic costs
associated with alien-invasive species in the United States. Ecological Economics 52, 273–288. # 2005 Elsevier B.V.
indication of the possible invasiveness of a species many sites in repeated efforts will have a greater chance of
(Rejmánek and Richardson 1996). success.
Other principles also have helped to explain the success of Many invasive species demonstrate the importance of
invaders in multiple contexts. Specifically, in planned “ecological match” or “pre-adaptation” to a novel environ-
introductions, an introduced species is more likely to be ment. Some aspects of ecological match occur at extremely
successful if (1) more individuals are released rather than broad levels. For example, among herbaceous plants, the best
less; (2) more release sites are used rather than fewer; and indicator of ability to invade a new area has historically been
(3) releases are repeated many times (Veltman et al. 1996). latitudinal range (Forcella et al. 1986, Rejmánek 1995). The
Thus, non-native species that can invade in large numbers at greater the spread of latitude (and, by inference, climatic
Fig. 3.15 The process and stages associated with the invasion, establishment, and spread of a non-native species. (Reproduced by permission
from McGraw-Hill Companies)
fishes in California (USA), Moyle and Light (1996) deter-

mined that abiotic conditions in streams, not characteristics of
native biotic communities, were the most important factors in
successful invasion (Moyle and Light 1996). The most suc-
cessful invaders were those adapted to the local hydrologic
regimes, specifically to patterns of seasonal changes in water
flow (Moyle and Light 1996).
Another dimension of ecological match is seen in species
that can in some way alter the habitat itself, effectively
creating their own niche where one did not previously exist.
An example of this can be seen in the nitrogen-fixing tree
Myrica faya (Fig. 3.17), an invasive species on the US island
of Hawai’i. Myrica faya is adept at colonizing volcanic ash
and open native forests, both of which are nutrient limited
Fig. 3.16 The brown tree snake (Boiga irregularis), an introduced systems. In these systems, Myrica faya can increase inputs of
predator to the Pacific island of Guam, that has exterminated all species
of native birds. (Photo courtesy of P. Kirillov, under CC BY-SA 2.0)
nitrogen up to four times (Lodge 1993). One would think
nitrogen additions might be beneficial to native species in
conditions) that an herbaceous plant can tolerate in its indig- these nutrient-limiting environments. However, for some
enous range, the more likely it is to invade new areas. species, effects of shading and high rates of litter accumula-
Structural habitat components have played a major role in tion under and around Myrica faya outweigh these benefits
creating an ecological match that facilitates invasion success and lead to their decline (Lodge 1993). Thus, Myrica faya
in some groups of organisms. In their review of non-native alters the habitat by changing rates of nutrient cycling as well
population demography, such as exponential or logistic

growth equations, to predict changes in numbers of the inva-
sive species through time. Spatially implicit models,
discussed further in Chap. 8, include spatially structured
data, such as the correspondence between spatial variation
in the environment and population growth, predicted in sim-
ple regression equations, to forecast rates of change in inva-
sive populations as they encounter physical and ecological
variation in a new environment. Spatially explicit models
include spatial locations of invasive organisms and knowl-
edge of “contagious processes” (site- or habitat-specific rates
of movement, birth, or death, among others) to predict the
rate and manner of invasive spread (Peters 2004).
Biometrician J. G. Skellam, in his classic paper “Random
Dispersal in Theoretical Populations” (1951), described the
Fig. 3.17 Myrica faya, a nitrogen-fixing tree species not native to the spread of an invading organism as a type of reaction-diffusion
US island of Hawaii, but successful as an invasive species there because
of its capacity to exploit nitrogen-limited soils, such as those found in
equation, predicting that the advancing front of the organism
open areas or in volcanic ash. (Photo reprinted under CC0 1.0 Universal should travel as a wave at a velocity (V) expressed as
Public Domain)
pffiffiffiffiffiffi
V ¼ 2 rD
as physical structure and light penetration of forests, creating
a new niche favorable to itself, but which native species where r is the population’s intrinsic rate of increase and D is
cannot tolerate. the diffusion coefficient, equal to one-half the mean squared
Invasive species can not only change ecological processes, distance moved in a time unit by an organism (Godfray and
but alter long-term evolutionary pathways through competi- Crawley 1998). For example, D has often been expressed in
tive exclusion, niche displacement, hybridization, introgres- km2/year (e.g., Grosholz 1996). For all its simplicity, the
sion, predation, and, ultimately, extinction. The invaders predictions of this equation have some record of success in
themselves evolve through their interactions with native spe- matching observed patterns of invasive spread in many spe-
cies and their new environment. Some invasive species cies (Grosholz 1996, Godfray and Crawley 1998). The speed
accomplish evolutionary effects by eliminating native species of advancement in a muskrat (Ondatra zibethicus) invasion,
altogether, usually either through competitive exclusion or for example, was locally constant, but influenced by topogra-
predation. For example, in Texas (USA), the invasive fire ant, phy and habitat preference (i.e., the muskrat’s affinity for
(Solenopsis invicta), a native of Bazil, reduced native (Texas) wetlands) (Skellam 1951). The model, however, can be
ant diversity by 70%, total number of native ant individuals modified to account for spatial heterogeneity and the patchy
by 90%, non-ant arthropod diversity by 30% and numbers of distribution of habitats by taking the form.
non-ant arthropod individuals by 70% through competitive pffiffiffiffiffiffiffiffiffiffi
exclusion (Mooney and Cleland 2001). V ¼ 2 r a Dh
where ra is the arithmetic mean of the population’s intrinsic

3.3.5 Forming a “Theory of Invasion Biology” – growth rate across patches and Dh is the harmonic mean of
Past Efforts the diffusion coefficient across patches (Shigesada et al.
1986). Both values can be calculated if one knows the rates
Scientists concerned with the dynamics of invasive processes of growth and spread in a sufficient number of habitat patches
have traditionally resorted to one of three categories of to calculate a reliable average. This model also has performed
models to explain invasion behavior, spread, and success. well when subjected to more stringent experimental tests in
Nonspatial models are the simplest. In these, knowledge of which the value of r was calculated from a life table and the
the spatial locations of invading organisms is not known and value of D from controlled experiments of dispersal, then
the “contagious processes,” such as transfer rates of compared to data associated with an actual invasion. In a
individuals from one point to another, also are unknown. classic review of the model, Andow et al. (1990) subjected
These models simply use projections derived from the wave model to such tests with an invasion of muskrats,
the cereal leaf beetle (Oulema melanopus, a European insect from source point y to destination point x is a function of the
introduced in the US in 1958) and a butterfly (Pieris rapae, shape of the dispersal “kernel,” k. Interestingly, IDE models
also European and introduced to the US several times in the reveal that it is the long-distance component of dispersal that
nineteenth century). The model performed well for the musk- ultimately governs invasive speed, even when long-distance
rat and butterfly, but drastically underestimated the rate of dispersal is rare (With 2002). Models that lack the element of
spread in the beetle. Andow et al. (1990) believed that the stage-structured dispersal, like reaction-diffusion models,
model failed for the beetle because it did not incorporate the overestimate the speed of invasion for many species.
effects of rare, long-distance dispersal by a few individuals. IDE models are examples of stratified diffusion models in
Such events, although uncommon, can affect the rate of which elements such as long-distance dispersal and density-
spread. More sophisticated models attempt to incorporate dependent rates of spread are incorporated more explicitly.
such effects, along with effects of density dependence and Populations at different distances from the source of invasion
carrying capacity as inner circles become saturated with are assigned different values of r and D, creating different
individuals. Veit and Lewis (1996) constructed a model strata in the dispersing population. Thus, stratified diffusion
with such elements to explain the spread of the house finch models resemble age-structured models of population growth
(Carpodacus mexicanus) from a small population of about (Chap. 6), except that the founding of each colony in succes-
250 birds in New York to a population now covering most of sive strata takes the place of “birth” and colony growth takes
the US. Their model predicted that range expansion would be the place of aging. If new colonies form near existing
slower than expected by traditional models due to Allee colonies and coalesce, the rate of spread changes from
effects (effects associated with decreased per capita growth accelerating to linear, creating a so-called “starburst” to dis-
rates in small populations), but speed of distribution would tinguish it from the traditional “traveling wave” pattern.
increase as the population grew. Both predictions were con- Godfray and Crawley (1998) note “Both types of spread
firmed in historical data. (“traveling wave” and ‘starburst’) are seen as two ends of a
A problem with simple reaction-diffusion models is that, single continuum; the key parameter is the distance between
although the population density is allowed to vary across the successive foci of establishment. When this distance is small,
landscape, the landscape itself is assumed to be spatially the assemblage behaves like a traveling wave, but when it is
homogeneous, redistribution of invading individuals is large starburst effects predominate.”
assumed to occur as a random dispersal process and dispersal Other patterns exist in which factors other than population
and reproduction of the invading species is assumed to occur growth and diffusion coefficients are controlling agents of
simultaneously and continuously. These conditions are met spread. For example, With (1999) developed early landscape-
in some species, but many invasive species, especially based models that examined species with differing dispersive
insects, disperse in one life stage and reproduce in another. abilities in landscapes with random, fragmented, and
And many studies of invading organisms reveal that clumped habitat dispersion. With’s models indicated that
differences in landscape characteristics are not only present, potential for the spread of an invasive species is enhanced
but important to an invading organism. Therefore, long- past a threshold level of landscape fragmentation (see
distance dispersal events, in which invaders “leapfrog” examples in Chap. 7), but the absolute level of such a thresh-
ecological and landscape barriers, although rare, are old is affected by characteristics of both the landscape and the
significant. invasive species. Once the disturbance threshold has been
Integro-difference equations (IDE) offer one way to incor- reached, an invasive species will spread faster in more
porate long-distance dispersal functions and stage-specific clumped, contiguous habitat than in more fragmented habitat,
reproduction and dispersal. IDE models break dispersal and suggesting that when an invasive species with limited disper-
population growth into separate stages through (1) a differ- sive abilities is also a habitat specialist, deliberate habitat
ence equation that describes population growth at each point fragmentation could slow or stop invasive spread at relatively
in the landscape and (2) an integral operator that accounts for low levels of landscape disturbance. For example, the
the pattern of dispersal of organisms in space (the so-called US Forest Service creates “barrier zones” at invasion fronts
“dispersal kernel”). Thus, all IDE models have the general of advancing gypsy moths (Lymantria dispar), a Eurasian
form of species that invaded North American in the late nineteenth
century, and whose larvae damage many kinds of trees.
Nt þ 1 ðxÞ ¼ kðx, yÞf ½Nt ðyÞdy Forest Service workers at the barrier zone employ direct
suppression or eradication actions against the moth (Sharov
Where Nt + 1 (x) is the population density at some destination and Liebhold 1998). The same strategy could be used by
point x, which is a function of the population growth at each creating habitat barriers that a habitat-specialist invasive spe-
source point y (f[Nt(y)]) and the movement of individuals cies could not cross (With 2002).
3.3.6 State of the Art: Current Theories

and Management Paradigms for Invasive
Species
Invasive species are very much an “Anthropocene” problem,

so their management must also be understood from an anthro-
pogenic standpoint – how do invasive species interact, and
often take advantage of, human modifications of natural
environments? One contribution has been offered by
biologists Christopher Kueffer and Curtis Daehler (Kueffer
and Daehler 2009), who propose a habitat-classification
framework based on four habitat types, with categories
defined by differences in degree of human habitat modifica-
tion and valuation of these habitats (Fig. 3.18, Table 3.4). The
Fig. 3.18 Four habitat types (anthropogenic, reference, abandoned,
first type, what we might call “pure” anthropogenic habitat, designed) that represent different environmental and management
consists of highly disturbed areas such as agriculture, planta- contexts of relevance for biotic invasions. The habitats are arranged
tion forestry, or urban settlements. The second type of habi- according to degree of human influence (unassisted vs. human-assisted
tat, which Kueffer and Daehler refer to as “reference habitat,” processes) and relevance for nature conservation (core vs. matrix nature
conservation areas). See text for further explanation. (Reprinted by
represents relatively undisturbed habitats dominated by permission from Springer Nature, from Kueffer, C., Daehler, C.C.,
native species. Between these two extremes, there is “aban- 2009. A Habitat-Classification Framework and Typology for Under-
doned habitat,” which currently experiences little human standing, Valuing, and Managing Invasive Species Impacts, in: Inderjit
interference but have been disturbed or managed in the (Ed.), Management of Invasive Weeds. Springer Netherlands, Dordrecht,
pp. 77–101. # 2009 Springer Science + Business Media B.V.)
past, such as agricultural land, abandoned plantation forests,
or degraded urban areas. Abandoned habitats may be of
actual or potentially high conservation value but cannot be
classed as reference habitat because their current condition when conditions are favorable and their numbers support
cannot be considered a reference for “high quality nature.” A greater dispersal. In fact, with sufficient population increase
fourth habitat type, “designed habitat,” is habitat that humans and local adaptation, invasive species may be able to
deliberately manipulate to create new habitat directed to “launch” a colonizing effort directly into native “reference”
specific conservation objectives (such as restoration of a habitat.
former native habitat). Some invasive species possess traits that can make them
These four classifications match four main strategies used successful even in these invader-resistant environments,
in nature conservation today. These are (1) maintaining bio- including reference habitat. As Kueffer and Daehler note,
diversity in cultural and urban landscapes, (2) protecting “invaders of reference habitat may have a combination of
natural areas, (3) practicing nature conservation on aban- traits similar to the native biota and novel traits. It can be
doned land, or (4) actively restoring habitats that have been expected that species that lack the latter may have minor
degraded restoration. This four-habitat classification scheme ecological impacts. . .Species with novel traits, in contrast,
also lends itself to classifying invasive species according to can have substantial impacts. . .” (Kueffer and Daehler
four parallel categories of life history. In the first category, 2009:87). And if the natural habitat contains rare native
invaders adapted to anthropogenic habitat, Kueffer and species, even minor effects of non-native species can increase
Daehler note that such species are “resource-demanding, risk of extinction.
fast-growing, and fast-reproducing ruderal or early succes- In abandoned habitat, one often sees unique combinations
sional species. If plant species, they usually have high seed of native and non-native species. Thus, abandoned habitat
output, and an efficient seed dispersal mechanism (particu- may be the ideal “breeding ground” for the creation of the
larly wind or birds)” (Kueffer and Daehler 2009:84). These novel ecosystems discussed in the previous section of this
are the “weedy invaders,” species that have offered the clas- chapter. “Abandoned habitat will initially have conditions
sic case histories that supported the development of invasive similar to anthropogenic habitat, and alien species invading
species biology. Given these species traits, anthropogenic anthropogenic habitat may also most commonly invade aban-
habitat itself is a critical component of successful invasion. doned habitat in an early phase of secondary succession”
Whereas natural habitat with a well-established array of (Kueffer and Daehler 2009:90). In abandoned habitat, inva-
native species may be resistant to the invaders, anthropogenic sive species can cause changes in ecosystem properties, such
habitat is likely to have few, or even no, native species. This as fire return interval, which can in turn change species
permits the establishment of an invasive “outpost,” where composition of the community as well as patterns of nutrient
invaders can build up their populations, adapt to local cycling, and which then further alter the frequency of fires
conditions, and “wait for an opportunity” to spread further (Vitousek 1990; D’Antonio and Vitousek 1992; Brooks et al.
Table 3.4 Four habitat types of relevance to biological invasions

Habitat
Habitat type characteristics Traits of invader Impacts of invader Management action Research focus
Anthropogenic Disturbed or “Weedy” invaders Reservoir for Reduction of propagule Upscaling of
habitat anthropogenic such as ruderal, or invasions into pressure control methods to
early successional biodiversity areas large areas
species Competition with Mainstreaming invasive Socio-ecological
native ruderal species control in different research on plant
species production sectors invasions,
e.g. urban and
agricultural ecology
Increase of
biodiversity of
anthropogenic
habitat, and
substitute for native
species
Reference Undisturbed, “high Traits similar to May be low, if no Early detection and Long-term and
habitat quality nature” native species, but novel traits are eradication indirect impacts
possibly with some involved
novel traits May be indirect or Monitoring of native biota Indicators of
cryptic and ecosystem functioning ecosystem health
Long-term effects Ecology of rare
uncertain native species and
communities
Abandoned Abandoned land Depends on initial Positive, negative, Directing secondary Stability and
habitat after prior exposure state and or both succession according to functioning of
to strong successional stage of management goals novel assemblies of
anthropogenic the habitat (time native and alien
disturbance since disturbance) species
Sustaining ecosystem Provision of
functioning rather than ecosystem services
restoring native dominated and products
habitat including native
biodiversity
Designed Strongly managed Management and Depends on Ecosystem design: Restoration
habitat for nature habitat conditions management goals Manipulation of native and techniques
conservation and may select for and stage in the alien species to attain
constantly depending specific types of restoration process specific conservation-
on management invaders oriented objectives
Deliberately Ecology of
introduced alien community
species with traits assembly
that facilitate Risks versus
restoration benefits of alien
species use
Socio-ecological
research on natural/
artificial dichotomy
Reprinted by permission from Springer Nature, from Kueffer, C., Daehler, C.C., 2009. A Habitat-Classification Framework and Typology for
Understanding, Valuing, and Managing Invasive Species Impacts, in: Inderjit (Ed.), Management of Invasive Weeds. Springer Netherlands,
Dordrecht, pp. 77–101. # 2009 Springer Science + Business Media B.V.
2004). But invasive species, especially invasive plant spe- present ideal conditions for testing hypotheses about invasive
cies, can also play a role in ecological restoration, if they species biology and behavior (Kueffer and Daehler 2009).
perform ecological functions similar to native species that are Although they may have very different functions and
no longer present, as in Mascaro et al.’s (2012) previously purposes from other anthropogenic habitats, designed
discussed study of novel forest ecosystems in Hawaii. In habitats also have similarities to them. The process of resto-
doing so, invasive species change (sometimes for the better) ration, which is common, indeed, typical, in designed
abiotic conditions in abandoned habitat, such as soil fertility, habitats, often simplifies the natural or reference habitat in
soil stability, or proportion of organic matter (Schlaepfer both species composition and ecological processes. Because
et al. 2011). For these reasons, abandoned habitat may existing vegetation may be removed to initiate restoration,
designed habitats, like other anthropogenic habitats, often especially if the land has been covered by an impervious sur-
have high light availability, a condition favorable to many face such as asphalt or concrete. The daily commuting radius
invasive species of plants. It is both ironic and unfortunate is large, often making urban areas centers of congested traffic
that, because restoration efforts and research are usually and accompanying high rates of greenhouse gas emissions
focused on native species brought back to the target site, from vehicles (take it from one of us who lives near the US
they have tended to give relatively little attention and effort Washington DC-Baltimore corridor). By 2030, it has been
to research on invasive species which may enter during the projected that urban land cover (the “built up” environment)
restoration process. will increase by 1.2 million km2, a value that will be nearly
The habitat classification approach to understanding the three times what urban coverage was at the beginning of the
traits and vulnerabilities of different categories of habitat present millennium (Seto et al. 2012). By 2050, the propor-
provides a management-oriented perspective that suggests tion of humans living in urban environments is projected to
specific actions that could stop, remove, or control invasive be 70% (United Nations Human Settlement Program 2012).
species in each setting. These recommendations would be to Such increase will not consist simply in the expansion of
(1) in anthropogenic habitat, manage invasive species like current urban areas, but in the development of new urban
“weeds,” (which, if plants, will be an apt and accurate areas in natural environments, including many of the world’s
description) by targeting removals that have the most effect biodiversity hotspots (Chap. 2, Seto et al. 2012). A develop-
on limiting economic damage and “nuisance” effects; (2) in ment of greater consequence is the relationship between the
reference (natural) habitats, attempt complete eradication increase of the urban area and the increase of the urban
when the infestation of invasive species has just begun and population. Historically, urban areas grew slowly, often in a
its populations are small, because even minor effects of pattern of concentric spread. Within the last 30 years they
invasive species can have disproportionately negative have grown much more rapidly, increasing in area twice as
consequences for native species, especially those that are fast as urban populations have been increasing in size (Seto
already threatened or endangered; (3) in abandoned habitats, et al. 2012).
consider making deliberate but carefully chosen The effects on global biodiversity forecast by such
introductions of non-native species if they can perform spe- changes are consequential. Among the sites designated of
cific ecological and environmental functions that can no high value by the Alliance for Zero Extinction (AZE) – a
longer be performed by native species which have been network of over 100 non-governmental biodiversity conser-
eradicated from the site, and use an adaptive management vation organizations working to prevent species extinctions
approach to monitor their effects and test hypotheses about by identifying and safeguarding sites where species evaluated
their ecological roles; and (4) in designed (restored) habitats, to be Endangered or Critically Endangered under IUCN
monitor the presence or appearance of non-native species criteria – are areas affected by urban expansion. Comparing
carefully, removing unwarranted non-natives, but planting areas of high urban expansion with AZE designated areas,
or permitting the presence of selected non-native species Karen Seto of Yale University (USA) and her colleagues
that may advance restoration objectives through the ecologi- determined that “More than a quarter of species in amphibian,
cal functions they perform (Kueffer and Daehler 2009). mammalian, and reptilian classes each will be affected in
varying degrees from urban expansion in AZE sites. . .
Habitats are expected to be encroached upon or destroyed
3.4 Biodiversity Conservation in Urban by urban expansion for 139 amphibian species,
Landscapes 41 mammalians species, and 25 bird species that are on either
the Critically Endangered or Endangered Lists of the Interna-
3.4.1 Growth of the Urban Landscape tional Union for Conservation of Nature (IUCN)” (Seto et al.
2012:16084).
As anthropogenic biomes (“anthromes,” Sect. 3.1.3) now Seto et al.’s (2012) analysis exemplifies the more general
dominate the habitable surface of the Earth, none is more threats that urban areas pose to biodiversity on multiple
human-dominated than the urban environment, and none is fronts, not simply by their expansion, but in their present
having a greater effect on global conservation. In land area, in condition. Urbanization of an area is equivalent to permanent
human population density, and in proportion of Earth’s total or near-permanent changes in land use and effective elimina-
human inhabitants, urban environments are growing. Today tion of natural ecosystems on that site (Beninde et al. 2015).
more than half (55%) of the world’s nearly 8 billion As such, urbanization has been understandably perceived as a
inhabitants live in urban areas. Urban areas are characterized major threat to global biodiversity (McKinney 2002, Grimm
by high densities of humans as well as built up areas in which et al. 2008). Measurable variables associated with negative
natural habitat has been “modified” (more honestly, “lost”) in effects of urbanization on biodiversity increase as one moves
ways that will make it difficult or impossible to reclaim, closer to core urban areas. These include increases in human
3.4 Biodiversity Conservation in Urban Landscapes 107
population density, road density, air and soil pollution, aver- applicable to urban settlements as it is to areas in remote
age ambient temperature, average annual rainfall, soil com- wilderness or uninhabited locations” (Pickett and Cadenasso
paction, soil alkalinity, and other indicators of anthropogenic 2006:115).
disturbance (McKinney 2002). The percentage of impervious What looms larger in cities than in other environments is
surface increases from 20% at the fringe of urban areas to people – not only their higher densities but their even higher
50% in urban core areas (McKinney 2002). Similarly, in impacts. As such, cities also immediately create a problem for
those taxa that have been carefully studied in urban areas, the traditionally educated ecologist who has been trained to
species richness declines in plants, birds, many insect groups, exclude human effects from a “correct” or “pure” understand-
including butterflies, and mammals. In all of these taxa, the ing of how the ecosystem is “supposed to work.” Pickett and
number of species at the urban core is reduced to less than Cadenasso again offer helpful insight, stating, “. . .urban, and
half of those found in the rural, more natural areas indeed all ecosystems inhabited by humans, must be modeled
(summarized in McKinney 2002). The volume of vegetation to include the human individuals as well as the social
coverage is reduced by mowing, pruning, and other landscap- aggregations they generate or influence. It is not sufficient
ing practices that are intensified in urban areas. to include humans only as demographic entities,
Urban areas exemplify the most intensive cases of anthro- characterized by density, birth, death, and migration. Humans
pogenic environments, as well as sites that generate multiple must also be represented by the institutions that organize and
forms of novel ecosystems, concepts we have introduced in affect daily life, by the cultural and social resources they and
earlier sections of this chapter. Compared to non-urban areas, those institutions rely on, and by the temporal dynamics
the reduced coverage of vegetation, and, in its place, large exemplified by individuals, families, organizations,
amounts of land covered by impermeable surfaces, such as neighbourhoods, economies, etc. . .” (Pickett and Cadenasso
asphalt or concrete, increase rates of runoff and decrease heat 2006:115). And such “institutions” are one of the signature
capacity (the ability to absorb energy without change in characteristics of urban environments. Cities are centers of
temperature). As a result of their impervious surfaces, urban human institutions – political, educational, social, and
areas discharge high levels of water into surrounding streams, scientific – and in cities the organizational effects of such
rivers and lakes –water often contaminated by the surfaces it institutions are high and their influence is strong. Human
could not penetrate but from which it absorbs nutrients and institutional organization and operation in urban
chemicals, some of them toxic, which are part of the urban environments strongly influence ecosystem processes as
environment. As a result of decreased heat capacity, urban well as ecosystem composition.
areas are usually “heat islands” with average temperatures To incorporate human effects more explicitly into under-
higher than surrounding rural landscapes, and so contribute standing ecosystem processes, scientists from multiple
disproportionately to regional climate change. disciplines have collaborated to develop the Human Ecosys-
Even with continued rapid increase in human population tem Framework (HES), a conceptual model that integrates
levels and densities, urban areas themselves still take up only the human social system with the resource system and bio-
about 8% of the world’s land cover, about two acres or one physical resources which it affects, but upon which it also
hectare in 12. The greatest effects of urban areas do not arise depends (Fig. 3.19). Changed understanding leads to
from the land they occupy, but from the effects created by changed thinking, and changed thinking to changed termi-
such intensive use and the amount of resources consumed nology. In the past, studies of ecological processes and
and processed to sustain it. effects in urban areas were often referred to as studies of
“urban ecology.” Today most scientists studying the same
processes and effects in an urban area refer to their
3.4.2 Understanding Cities as Ecological investigations as studies of “urban ecological systems” and
Systems “urban ecological research” because urban areas do not have
a special “ecology” that involves different ecological pro-
Most established ecologists and conservationists have been cesses compared to non-urban systems. Rather, urban eco-
trained in their craft far from urban areas and think of cities logical systems are distinguished by the fact that such
as, at best, “non-habitat.” Yet ecological processes that trans- processes are operating in a different system of landscape
form energy and matter and mediate species interactions and human interaction, one which is distinctive of the urban
occur in cities with the same inexorable certainty that they environment.
occur in wildlands and rural areas. Pickett and Cadenasso Consistent with such thinking, the US National Science
(2006) offer a first step toward changed thinking. “Ecologists Foundation’s (NSF) Long-term Ecological Research (LTER)
are quite comfortable studying complex systems, comprising program now includes two major US metropolitan areas,
biotic and abiotic components, which are linked by flows of Baltimore, Maryland and Phoenix, Arizona. Studies in
matter, energy, and information. The basic concept is as Baltimore, for example, have led to new insights about how
Fig. 3.19 The Human Ecosystem Framework (HES), which includes a Wiley, from Pickett, S.T.A., Cadenasso, M.L., 2006. Advancing urban
complete roster of the structural and functional variables that can moti- ecological studies: Frameworks, concepts, and results from the
vate hypotheses and inform models addressing inhabited or managed Baltimore Ecosystem Study. Austral Ecology 31, 114–125. # Ecologi-
ecosystems, and which provides an excellent conceptual understanding cal Society of Australia)
of the urban ecological system (UES). (Reprinted by permission from
the urban environment changes (and, in some cases, does not 3.4.3 Explaining Urban Ecological Responses:
change) basic ecological processes. Pickett and Cardenasso, Traditional and Contemporary
who have studied the Baltimore system for many years, note Approaches
that, for example, in the case of nitrogen cycling, “The
amount of N retained on an annual basis is, as expected, Habitat changes that accompany increasing levels of urban
greater in the forested watershed [in the greater Baltimore land uses operate as key “filters” in determining the species
area] (94% on a mass basis). Agriculture retains 85% of N composition or urban areas (Kowarik 2011). Given the filter-
applied to that land cover type, while suburban watersheds ing effect that operates in this manner, one of the most
retain 71% of N inputs. The suburban retention rate is sur- common tools for interpreting the effects of urbanization on
prisingly high, and suggests that the combination of ecosystems and their biodiversity is the use of the rural-to-
maintained lawns, volunteer vegetation, and engineered urban gradient (RUG). This tool is widely used because it has
retention basins as biophysical infrastructure trap substantial proven widely useful, providing a way of understanding not
amounts of N. This suggests that human structures have not only general patterns, but, more specifically, land coverage
obliterated all biological processes that determine ecosystem (an index of habitat modification and loss, as well as species
function in this suburban catchment. . .” (Pickett and richness (Fig. 3.20, McKinney 2002). A RUG approach also
Cadenasso 2006:122). invites straightforward interpretation in changes of such
Fig. 3.20 A generalized

depiction of the rural-to-urban
gradient (RUG). Species richness
declines toward the urban core as
anthropogenic land uses
intensifies, reflected in the
increasing proportion of
impervious surface (mainly
asphalt and concrete). Not only
does the number of species change
(species richness), but a pattern of
species replacement occurs such
that species in the urban core are
uniquely adapted to exploit
particular characteristics of the
urban environment. (Reprinted by
permission from Oxford
University Press, from McKinney,
M.L., 2002. Urbanization,
Biodiversity, and Conservation.
BioScience 52, 883. # 2002
Oxford University Press)
variables as the amount of subsidized energy and matter relying on a one-dimensional linear gradient to describe
imported for use by humans and available to other species, ecological response, the DUF approach integrates under-
both of which increase from a city’s rural fringe to its urban standing of past land use in the urban environment
center. The RUG also shows a clear pattern of change for (so-called “land-use legacies”), the spatial configuration of
nonnative species, which increase with movement toward remnant natural environments within the city, the period of
urban core habitat. time over which urban development has been taking place
As faithful as the RUG has been as an explanatory work- (“urbanization age”), the local environments characteristics
horse for effects of urbanization on ecology and biodiversity, of geology, soils, and disturbance regimes, and socio-
some investigators are critical of it as too simplistic and no economic characteristics of the specific urban environment
longer applicable to the current pattern of urban expansion. In that drive patterns of current land use (Ramalho and Hobbs
the past, cities have been well-defined areas that expand 2012) (Fig. 3.21). The DUF, and other frameworks derived
through concentric growth, making the RUG approach sub- from or inspired by it, are likely to become more common in
stantially accurate in tracking change along a rural to urban evaluating ecological characteristics of cities, and increas-
gradient. Modern urban expansion often follows a different, ingly replace the RUG method and other linear gradient
non-linear pattern of growth. Today’s cities are expanding approaches.
more rapidly than in the past and are more dispersed in their
configuration (Ramahlo and Hobbs 2012). As a result, linear
gradients no longer fit or properly explain variation in species 3.4.4 Biodiversity in Urban Landscapes
diversity and other ecological variables in more modern
cities. As ecologists Cristina Ramahlo and Richard Hobbs 3.4.4.1 Effects of Urban Landscapes
put it, “. . .the fact that contemporary cities grow in a rapid, on Biodiversity – A Review of Traditional
complex, non-linear, dispersed and expansive manner, means Perceptions
that urbanization intensifies and ages in patchy and complex Conservationists are accustomed to thinking that their pri-
spatial patterns across the landscape, rather than in a linear mary goal is to preserve “natural” environments. Some cities
gradient . . .the environmental or ecological conditions in one have none, but the biological environments they do present
focal remnant patch depend not on its position along the may be the only ecosystems that many people will ever
linear gradient, but on the characteristics of the neighboring know, and the only places in which they will encounter
patches” (Ramalho and Hobbs 2012:181). Beninde et al. nature’s biodiversity in some form. Thus there can be only
(2015) criticize the RUG approach because “Despite its agreement with the words of Sophie Parker, a senior scientist
practicability, it greatly generalizes the urban landscape and with The Nature Conservancy (TNC), who stated that “The
makes an interpretation of results ambiguous with regard to conservation of biological diversity is closely linked with the
decisive underlying specific features of the urban landscape, fate of the world’s cities. While the protection of sensitive
which are not quantified. . .” (Beninde et al. 2015:581). One and threatened species and habitats has often taken place in
alternative to the RUG is the Dynamic Urban Framework natural landscapes largely devoid of people, strategies for
(DUF) approach (Ramalho and Hobbs 2012). Instead of preserving the Earth’s biodiversity that can be employed
Fig. 3.21 The Dynamic Urban Framework (DUF), an emerging para- urban area based on their length of time in an urban landscape and their
digm for research in urban ecological systems. The traditional rural-to- past spatial configurations and land uses. (d) The DUF also uses a more
urban (RUG) framework (left half of figure) classifies degree of urbani- multi-dimensional ecological perspective that evaluates range of varia-
zation on a linear scale based on distance from the urban center (a) or tion in the ecological community, past and present disturbance regimes,
metrics indicative or urbanization such as road density (b). Ecological variation in local conditions (e). These and other factors are evaluated to
responses are then assumed to be functions of the amount of remnant understand causal and interactive relationships of drivers that the eco-
natural areas and their connectivity to one another or of single-effect logical response. (Reprinted by permission from Elsevier,
relationships between variables associated with land use of socio- from Ramalho, C.E., Hobbs, R.J., 2012. Time for a change: dynamic
economic characteristics of the urban environment (c). A DUF uses a urban ecology. Trends in Ecology & Evolution 27, 179–188. # 2011
time-sensitive perspective that evaluates focal natural remnants in the Elsevier Ltd)
within cities are likely to become more common as urban 3.4.4.2 Non-native Species, Endangered Species
areas continue to increase in size and number” (Parker and Urban Biodiversity – Cause and Effect
2015:683). It is foolish and false to think that urban areas or Effect and Cause?
and biodiversity hotspots are remote from and widely Urban areas are often environments with high concentrations
separated with respect to one another. Over 20% of the of non-native species. Recall that invasions by alien species
world’s human population resides within areas designated are widely accepted as a leading cause of biodiversity loss.
as biodiversity hotspots since 2000, which includes many However, much evidence for this claim is based on
living in dense urban areas (Cincotta et al. 2000, Balmford correlations between, as Kowarik puts it “exotic dominance
et al. 2001, Parker 2015). and native species decline in degraded systems” (Kowarik
2011:1980). But often the cause and effect relationship is the (McKinney 2002). This categorization scheme originated
other way around. “The decline of previously occurring from studies that examined species presence along a rural-
native species is often due to habitat loss and transformation to-urban gradient (RUG) described earlier. Return now to
associated with urban land uses, which in turn provide Fig. 3.19 to see a visual representation of one way of
beneficial conditions for introduced species” (Kowarik categorizing urban species.
2011:1980). Therefore, broad generalizations about cities Along the RUG, urban avoiders are animal species sensi-
and non-native species may be inaccurate. Yet cities remain tive to human persecution and habitat disturbance, such as
not only important points of entry for many non-native spe- large mammals, especially predators. Among plants, urban
cies but also points of secondary release of such species to avoiders would include species sensitive to human activities
other areas, as has been documented in many ecological such as late-successional (old-growth) and wetland plants
studies (summarized in Kowarik 2011). Urban land use (Stein et al. 2000), as urbanization leads to increasing levels
patterns, including destruction of native habitats, coverage of deforestation and drainage of wetlands for agriculture and
of land by impervious surfaces, and climate modifications residential development. Urban adapters are often found in
can all act to facilitate entry and spread of non-native invasive suburban landscapes. In this context, early successional spe-
species (Dukes and Mooney 1999, Kowarik 2011). Not only cies of plants may not only be common but deliberately
can invasive species be spread by changes in land use introduced and managed in residential yards and public
associated with urban environments, but equally by human parks, or they may be the first to colonize abandoned habitats
activities in urban areas that affect species selection (which (vacant lots). These early successional plants include both
may favor non-native species over native species), manage- cultivated species favored by humans (e.g., turfgrass, fast-
ment of urban green spaces (which may intentionally intro- growing ornamental shrubs, and trees), as well as weedy
duce large populations of non-native species) (Kowarik species common in both managed and unmanaged habitats.
2011) and increased use of multiple means of transportation Among animals, urban adapters include many species often
within and around cities (which may provide translocation of referred to as “edge species,” especially species adapted to
non-native species and dispersal of their seeds or propagules) boundaries between forests and open areas. Urban adapter
(multiple studies summarized in Kowarik 2011). animals often exploit human-subsidized foods, such as
Cities themselves are often located in areas of high biodi- cultivated plants and garbage. As human density increases,
versity, and their conditions often create endangerment for so does the abundance of such subsidized food, which can
resident species, so it is not surprising that cities may have lead to high densities and biomass of species in this category
high levels of endangered species – not because the cities are that can exploit it (the so-called “garbage can” guild), much
a refuge for those species but because they are the reason higher than would be found in natural areas (Adams 1994,
for their endangerment. Likewise, cities are often richer in Marzluff 2001).
plant species than rural areas, but such richness is often an Urban exploiters probably represent the most
artifact of high numbers of non-native species in the urban homogenized of the world’s biotas (Blair 2001). Unlike
environment, a fact that often “homogenizes” urban plant urban adapters, which are largely composed of early succes-
biodiversity in cities from different regions, or even sional species from nearby ecosystems, urban exploiters are
continents (Kowarik 2011). In a study of Australian cities, composed of a very small subset of species well adapted to
Christopher Ives of Leuphana University (Germany) and his intensely modified urban environments wherever humans
colleagues determined that “Australian cities support sub- construct them (Adams 1994, Johnston 2001, Marzluff
stantially more nationally threatened animal and plant species 2001). Urban environments typically have more in common
than all other non-urban areas on a unit-area basis. Thirty with other cities than with adjacent natural ecosystems, so
percent of threatened species were found to occur in cities” urban exploiters like rock doves, starlings, house sparrows,
(Ives et al. 2015:117). This pattern is not exclusive to Norway rats, and the house mouse are found in all cities in
Australian cities. Twenty-two percent of the occurrences Europe (Mackin-Rogalska et al. 1988) and North America
of endangered US plant populations were located in the (Adams 1994), a pattern also characteristic of urban plants
40 largest US metropolitan areas (Hahs et al. 2009, cited in (Whitney 1985, Kowarik 1995). Such species are particularly
Ives et al. 2015). But how do different species relate to cities, adept at non-linear expansion in urban environments (Sect.
and are there patterns of species success and failure that 3.4.3), and can also “leapfrog” from city to city despite large
would permit us to categorize responses to urban distances between.
environments? Many species in urban areas are uniquely urban species,
often cosmopolitan in distribution, and so considered of
3.4.4.3 Categorizing Urban Biodiversity “Least Concern” in conservation priority by organizations
A common categorization scheme to describe species’ like IUCN. But are there species unique to cities that deserve
response to cities consists of classifying them as urban conservation protection, and, if so, can cities make unique
avoiders, urban adapters, or urban exploiters (synanthropes) contributions to global biodiversity by protecting them?
3.4.5 Can Urban Areas Be Managed “protect nature from people” is no longer effective as a sole
for Biodiversity Conservation? justification for protecting urban nature. The modern city
offers no “nature without people.”
Regarding urban ecological systems, ecologist Ivan Kowarik Today there is increasing interest in “urban conservation,”
noted, “These novel urban ecosystems may challenge the a subject that has grown so large and so rapidly it is difficult
perception of conservationists in different ways. First, one to define it. Some of the key emerging ideas in urban conser-
must consider whether ecosystems that represent profound vation, however, include preserving biodiversity within cit-
human-induced changes in natural systems have value or ies, protecting natural areas providing resources and
whether they are per se to be negatively viewed because ecosystem services upon which cities depend, conserving
they diverge from natural settings” (Kowarik 2011:1979). resources within cities to reduce the city’s ecological foot-
That is indeed the foundational question for conservation of print and enhance its sustainability, and gathering and
urban ecological systems – do they have value in conserving organizing support for conservation efforts from urban-
biodiversity? Traditionally, conservationists have given two dwelling constituencies through educational outreach and
reasons for promoting conservation activities in urban advocacy (Dearborn and Kark 2010; Parker 2015). The prob-
settings: (1) as a contribution to biodiversity conservation lem with this burgeoning diversity of strategies are that many,
and, (2) to enable social functions and provide ecosystem perhaps most, conservationists have been educated and
services for humans living in an urban environment trained in their knowledge and practice of conservation in
(summarized in Kowarik 2011). First we will address the relatively natural areas, often far from urban environments.
question of conservation value. Can cities be managed to Because such deficiency of education is widespread, many
enhance biodiversity? conservationists do not know how to interpret urban
Like most of the world, conservation in cities has tradi- landscapes in conservation perspective, either in recognizing
tionally followed a strategy of finding what natural areas the problems to be solved or the opportunities to be exploited
remain and preserving these as “parks” or “reserves,” and with respect to conservation concerns.
many cities understandably take pride in this expression of Such a perception, or perhaps, lack of perception, is why,
conservation. In the US county of DuPage, adjacent to met- even today, many of the working maps used by conservation
ropolitan Chicago in the state of Illinois, local citizens practitioners display urban areas as “nonhabitat,” devoid of
established a forest preserve system over 100 years ago to value for biodiversity conservation. In fact, most cities have
protect natural areas around their local communities, which areas within them that contain or could contain high levels of
were then still widely separated from one another by species richness (Kuhn et al. 2004) and high-quality habitat
unpreserved natural areas and agricultural lands. By 2010, for species of conservation concern (Gustafsson 2002; Kuhn
95 years later, the DuPage Forest Preserve included over et al. 2004). A first step toward true “urban conservation” is
60 preserves protecting 10,200 ha, about 12% (one eighth) the ability to recognize precisely what elements of an urban
of DuPage County (Fig. 3.22). But, by 2010, any land that environment can and should be included in conservation
was not secured within this preserved system was being strategies designed to protect biodiversity and develop con-
intensively used. Thus, although traditional urban conserva- servation programs sensitive and specific to a particular city
tion efforts were admirable, their traditional approach to (Parker 2015:685).
Fig. 3.22 Pratt’s Wayne Woods,

one of a network of forest
preserves within DuPage County,
Illinois (USA), bordering the
more intensely urban Cook
County, both part of the Greater
Chicago Area. By 2010, the
DuPage County Forest Preserve
system, started in 1915, included
over 60 preserves protecting
10,200 ha, about 12% (one eighth)
of the land area in DuPage
County. (Photo courtesy of the
Forest Preserve District of
DuPage County)
Urban habitat types can contribute to biodiversity conser-

vation of non-human species. But without effective biodiver- Information Box 3.2 (continued)
sity conservation and nature protection in cities, there will not
only be an extinction of species, but an accompanying
“extinction of experience” for human beings in their
interactions with nature. It is this second extinction that will
negatively affect the human perception of nature and the
number of people who pursue conservation as career and
vocation (Chap. 13). Although there is increasing need to
manage urban areas for biodiversity conservation, can such
efforts succeed in light of the problems and challenges cre-
ated in the urban landscape? Specifically, what particular
traits of the urban environment can be used to accomplish
conservation goals?
Information Box Fig. 3.2 Over one million Mexican free-

tailed bats (Tadarida brasiliensis) depart nightly from beneath
3.4.6 Changing Liability to Asset: the Anne W. Richards Congress Avenue Bridge over the
Incorporating People into Urban Guadalupe River in the US city of Austin, Texas (USA) from
Conservation March through October. (Photo courtesy of Ann Froschauer,
US Fish and Wildlife Service, public domain)
One of the ways that urban residents differ, on average, from
people residing in rural areas is the value they place on
non-human species. The late Stephen Kellert, in his This was not always the case. The origin of the enor-
pioneering work in assessing the ways in which people mous bat colony was an unintended consequence of a
value wildlife (Chap. 10), demonstrated that urban residents 1980 reconstruction that involved the addition of
tend to place higher value on species conservation than crevices on the underside of the bridge, structures
residents of rural areas (Kellert 1996) (Information Box which the bats immediately began to exploit. Initially,
3.2), and, not surprisingly, their legislators tend to be more the rapid increase in the number of bats caused con-
supportive of conservation legislation, such as strengthening cern, and some citizens petitioned to have them
the US Endangered Species Act, than legislators representing eradicated. At the invitation of US ecologist, conserva-
people from rural areas (Mehmood and Zhang 2001). These tionist, and wildlife photographer Merlin Tuttle,
potential advantages for biodiversity conservation, however, representatives from Bat Conservation International
may be offset, in part, by the fact that urban residents tend to (BCI), a conservation NGO specializing, as its name
have less knowledge about nature and species than rural suggests, in bat conservation, came to Austin. BCI
residents (McKinney 2002), and may be less directly affected began a campaign of public education about bats, con-
by conservation legislation and its enforcement. This fact vincing local residents that they were not only harmless
identifies one of the first problems to be solved in urban but fascinating creatures, who, for a living, would
biodiversity, and suggests an obvious remedy. Environmen- consume 10,000 to 20,000 pounds of insects every
tal and wildlife-oriented education will be vital to the success night during their evening foraging along the
of conservation efforts in the urban environment, as will Guadalupe, an action that was not only appreciated
intentional efforts to increase direct experience with and in by residents at an individual level, but as a welcome
nature for urban residents (Dearborn and Kark 2010). help to local agricultural producers.
Local citizens and government officials changed
their minds. The bats were spared and their population
Information Box 3.2: Austin Goes “Batty” Over
has grown to one and a half million individuals, the
Urban Biodiversity
largest urban bat colony in the world. Today over one
Each evening, from March through October, about
hundred thousand people visit Austin each year just to
20 min before sundown, over 1 million Mexican free-
see the bats make their spectacular evening departure.
tailed bats (Tadarida brasiliensis) emerge from
Austin’s newspaper, the Austin American-Statesman,
beneath the Anne W. Richards Congress Avenue
has created the Statesman Bat Observation Center adja-
Bridge spanning the Guadalupe River in the US city
cent to the Congress Bridge, giving visitors a dedicated
of Austin, Texas (Information Box Fig. 3.2).
area to view the nightly emergence. And the bats have
(continued)
(continued)
environmental knowledge in the process. Because urban

Information Box 3.2 (continued) residents tend to value wildlife more than rural residents,
now “spawned” their own cottage industry of business they often feel a sense of ethical obligation to “take care of”
specializing in “bat tours” to enhance bat viewing, natural areas and the animals that inhabit them (Dearborn and
including Austin Bat Tours, which takes tourists on Kark 2010). Given that people can form such an important
boats into the Guadalupe to be in just the right position element in urban biodiversity conservation efforts, what actu-
for an optimal view of this extraordinary event. Public- ally motivates people to become involved in conservation
ity about the bats has been so successful that Austin education and activity?
now has a bat statue, an annual bat festival (complete From a psychological standpoint, the most important qual-
with two stages of live music, over 50 arts and crafts ity necessary for involvement is commitment, which has been
vendors, and a bat costume contest), and, had, from defined as “a force that stabilizes individual behavior under
1996 to 2008, a minor league hockey team (the Austin circumstances where the individual would otherwise be
Ice Bats) with a bat mascot. tempted to change that behavior” (Asah and Blahna
Not every city has one and half-million bats that will 2013:867–868). Conservation scientists Stanley Asah and
make a nightly spectacle of themselves in places ideal Dale Blahna assessed this problem analytically by thinking
for public viewing. But the Austin example carefully about what functions are served in an individual’s
demonstrates that urban residents can display high life by changing their attitudes and behaviors. As they put it,
levels of interest in, support for, and even unique “success of efforts to change attitudes and behaviors depends
pride in biodiversity unique to their city if they are on the extent to which such efforts address the functions
taught to appreciate it and learn how to observe those attitudes and behaviors serve” (Asah and Blahna
it. Bats might not the first thing that comes to mind 2013:868). To better understand motivations and
when one thinks of a “flagship conservation species,” commitments, Asah and Blahna surveyed 322 urban conser-
but they are just that in Austin, and the pride of the vation volunteers and evaluated their responses to questions
entire community. about their motivation and commitment to determine how
volunteers structured motivations and commitment in rela-
tion to urban conservation activities. Affective commitment
related to those factors that motivated their emotional attach-
Motivations for biodiversity conservation in urban areas
ment to volunteering (“showing up” commitment). Norma-
are many (Fig. 3.23). Plants and animals in cities provide
tive commitment related to their ability to resist urges to avoid
ecosystem services, although, in urban areas, such services
volunteering (the strength of conviction that volunteering
might not be provided by native species (recall our example
was “the right thing to do”). Overall commitment related to
of the novel forests in Hawaiian ecosystems, which actually
their long term motivation to continue to volunteer. In terms
provided high levels of some ecosystem services than native
of “behavioral function,” volunteers identified functions of
forests). Plants, for example, may provide temperature
their volunteer activities as motivated by desires to help the
moderation and carbon sequestration (Chap. 4). Plants and
environment, defend and enhance the ego, offer career and
animals in urban areas, especially if incorporated into well
learning opportunities, provide “escape” and exercise,
designed environmental education programs, can help people
experience connection to nature, and enhance their level of
Fig. 3.23 Some important

motivations for biodiversity
conservation in urban areas,
which range in relative
importance from those that
primarily benefit nature
(ascending arrow) to those that
primarily benefit humans
(descending arrow). (Reprinted by
from Dearborn, D.C., Kark, S.,
2010. Motivations for Conserving
Urban Biodiversity. Conservation
Biology 24, 432–440. # 2009
Society for Conservation Biology)
facilitate social interactions, and provide opportunity for conservation behaviors, even if the motivating influences are
community building. something other than saving species. As Asah and Blahna
The key motivators identified by the volunteers were not summarize, “Conservation initiatives are increasingly depen-
what conservationists might expect, or, perhaps, want to hear. dent on volunteers to accomplish goals. Traditionally, con-
“The motivation to help the environment,” noted Asah and servation practitioners used environmental problems to incite
Blahna, “was a marginally significant predictor of affective people’s involvement in conservation initiatives.
commitment and did not significantly predict normative and Practitioners could be more effective and efficient if they
overall commitments. . .[in contrast] The more volunteers were to use personal and social functions, especially commu-
wanted to socialize with others, the more they were emotion- nity motivations, as a means to secure volunteers’ commit-
ally attached to, identified with, and got involved with ment to conservation issues. These personal, social, and
volunteering (affective commitment) and were likely to con- community motivations could be planned to occur during,
tinue to volunteer under circumstances they would otherwise or related to, voluntary conservation efforts” (Asah and
be tempted not to do so (overall commitment). And the more Blahna 2013:873).
volunteers wanted to build and enhance community, the more
they affectively committed to volunteering, felt obligated to
volunteer (normative commitment), and had overall commit- 3.4.7 How to Do It – Six Strategies
ment to volunteer. The more volunteers wanted to feel less for Conservation Practitioners in Urban
guilty about harm humans cause to the environment (protect Areas
the ego against negative features of the self) and make a
difference in that respect, the more they were normatively As urban ecologist Ivan Kowarik noted, “Existing conserva-
committed to volunteer” (Asah and Blahna 2013:871–872). tion strategies mostly aim at preserving native ecosystems by
In other words, volunteers were more motivated to engage curbing urban growth and resulting habitat losses, preserving
in urban conservation activities in order to socialize with more or less transformed relicts of natural habitats within
others and “feel better” about themselves than they were to cities or restoring native species in urban habitats. . .most
engage in the activities to benefit the environment itself, or papers on urban conservation approaches appear to be biased
the creatures that comprised its biodiversity. Although to some extent by conservation, or restoration, efforts that
conservationists might view these findings with discourage- prioritize relicts of pristine (“native”) ecosystems or native
ment, they provide an important insight into how to recruit species in urban regions and tend to neglect other urban
and retain people for conservation activities. As Asah and systems” (Kowarik 2011:1979).
Blahna put it, “. . .conventional management structures [such The distinctiveness of an urban landscape with respect to
as a conservation agency or organization might impose on or conservation is composed of (1) its bio/geo/ecological realm
prescribe for a volunteer event] may hinder volunteer engage- or “natural” environment, (2) its human realm, and (3) its
ment with environmental causes . . .Volunteer events are built environment. Conservationists have traditionally
often planned and managed in ways that ineffectively match focused on the first realm, but the second and third categories
volunteers’ most salient motivations (i.e., they provide little provide opportunities for biodiversity conservation that may
or no explicit socially interactive and community-building be even more important, particularly with the help of conser-
activities). Thus, it is likely that difficulties retaining vation social scientists and ecological and environmental
volunteers is partially explained by conventional manage- engineers. Sophie Parker of The Nature Conservancy offers
ment practices” (Asah and Blahna 2013:873). six strategies for a manager working to achieve conservation
A second insight is also latent in these findings. It is that goals in an urban environment (Table 3.5). (1) Focus on
education about conservation is not, in itself, sufficient to Adaptive Management – This is good advice in any conser-
produce engagement with conservation. As we will see vation context, but is particularly appropriate in an urban
throughout the examples used in every chapter of this book, environment in which land use, land ownership, and environ-
conservation is not advanced solely or even primarily by mental conditions can and do change rapidly, and new con-
increasing knowledge or better science. It is advanced only servation efforts are unlikely to be initiated unless previous
by changes in human behavior. And motivation is the most conservation efforts have demonstrated that the conservation
important factor in behavioral change. Therefore, as Asah strategy is achieving the desired goals. An adaptive manage-
and Blahna point out, “. . .calls for education as the means to ment approach is vital in an urban setting because a city’s
conservation ends are somewhat misplaced” (Asah and built environment and high human densities may limit the
Blahna 2013:874). The best strategy, particularly in an array of management actions more than in a rural area, park
urban setting in which people are accustomed to a high or preserve. (2) Recognize the Urban Environment as an
density and frequency of social interaction, is to Ecological System – Downtown Los Angeles (California,
empahsize activities and events that will motivate positive USA) may not look like an ecosystem to a biologist trained
Table 3.5 Six strategies for conservation practitioners working in urban areas
Strategy Explanation
Employ adaptive management Use a robust conservation planning framework that allows for continual refinement of conservation
strategies
Embrace the ecosystems concept When defining the scope of an urban conservation planning area, pay special attention to ecological pattern/
process
Abandon the gray/green Incorporate gradations in the suitability of urban habitat for various species into project maps
dichotomy
Use socioeconomic/cultural Use pre-existing surveys of human communities’ perceptions of nature to help guide implementation and
factors in planning educational outreach
Understand and engage with Strong ties with government, NGOs, and the public can go far towards advancing conservation strategies
power
Make use of technology Collection of some data can be easier in cities because of the ubiquitous presence of mobile
telecommunications applications and citizen science
Reprinted by permission from Springer Nature, from Parker, S.S., 2015. Incorporating critical elements of city distinctiveness into urban biodiversity
conservation. Biodiversity Conservation 24, 683–700. # 2014 Springer Science+Business Media Dordrecht
in Yellowstone National Park, but it is one just the same. associated with permitted land use and zoning restrictions.
Matter and energy are being processed, transported and (6) Make Use of Technology and Comprehensive Data
cycled in a myriad of ways, but in a fundamentally similar Sources – High levels of remote sensing technology, mobile
manner as in non-urban environments. Non-human species networks and sophisticated analytical software are part of the
are present (if not always apparent) and interacting with their cultural resources of an urban environment, and their integra-
environment, and climatic and hydrological regimes are tion in planning is almost always necessary to convince city
affecting environmental conditions. An urban environment officials and municipal planners that the conservation strat-
is particularly vulnerable to outside inputs of energy and egy is workable and well-conceived. But any technology is
matter, even (perhaps especially) energy and matter that only as good as the data it collects and analyzes. Urban areas,
enters the system from far distant sources. Think beyond because they are invariably rich in cultural and administrative
the immediate surroundings when planning the conservation resources, often provide high accessibility to detailed, high
strategy. (3) Abandon the Gray/Green Dichotomy – See the quality datasets that are needed to inform conservation
urban landscape as a gradation of environments from the planning (Table 3.6).
most natural areas to the most developed, but not as only Conservation matters everywhere. Conservation in an
two categories (natural versus developed) in which all con- urban context matters now more than ever, and conservation
servation effort goes into the first category and none in the must relate to life experience. If all of a person’s life
second. Integrate conservation and development strategies in experiences have occurred within an urban environment,
the same places and in the same projects. (4) Use socioeco- they cannot and will not have the experience of conservation
nomic and cultural factors in planning – No conservation of nature except and unless it takes place in an urban envi-
effort anywhere is likely to succeed without local support, ronment. As James Miller and Richard Hobbs note, “From
and this is particularly true in an urban landscape. Ownership the perspective of someone who lives in a city or suburb,
of land is private. Individual holdings are small. No “mega- conservation is too often something that happens somewhere
landowner,” such as a government agency, can dictate con- else—in a national park, wilderness area, or rainforest—and
servation policy. Local support is needed, especially in the is experienced second-hand (if at all) on television or in a
form of volunteers, But local residents will only support magazine. . .” (Miller and Hobbs 2002:334). As Aldo
conservation values they actually perceive through conserva- Leopold observed, “conservation is not merely a thing to be
tion plans and strategies which they help to create. They will enshrined in outdoor museums, but a way of living on the
be involved in the actual work of conservation only to the land” (Meine 1988:310), even if the land is encased in pave-
extent that their own commitments and motivations make it ment. “Like all real treasures of the mind,” wrote Leopold,
reasonable for them to invest in that work. (5) Understand “perception can be split into infinitely small fractions without
and Engage with Power – Although city governments will losing its quality. The weeds in a city lot convey the same
not have the same breadth of jurisdiction as federal lesson as the redwoods” (Leopold 1949:174). From now on,
administrations, their endorsement, sponsorship, and finan- conservationists must perceive such weeds, and city lots,
cial support, along with local conservation organizations, will with the same insight and perception they have formerly
legitimize urban conservation efforts and help clear obstacles devoted to redwoods and wildlands. They must make urban
3.5 Whither the Anthropocene? What Strategy Creates a Place for Nature? 117
Table 3.6 Examples of data available from various sources relevant for incorporation into urban biodiversity conservation efforts
Data category Data description Example sources
Natural history, biological, and ecological data Occurrence data for species State Natural Heritage Programs
Soils and geology maps USDA Soil Survey
Historical ecology studies San Francisco Estuary Institute
Footprint maps The Nature Conservancy
Citizen science efforts eBird
People Demographic data US Census Bureau
Built environment Housing density, zoning, infrastructure maps County planning departments
Reprinted by permission from Springer Nature, from Parker, S.S., 2015. Incorporating critical elements of city distinctiveness into urban biodiversity
conservation. Biodiversity and Conservation 24, 683–700. # 2014 Springer Science+Business Media Dordrecht
environments places where nature flourishes and where peo- systems: climate change, ocean acidification, stratospheric
ple understand and enjoy its flourishing. That must be ozone depletion, nitrogen-phosphorus cycle, global freshwa-
conservation’s ideal for the modern city. ter use, change in land use, biodiversity loss, atmospheric
aerosol loading, and chemical pollution (Fig. 3.24)
(Rockström et al. 2009) with prescribed boundaries for each
3.5 Whither the Anthropocene? What system, some of which they judge to have already been
Strategy Creates a Place for Nature? crossed. Regarding climate change and associated
CO2 emissions, the authors assert, “We propose that human
3.5.1 The Emergence of “Neoprotectionism” changes to atmospheric CO2 concentrations should not
exceed 350 parts per million by volume, and that radiative
Cities are arguably the most “novel” of all ecosystems, and forcing should not exceed 1 watt per square meter above
the most anthropogenic. But the urban ecological system is pre-industrial levels. Transgressing these boundaries will
but one of many signatures of humanity on the Earth, and increase the risk of irreversible climate change, such as the
arguably not the most extensive or pervasive. That one spe- loss of major ice sheets, accelerated sea level rise and abrupt
cies should make such a mark on the planet as to be shifts in forest and agricultural systems.” (Rockström et al.
immortalized in its own geologic epoch might be considered 2009). As of 2019, the atmospheric CO2 concentration was
a great achievement. But when we look at what humanity has 414 ppm and the change in radiative forcing was 2.29 W/m2
done to nature to reach this point, it is nothing to be proud (NOAA 2020), making Rockström et al.’s warnings pro-
of. The ravaging damage that has characterized the human phetic. “We do not have the luxury of concentrating our
presence in the last 500 years, and particularly in the last efforts on any one of them in isolation from the others. If
70, has given us the Extinction Crisis, Global Climate one boundary is transgressed, then other boundaries are also
Change, and Tropical Deforestation, to name a select few. under serious risk” (Rockström et al. 2009). If such warnings
To this point, humanity has produced an irresponsible are warranted, how will humanity stay within its “safe
Anthropocene. The question is, can there be a responsible operating space”? And, of particular concern to conservation
Anthropocene? science, how will it preserve the Earth’s biodiversity?
Johan Rockström of Stockholm University (Sweden) and One comprehensive proposal is offered by the eminent
his colleagues crystalized such concern, and a possible biologist E. O. Wilson (Fig. 3.25). That solution is what
response to it, in their landmark paper, “A Safe Operating Wilson has called the “Half Earth,” proposal, a plan detailed
Space for Humanity (Rockström et al. 2009). They begin in his Pulitzer Prize-winning book, Half-Earth: Our Planet’s
with dire but warranted concern. “Since the Industrial Revo- Fight for Life. Wilson also believes, like many, that the
lution, a new era has arisen, the Anthropocene, in which Anthropocene is a present global reality. Its problems, there-
human actions have become the main driver of global envi- fore, must be addressed through a global strategy. His solu-
ronmental change. This could see human activities push the tion: dedicate one half of the Earth to nature and natural
Earth system outside the stable environmental state of the processes. Humanity would carry on its life within the “safe
Holocene, with consequences that are detrimental or even operating zone” of the other half. Using a rough, but well-
catastrophic for large parts of the world. . .The result could be established rule of thumb to express the relationship between
irreversible and, in some cases, abrupt environmental change, percent habitat preserved and percent species preserved, as
leading to a state less conducive to human development” observed at smaller scales, Wilson estimates that the preser-
(Rockström et al. 2009:472). In response to Earth’s current vation of 50% of the Earth’s land area in a natural state would
condition, they propose a “safe operating space” for human- preserve 84% of its species (Wilson 2016a). We have already
ity, defined by criteria associated with nine planetary passed this point if we are talking about preserving wildlands,
Fig. 3.24 A representation of nine critical Earth planetary systems and Lambin, E.F., Lenton, T.M., Scheffer, M., Folke, C., Schellnhuber, H.J.,
a “safe operating space” (green region) for humanity within those Nykvist, B., de Wit, C.A., Hughes, T., van der Leeuw, S., Rodhe, H.,
systems. Red wedges represent estimation of current position of Sörlin, S., Snyder, P.K., Costanza, R., Svedin, U., Falkenmark, M.,
human effect in each system, with three systems (biodiversity loss, Karlberg, L., Corell, R.W., Fabry, V.J., Hansen, J., Walker, B.,
climate change, and the nitrogen cycle) already beyond safe boundaries Liverman, D., Richardson, K., Crutzen, P., Foley, J.A., 2009. A safe
in human effect. (Reprinted by permission from Springer Nature, operating space for humanity. Nature 461, 472–475. # 2009
from Rockström, J., Steffen, W., Noone, K., Persson, Å., Chapin, F.S., Macmillan Publishers Limited)
which now comprise less than 25% of the Earth’s ice free
surface (Sect. 3.1.2), so we must take care in how we would
define nature’s “natural state” and the role of human agency
that in creating that state. Wilson’s tactic to preserve nature’s
natural state would be to increase the size of existing nature
refuges. “The only way to save upward of 90% of the rest of
life is to vastly increase the area of refuges, from their current
15% of the land and 3% of the sea to half of the land and half
of the sea. That amount, as I and others have shown, can be
put together from large and small fragments around the world
to remain relatively natural, without removing people living
there or changing property rights. This method has been
tested on a much smaller scale at the national and state park
levels within the United States” (Wilson 2016b).
Wilson’s plan, and variations of it, have been described as
strategies of neoprotectionism – new forms and variations of
the historical approach to conservation as it originated in the
Fig. 3.25 Using a common rule-of-thumb for the relationship between late nineteenth century in North America (Chap. 1) and
habitat area and the number of species it can sustain in the long term, developed in various forms through the twentieth century
Harvard biologist E. O. Wilson has proposed setting aside half of the worldwide. Like traditional protectionism, neoprotectionism
Earth to preserve 84% of Earth’s current species. (Photo courtesy of
J. Harrison, under CC BY 2.5)
operates on a fundamental premise that conservation is a way
to protect nature from people. One emerging alternative own lives. In summary, we are advocating conservation for
viewpoint is what some have called “natural capital” people rather than from people” (Kareiva and Marvier
conservation – the goal of which is protecting nature for 2012:968). Kareiva and Marvier advocate a vision of conser-
people. vation that is cooperative with capitalism and development.
“Conservation should seek to support and inform the right of
development – development by design, done with the impor-
3.5.2 Conservation as Human and Economic tance of nature to thriving economies foremost in
Development mind. . .Instead of scolding capitalism, conservationists
should partner with corporations in a science-based effort to
Various forms of neoprotectionism, endorsed by many integrate the value of nature’s benefits to their operations and
conservationists, are also criticized and even ridiculed by cultures” (Marvier et al. 2019).
others. Although the critics, and their criticisms, vary, the
largest group might be called the “natural capital” conserva-
tionists, the most vocal and well known being Peter Kareiva, 3.5.3 “Convivial” Conservation – Local
Chief Scientist and Vice President of The Nature Conser- Autonomy for Local Benefit
vancy, and Michelle Marvier, Professor of Environmental
Studies and Sciences of Santa Clara University (USA). Perhaps the most fundamental difference between the
They argue that what Wilson proposes to save the Earth’s neoprotectionist approach exemplified by Wilson and the
biodiversity has already proven to be a failed strategy. Put “new conservation” espoused by Kareiva and Marvier is
bluntly, “. . .conservation cannot promise a return to the pris- their view of humans as drivers of ecological and environ-
tine, prehuman landscapes. Humankind has already pro- mental change. New conservationists see human-driven
foundly transformed the planet and will continue to do change as potentially positive. Neoprotectionists do not.
so. What conservation could promise instead is a new vision They see it instead as the primary threat. And, based on the
of a planet in which nature – forests, wetlands, diverse human track record, that view is not without warrant. Not all
species, and other ancient ecosystems – exists amid a wide conservationists endorse the new conservation’s repudiation
variety of modern human landscapes. For this to happen, of conservation’s historic values, the apparent equation of
conservationists will have to jettison their idealized notion biodiversity with “natural capital,” or the sometimes uncriti-
of nature, parks, and wilderness – ideas that have never been cal acceptance new conservationists at times appear to give to
supported by good conservation science – and forge a more development as the best pathway to saving nature. One
optimistic human-friendly vision” (Marvier et al. 2019). school of thought, differing from both, is the paradigm of
Kareiva and Marvier consider the “half Earth” approach “convivial conservation,” (convivial – adjective – friendly,
not only outdated but insensitive to human needs and human lively, enjoyable) most prominently advocated by Bram
rights. Marvier and Kareiva disparage what they see as the Büscher and Robert Fletcher of Wageningen University
traditional approach advocated by Wilson and others. “By (The Netherlands). Büscher and Fletcher argue that neither
pitting people against nature, conservationists actually create the “traditional” nor “new” conservation strategies take polit-
an atmosphere in which people see nature as the enemy. If ical and economic realities of the present time seriously
people don’t believe conservation is in their own best enough to succeed. “The crucial difference,” assert Büscher
interests, then it will never be a societal priority. Conserva- and Fletcher, “. . .is that we explicitly start from a political
tion must demonstrate how the fates of nature and of people ecology perspective steeped in a critique of capitalist political
are deeply intertwined – and then offer strategies for promot- economy. . . This critique is built on a rejection of both
ing the health and prosperity of both” (Marvier et al. 2019). nature-people dichotomies and a capitalist economic system
As such, Kareiva and Marvier see effective future conserva- demanding continual growth via intensified consumerism”
tion as an expression of enlightened human social and eco- (Büscher and Fletcher 2019). Like new conservation,
nomic development. They advocate a view of conservation Büscher and Fletcher insist that conservation should shift its
that emphasizes human rights on equal or greater footing with focus away from protecting nature from people. Unlike new
nature’s rights. “Forward-looking conservation protects nat- conservation, they advocate promoting nature “for, to and by
ural habitats where people live and extract resources and humans” (Büscher and Fletcher 2019) in non-capitalist terms,
works with corporations to find mixes of economic and not marketing them for potential capital accumulation and
conservation activities that blend development with a con- exploitation (ecotourism or natural capital), but rather as
cern for nature. . .Our vision of conservation science differs “promoted areas” where people “are considered welcome
from earlier framings of conservation biology in large part visitors, dwellers or travelers rather than temporary alien
because we believe that nature can prosper so long as people invaders upon a nonhuman landscape” (Büscher and Fletcher
see conservation as something that sustains and enriches their 2019). And from this perspective their paradigm derives its
Fig. 3.26 One example of an

Indigenous Community and
Conservation Area (ICCA) can be
seen in a recent concession of
112,850 acres (45,699 hectares) of
remote, intact lowland jungle to
indigenous tribes of Peru’s Yurua
River. The concession of land
establishing this ICCA will be
managed by the new Asociación
de Conservación Comunal Yurúa,
which is comprised of local men
and women from the Ashéninka,
Asháninka, Yaminahua and
Amahuaca tribes. The area
harbors many rare species and
now protects tribal homelands
from logging, ranching,
commercial agriculture and other
causes of deforestation which
have become prevalent in the
western Amazon. (Photo courtesy
of Jason Houston/Upper Amazon
Conservancy)
name – a conservation based on conviviality, “the building of further advocate a “conservation basic income” (CBI) to local
long-lasting, engaging and open-ended relationships with people which would consist “monetary payment to individual
nonhumans and ecologies” (Büscher and Fletcher 2019). community members living in or around promoted areas that
They advocate that such “promoted areas,” indeed all areas, allows them to lead a (locally defined) decent life (Bücher
would be managed, not by government agencies or national and Fletcher 2019). Finally, Bücher and Fletcher advocate an
or international conservation organizations, but by local ideological and practical separation of conservation from
residents who would govern such spaces as “Indigenous capitalism, meaning that, in terms of engagement between
and Community Conservation Areas” or ICCAs, which conservation organizations and corporate sponsorship of con-
have been defined by the IUCN as natural and/or modified servation projects, “such engagement should proceed only
ecosystems containing significant biodiversity values and under strict conditions. One of these is that conservation
ecological services, voluntarily conserved by (sedentary NGOs should only work with companies if the latter pledge
and mobile) indigenous and local communities, through cus- that they understand the necessity of moving towards a dif-
tomary laws or other effective means (IUCN 2010) ferent economic model beyond capitalist accumulation and
(Fig. 3.26). As Bücher and Fletcher put it, “Under convivial GDP-based economic growth” (Bücher and Fletcher 2019).
conservation, the emphasis will be on long-term democratic Certainly no one can accuse Bücher and Fletcher or other
engagement rather than on short-term voyeuristic tourism or convivial conservationists of a lack of originality. But how-
elite access and privilege. . . we cannot afford to continue ever effective such envisioned changes might be in producing
flying around the world in climate-changing airplanes in conviviality between humans and nature, these ideas, as
order to save nature through (eco)tourism..[we must rather] recommendations for action, will not produce any such
encourage long-term visitation focused on social and ecolog- thing without further discussion, especially since Bücher
ical justice. . .in relation to the natures close(r) to where we and Fletcher support monetary payments to create a CBI,
live” (Bücher and Fletcher 2019). bringing up the level of “decent life” for local peoples,
In the same vein, convivial conservation emphasizes while at the same time advocating cutting ties with corporate
“everyday nature” and “local nature” rather than “nature as sponsors, which might be one of the few sources with the
spectacle.” It advocates that local people, rather than conser- money to fund such a plan. And, to make local residents the
vation organizations or government authorities, be the key sole or primary authority for management of conservation
conservation decision makers. But Bücher and Fletcher go areas would mean that local considerations would receive
further. They want “historic reparations” of land to indige- priority in the decision making equation, when in fact many
nous people in which they “receive (access to) their land back conservation areas and protected refuges, such as national
or at the very least obtain co-ownership or of co-management parks, have been established to reflect national or interna-
responsibilities over it” (Bücher and Fletcher 2019). They tional priorities which were not important to local residents.
preserving highly sensitive areas from most or all forms of

Point of Engagement Question human impact, they also work to engage people in the
Can you envision what government agency or conser- enhancement of biodiversity where they live, and in harmony
vation organization might have the desire and the finan- with a sustainable economic system, subjects and examples
cial resources to provide a “conservation basic income” we will explore in detail in Chap. 11. And, among many of
(CBI) to local residents living in and around one of the the most effective conservationists today, there is active and
“promoted areas” that Bücher and Fletcher describe? diligent effort to foster an ethical awareness and framework
Do you think that there is a possibility of unintended that creates genuine affection and concern for nature, with
consequences and perverse incentives arising from appropriate restraint of the kinds of human influence and
CBIs if such payments motivated more people to development that could destroy it (Chap. 10).
move into or adjacent to designated conservation
areas, increasing human density and human use in
Synthesis
and around environmentally sensitive areas? What
The wildlands of the Earth still teem with beautiful,
might be the advantages of such an approach?
varied, and fierce life. For those who have never beheld
it, such life is beyond the power of words to describe or
3.5.4 Can Different Approaches Find photography to depict. You must see it for yourself, for
Reconciliation or Resolution? it possesses power that deserves the dignity of direct
contact, meeting such life where its power resides. For
Whatever one may think about the merits of convivial con- those who have seen it, known it, even lived among
servation, neoprotectionism, natural capital conservation, or such creatures who possess it, sharing the power of
current mainstream conservation efforts, this is not a time of their lives in small ways and varied measures, such
conviviality among conservationists. It is a time of contro- life is a constant inspiration that can never diminish
versy, criticism, crisis and potentially increasing confusion. or fade in the story that it tells or the effort it sustains to
Mainstream conservation faces the challenges, and legitimate preserve and protect it. And such life must not perish
charges, that increasing the number and size of protected from the Earth.
areas has not produced effective results in reducing global But many of the Earth’s wild places have been
biodiversity loss, as does neoprotectionism. diminished, and much of their life has perished. The
Neoprotectionism and convivial conservation face significant recognition that we are now living in the Anthropocene
problems in implementing their most important (whether geologists ultimately choose to give our time
recommendations, although for different reasons. Some this name or not) is not simply another variant to the
forms of natural capital conservation, or conservation as ongoing story and struggle of conservation as we have
development, face the difficulty of redefining the objectives always known it. It is, and it requires, a fundamental
of traditional approaches to development, framing conserva- transfiguration of conservation strategy, conservation
tion primarily as advancement to human wellbeing and eco- targets, and, in some cases, conservation purposes. The
nomic self-interest (Chap. 11). Unless the conservation vast majority of human inhabitants on the Earth today
community can effectively integrate or resolve the immense will never know wildlands or the life they hold in any
differences in these approaches, it will not be able to send a meaningful or interactive way. The urban areas we
clear message to the human community about the values, have created for our own convenience must now them-
purposes, and goals of conservation efforts, or the appropri- selves become the areas to which we “bring back”
ate strategies for attaining them. nature and natural life, the biodiversity of the nonhu-
All of these approaches desire to create what we might man world. This cannot be done simply by a reconcili-
describe as a “Responsible Anthropocence.” But now, in an ation of compromise. It can be accomplished only by a
age in which human populations and technology create reconciliation of repentance. Human needs and devel-
effects that make them the equivalent of geological, climato- opment must now always be an integral part of conser-
logical, and ecological forces, the absence of a coherent vation effort, but meeting such needs and fostering
conservation strategy and the confusion of a divided conser- such development must never become the sole purpose
vation community could produce disaster for global biodiver- of conservation effort. The non-human life of the world
sity. There are many active conservationists today who, is more than natural capital. Our repentance lies in this
although not subscribing entirely to any of these or other recognition, and only in that recognition can there be
“camps” of conservation opinion, are nevertheless changing true reconciliation with nature. As modern
their approach in light of the realities of doing conservation in
an age of human-dominated nature. Recognizing the value of (continued)
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Biodiversity Conservation and Climate Change
4
Are we at the failsafe point? No. We still have time to act upon the recognition that our planet is an intricately
linked biological and physical system that holds yet-to-be-understood capacity to heal and clean itself. We still
have tools and opportunities to effectively manage the living planet and its biodiversity for the benefit of
humanity and all life on Earth.
Thomas Lovejoy and Lee Hannah (2018)
Keywords 4.1 Climate and Climate Change

Climate fingerprints · Phenological changes · Range
shifts · Climate-driven extinctions · Integrated vulnerabil- 4.1.1 Why Does Climate Change Threaten
ity assessments · Climate velocity · Bioclimate envelope · Biodiversity?
Dynamic Global Vegetation Model (DGVM) · Climate-
informed conservation strategies · Paris Agreement “Climate change represents a fundamentally different threat
to Canada’s national park system. Never before has there
been an ecological stressor that raised questions about the
adequacy of the system to protect representative samples of
Overview Canadian ecosystems” (Scott 2005: 344). In these remarks,
In this chapter, you will learn about: Daniel Scott, professor at the University of Waterloo, is
addressing the effects of contemporary climate change on
1. The scientific evidence for anthropogenic climate only one nation, Canada, and only one dimension of its
change. national conservation effort, national parks. However, Scott
2. The documented effects of climate change on could just as easily have been speaking of every global
biodiversity. conservation strategy. As you learned in Chap. 3, the most
3. The modeling tools used to project future climate- important threats to global biodiversity have traditionally
ecosystem interactions. been perceived as habitat destruction, overexploitation of
4. Climate-informed conservation strategies. species, and the competition, predation, and displacement
5. Policies to mitigate the effects of climate change and of indigenous species by invasive species. These threats
support biodiversity conservation. remain obstacles to biodiversity conservation. However,
superimposed as both an additional threat and an intensifier
of existing threats, global climate change has the potential to
overwhelm existing local or regional actions, however well
designed or intended, toward conservation goals.
It is important to note that the relationship between climate
and species, particularly species distribution is well
established. In 1917, the American zoologist Joseph Grinnell,
examining the Oregon jay (Perisoreus canadensis obscurus),
pika (Ochotona princeps), rosy finch (Leucosticte
tephrocotis), redwood chipmunk (Neotamias ochrogenys),

126 4 Biodiversity Conservation and Climate Change
and western meadowlark (Sturnella neglecta) (Fig. 4.1), of the past climate events and is actively challenging the
published his assessment of the role of climate in effectiveness of twentieth century conservation.
constraining the geographic ranges of these animals (Grinnell Climate change is, as Daniel Scott put it, a “fundamentally
1917). Grinnell did not call his work a “climate study,” but different threat,” an “ecological stressor” that, unlike any
noted, in his own words that “. . .upon the North American other, raises questions about the adequacy of any conserva-
continent, one single factor does happen to loom up as being tion strategy or reserve system to protect and preserve repre-
the most frequent delimiter of distribution, or even the ulti- sentative biodiversity. The reason that such a description is
mately effective one, in greater or lesser degree, even though warranted is because climate change is capable of altering the
other factors be effective also. This factor is temperature” very nature of what is being preserved. As Scott later stated in
(Grinnell 1917:128). Grinnell considered temperature a lim- the context of addressing the problem climate change
iting factor in species distribution, but not a changing factor. presents to Canadian national parks, “. . .the stated purpose
He, like other naturalists of that day, considered temperature of Prince Albert National Park is to ‘protect for all time the
and other climate factors to be stable and consistent, however ecological integrity of a natural area of Canadian significance
restrictive they might be to a particular species. representative of the southern boreal forest and plateau. . .’
In 1992, ecologist Peter Vitousek wrote in his introduction Yet all six vegetation change scenarios examined. . .
to that year’s Annual Review of Ecology, Evolution, and projected the eventual loss of boreal forest in this park. . .,
Systematics special issue on “Global Environmental suggesting that the park’s mandate would be untenable in the
Change,” that “climate change probably has the greatest long term” (Scott 2005:343).
potential to alter the functioning of the Earth system; its direct One survival strategy of species is to move, tracking their
effects on natural and managed systems ultimately could “climate niche” as it changes across space and over time. But,
become overwhelming. . .nevertheless, the major effects of in a world of increasing habitat destruction, fragmentation,
climate change are mostly in the future while most of the and isolation, movement to new areas is often impeded. Thus,
others are already with us” (Vitousek 1992:7). Vitousek’s acting in concert with habitat destruction, climate change can
statement might still be true, but climate change is no longer a create synergistic stresses that can lead to species extinction.
future scenario, nor are its effects. Climate change is here, it’s But climate change, even in the absence of direct habitat
because of us, and it’s serious (Molina et al. 2014). Although destruction, can change the surrounding environmental
we can learn from the impacts of past climate changes on conditions of a species so quickly that the species cannot
species distribution and migration (e.g., Huntley and Webb adapt in time. Thus, climate change can lead species down a
1989), the speed of contemporary changes is faster than most second pathway of extinction by exceeding their present
environmental tolerances, altering their environment more
Fig. 4.1 Some of the species the

early twentieth century
US naturalist Joseph Grinnell
assessed in relation to the role of
climate factors in their distribution
included the Oregon jay
(Perisoreus canadensis obscurus,
A), western meadowlark
(Sturnella neglecta, B), pika
(Ochotona princeps, C), and rosy
finch (Leucosticte arctoa, D). (Jay
image courtesy of Walter
Siegmund, under CC BY-SA 3.0.
Meadowlark image courtesy of
Missouri Department of
Conservation. Pika photo courtesy
of Will Thompson/USGS, public
domain. Jay photo courtesy of
Alan Wilson/Nature Pics Online,
under CC BY-SA 3.0)
4.1 Climate and Climate Change 127
quickly than their genetic resources can adapt to it. On their

own, species face three realities when faced with climate
change: disperse, adapt, or die. As we will discuss later in
this chapter, conservation biologists are often faced with
three choices in response: do nothing, continue conventional
conservation approaches, or create new and hybrid conserva-
tion strategies to account for specific climate changes and
their effects.
4.1.2 What Is “Climate” and What Is “Climate

Change”?
A dictionary would define “climate” as “the average course

or condition of the weather at a place over a period of years”
(Webster 1971:155). Thus, the first important distinction
when examining climate change is the distinction between
weather and climate. “Weather” refers to local, short-term
changes in variables such as temperature, precipitation, wind Fig. 4.2 (a) Milankovitch cycles are examples of regular periodic
speed, and humidity. Weather is constantly changing, but variation in climate. The periodicity of Milankovitch cycles is due to
changing within a range of historical norms, cycles or longer term (22,000 year), regular variations in the Earth’s distance from
the sun. (b) In contrast, discontinuities, jumps, trends, or increasing
fluctuations that, in the long term, are highly predictable.
variability can all represent different forms of climate change. In these
These long-term predictable fluctuations are what is meant cases, the means of climatic variables do not remain stationary within
by “climate.” the time domain of interest. After the jump or discontinuity, a new
Variations in climate, over the long history of the Earth, baseline of climate variability emerges that is different from the old.
(c) Increasing variability occurs when the means of climatic variables
generally take one of four overall forms (Fig. 4.2; Karl and
remain the same, the but their range of variation increases. Now individ-
Trenberth 2005). Regular periodic variation in climate is a ual “weather” events become more unpredictable and less tightly clus-
kind of predictable change manifested in annual seasonal tered around historic means. (d) Trends in climate refer to changes in
cycles and in longer-term regular fluctuations like the which the range of variation of climatic variables remains more or less
within historical norms, but there is a movement of the mean in a single
Milankovitch cycles which show periodic, regular variations
direction, consistently greater or less than the historic average. When
in temperature on the Earth over periods of thousands of such consistent, sustained movement occurs, it is appropriate to speak of
years. The periodicity of annual variation is due to the tilt such variation as climate change. (Reprinted by permission from Yale
of the Earth’s axis relative to the axis of the sun, such that the University Press, from T. R. Karl and K. E. Trenberth, “What is Climate
Change,” in Climate Change and Biodiversity, edited by Thomas
directness and intensity of light changes at different points in
E. Lovejoy and Lee Jay Hannah, 15–30. New Haven: Yale University
the Earth’s orbit, due to a change in the Earth’s inclination Press. # 2005 Yale University Press)
between 22.1 and 24.5 . The periodicity of Milankovitch
cycles, in contrast, is due to longer term, but regular 22,000-
year cycles caused by variations in the Earth’s distance from
the sun. Such changes, although capable of producing great climate variables, relative to the length of the temporal cycle,
differences in climatic variables in both space and time, are show a large change in magnitude over relatively short time
called “climate variation,” not “climate change.” Within such spans. After the jump or discontinuity, a new baseline
cycles, long-term averages of key climatic variables, such as emerges that is different from the old one. Increasing
temperature and precipitation, remain the same. Studying variability occurs when the means of climatic variables
climate over long time scales, scientists have been able to remain the same, but their range of variation increases. In
establish foundational relationships between changes in these cases, individual “weather” events become more unpre-
external forcing, such as changes in the sun’s strength and dictable and less tightly clustered around historic means.
how this change in forcing influences climate system Trends in climate refer to changes in which the range of
interactions, such as between atmosphere and ocean, and variation remains within historical norms, but with move-
results in an observable changes within each system ment of the mean in one direction, consistently greater or
component. less than the historic average. When such sustained direc-
In contrast, discontinuities, jumps, trends, or increasing tional movement occurs, such variation is called climate
variability can all represent forms of “climate change.” In change (Information Box 4.1).
these cases, means of climate variables do not remain station-
ary. Climate discontinuities and climate jumps occur when
Information Box 4.1: Protecting National Parks Information Box 4.1 (continued)
Outside of Historical Range of Variability generally falling within historical and multi-decadal
To determine the presence of a sustained climate trend, ranges (normal range of variability). Looking at
senior scientists William B. Monahan and Nicholas monthly averages of daily temperature and precipita-
A. Fisichelli, of the US National Park Service studied tion, percentage cloud cover, vapor pressure, as well as
289 natural resource parks in the system to compare the frequency of frost days and wet days, the authors
climate data from the last 10–30 years to their evaluated the total number of times over the past few
documented historical range of variability (HRV) decades these variables evidenced “extreme” values,
from 1901 to 2012 (2014). Since it is logistically chal- defined as either less than the 5th percentile or greater
lenging to manage parks for a range of possible future than the 95th percentile of the HRV.
climate conditions and their accompanying novelties, While there was variation and complex patterns of
managers have traditionally developed conservation exposure among parks, 91% of parks reported as
strategies and policies relative to “existing, observed “warm” with respect to their HRV, signified here as
HRV, rather than an entirely new and uncertain set greater than the 75th percentile. Even further,
of conditions.” In this study, Monahan and Fisichelli the temperatures of more than half of all parks were
wanted to determine if recent climate conditions in consistently greater than the 95% percentile (Informa-
these parks were consistently evidencing unusually tion Box Fig. 4.1). With parks overwhelmingly at the
low or high values for certain climate variables extreme warm end of historical temperature dis-
(suggesting climate change), or if conditions were tributions, managers are concerned about the ongoing
Information Box Fig. 4.1 The average (mean) and maximum differ- are greater than the 75th percentile). (Reprinted under the Creative
ence of recent temperature change with respect to a National Park’s Commons CC0, from Monahan, W.B., Fisichelli, N.A., 2014. Climate
historical range of variability (HRV, 1901–2012). Practically all natural exposure of US national parks in a new era of change. PloS one 9,
resource parks are warm within respect to their HRV (292 (91%) parks e101302)
The absorption of infrared radiation warms the Earth,

Information Box 4.1 (continued) making it a more hospitable place for life than if the influx
effectiveness of current planning decisions. The and outflux of solar radiation to and from the Earth were
authors state that “many parks are already extremely equal. Without such warming, the average near surface tem-
dry or wet; if these observed extremes are followed by perature of the Earth would be 19 C instead of its current
future changes in the same direction, then affected 14 C. Each greenhouse gas differs, not only in its capacity to
parks will experience precipitation regimes unlike any absorb longwave radiation, but in other traits that affect its
they have seen in over a century” (Monahan and contribution to the overall warming produced. The key
Fisichelli 2014). Moving forward, “management and factors in the process are: (1) the amount of gas released
planning decision will be greatly informed by an over- into the atmosphere per year, (2) the length of time that it
all assessment of vulnerability that integrate the present stays in the atmosphere before being destroyed or removed
climate change exposure analyses with others that con- (also known as residence time), (3) any indirect effect it has
sider both the sensitivity and adaptive capacity of park on atmospheric chemistry, and (4) the concentration of and
resources to climate change” (Monahan and Fisichelli interactions with other greenhouse gases. If all of these
2014). This work supports the hypothesis that contem- factors are collectively taken into account, H2O and CO2
porary warming trends are indicative of larger scale emerge as the greenhouse gases that contribute most to this
climate changes rather than simple outliers within a warming effect, with water vapor accounting for about 60%
historical temperature range. of the warming effect and carbon dioxide about 26% (Kiehl
and Trenberth 1997).
Humans, through various activities, make net additions of
4.1.3 How Is Contemporary Change Different
greenhouse gases into the atmosphere. Of these, the largest
from Past Climate Change?
additions are of CO2 produced through the combustion of
fossil fuels. This addition is significant, because, as noted
Temperature on Earth varies spatially, seasonally, and histor-
above, CO2 is, next to water vapor, the gas most responsible
ically. As an index of energy, the range of temperatures on
for the greenhouse effect. The potential for increases in
Earth, both now and in past times, is driven by the absorption
atmospheric CO2 to spur related increases in atmospheric,
of solar radiation from the sun. The net incoming energy from
oceanic and surface temperatures has been recognized for a
such solar radiation is, as an “Earth average,” 342 watts per
long time. As early as 1856, scientists such as Eunice Newton
square meter (W/m2) (Karl and Trenberth 2005). Over the
Foote, were speculating that modest increases in CO2
entire Earth’s surface, this energy input is equal to
concentrations could result in atmospheric warming (Foote
175 petawatts (1 1015 W). Given that the largest human-
1856). Several decades later, Nobel laureate Swedish chemist
constructed power stations have energy production capacities
Svante Arrhenius developed a model quantifying the radia-
of about 1000 megawatts, the incoming energy is approxi-
tion budget of the Earth’s atmosphere and surface (Arrhenius
mately equivalent to that which could be produced by
1896), and from this was able to create a climate model that
175 million such power stations (Karl and Trenberth 2005).
agrees with many aspects of modern climate models
About 31% (107 W/m2) of this energy is reflected back into
(Ramanathan and Vogelmann 1997). Arrhenius was
space by clouds and atmospheric particles (aerosols) and
motivated by his desire to understand past temperature
never reaches the Earth’s surface, but the remaining 69%
variations from the Quaternary Period, but he also applied
(235 W/m2) is available to warm both the Earth and its
his results to present conditions, and predicted the
atmosphere (Fig. 4.3).
consequences of industrial emissions of CO2 on future cli-
In the act of being reflected from the Earth’s surface, the
mate change, including the consequence of global warming
wavelength of the emitted radiation is changed to long wave
(Saavedra 2002).
“infrared” radiation. The form of the reflected radiation is
Direct measurements of atmospheric CO2 and other GHG
important because such wavelengths are absorbed by certain
concentrations began in 1958 at the Mauna Loa Observatory
gases (also called greenhouse gases) in the Earth’s atmo-
in Hawaii (USA), but older records can be estimated by
sphere, notably water vapor (H2O), methane (CH4), ozone
indirect indices, especially by measuring the concentration
(O3), nitrous oxide (N2O) and carbon dioxide (CO2). Such
of GHGs trapped in ice cores, some of which, such as those
absorption creates a net influx of energy to the Earth. To
from Antarctica, can provide a continuous record of such
conserve the energy balance of the system, there must be
concentrations over the last 800,000 years (Fig. 4.4). The
some change in non-radiative energy states of the system,
molecules of CO2 trapped in this ice also provide a record
including temperature. This change is known as radiative
of atmospheric temperature inferred through variations in the
forcing.
Fig. 4.3 The components of the global average annual radiation budget space by the atmosphere and clouds. (Reprinted by permission from
and how much radiation is absorbed, scattered and emitted within the Elsevier, from Loeb, N.G., Wielicki, B.A., 2015. Satellites and satellite
atmosphere and at Earth’s surface. At the surface, an energy balance is remote sensing: Earth’s Radiation Budget, in: North, G.R., Pyle, J.,
achieved through evaporation and convection. At the top of the atmo- Zhang, F. (Eds.), Encyclopedia of Atmospheric Sciences (Second Edi-
sphere, the radiation balance is achieved by the heat energy emitted to tion). Academic Press, Oxford, pp. 67–76. # 2015 Elsevier Ltd)
concentration ratios of different isotopes of oxygen atoms in ppmv during the interglacial periods when the temperature
their molecules. Such examination reveals three key facts. was similar to present values. . .This narrow range of varia-
First, atmospheric CO2 concentrations and temperature show tion in atmospheric CO2 is remarkable given that its concen-
tight correlation. Second, until recently, concentrations of tration is determined by a highly dynamic biogeochemical
atmospheric CO2 varied between 280 ppm (warmest periods) cycle....This generally tight domain of stability between
and 180 ppm (coldest periods) over the last 450,000 years. variations in CO2 and global temperature . . .suggests that
Third, current levels of atmospheric CO2, at 410 ppm, are the global carbon cycle has been controlled by powerful
higher than ever before recorded during this 800,000-year biological feedback processes that have maintained the cli-
period (Fig. 4.4b). mate in a habitable range.” (Apps et al. 2006:176).
M. J. Apps, climatologist and former senior scientist with
the Canadian Forest Service, and his colleagues, drawing on
ice cores and other inferential data permitting estimates of 4.1.4 The Implications of Rapidly Rising CO2
CO2 and temperature going back 1.5 million years, noted this
tight and highly predictable historical oscillation. Studying As already noted, the current atmospheric concentration of
these patterns, they remarked “Throughout at least the last CO2 of 410 ppm appears to be higher, by approximately
four glacial cycles, spanning nearly 1.5 million years prior to 42%, than any recorded levels of the past 1.5 million years.
the twentieth century, the atmospheric concentration of CO2 More troubling is the fact that the current rate of increase in
only varied between ~180 ppmv during glaciations, when the atmospheric CO2 is now five times greater than any known
global temperature was 8–9 C colder than today, and ~280 historical increase, and more than 100 times faster than the
Fig. 4.4 (a) The concentrations

of CO2 in the atmosphere from
hundreds of thousands of years
ago through 2015, measured in
parts per million (ppm). The data
come from a variety of historical
ice core studies and recent air
monitoring sites around the world.
(Figure A from US Environmental
Protection Agency. 2016. Climate
change indicators in the United
States, 2016. Fourth edition. EPA
430-R-16-004. www.epa.gov/
climate-indicators.) (b) Recent
monthly mean carbon dioxide
measured at Mauna Loa
Observatory, Hawaii. The dashed
red line with diamond symbols
represents the monthly mean
values, centered on the middle of
each month. The black line with
the square symbols represents the
same, after correction for the
average seasonal cycle. (Figure B
courtesy of NOAA/ESRL/GMD)
increase that occurred at the end of the last ice age (NOAA Carbon moves into and out of the atmosphere in a com-
2013). As the concentration of atmospheric CO2 has plex biogeochemical cycle, but one which can be
increased, so has the positive radiative forcing associated summarized in terms of “sources” and “sinks” and their
with it, now at a level two times what it was in 1800 (IPCC respective additions or removals. Humans add CO2 to the
2013). As is historically documented in ice cores, global atmosphere through their combustion of fossil fuels, but they
temperatures and atmospheric CO2 increases have been also contribute to net additions through the changes they
correlated, such that, since 1901, nearly every measured impose on the landscape (Information Box 4.2). When
location on Earth, both terrestrial and aquatic, has seen burned, the CO2 emitted by fossil fuels possesses a charac-
some level of temperature increase (IPCC 2013). In a recent teristic “signature” of carbon and oxygen isotope ratios that
special report, the Intergovernmental Panel on Climate are different from CO2 originating from other sources. Most
Change (IPCC) disclosed that the global mean surface tem- additions of CO2 to our atmosphere bear these signatures.
perature has increased approximately 1 C (0.8–1.2 ) above This is one manifestation of what many scientists have come
pre-industrial levels (IPCC 2018). to call the “global fingerprint” of human-induced climate
change.
Information Box 4.2: Contributions of Land-Based Information Box 4.2 (continued)

Carbon: Moving from a Carbon Sink to a Carbon Although recent studies have confirmed the presence of
Source major global carbon sinks, including those in the
Natural vegetation, and particularly forest vegetation, United States, their long-term role in maintaining cli-
removes CO2 from the atmosphere via photosynthesis, mate stability is not guaranteed. Mathematical ecolo-
during which CO2 is combined with water in the plant gist and geographer, George C. Hurtt, and colleagues
to produce glucose according the familiar reaction, conducted a study to project the future of the
6CO2 + 6H2O ! C6H12O6 + 6O2. In the complemen- US carbon sink given known ecosystem recovery pro-
tary reaction of respiration, plants, like animals, also cesses and a range of land-use management decisions.
release CO2 back to the atmosphere, but their overall Using a dynamic ecosystem model called the Ecosys-
effect is to remove more CO2 through photosynthesis tem Demography Model (or ED) (Hurtt et al. 1998;
than they release through respiration, thus acting Moorcroft et al. 2001), the authors model complex
as a carbon sink. When humans remove forests and future interactions between climatic, environmental
other kinds of intact vegetational communities, the and land-use conditions based on historical patterns
net effect is to reduce such carbon removal capacities.
Information Box Fig. 4.2 (a) Estimated average annual area suppression ceases. (Reprinted by permission from the National
burned from 1700 to 2100. (b) Projected average annual air-to- Academy of Sciences, from Hurtt, G. C., Pakala, S. W., Moorcroft,
ground net flux from 1700 to 2100. Positive values indicate a land P. R., Caspersen, J., Shevliakova, E., Houghton, R. A., & Moore,
sink and negative values indicate a source to the atmosphere. B. (2002). Projecting the future of the US carbon sink. Proceedings
Diamonds represent ED model estimates; squares represent Miami- of the National Academy of Sciences, 99(3), 1389–1394. # 2002
LU estimates. Solid symbols represent cases assuming continued fire National Academy of Sciences, USA)
suppression. Empty symbols represent cases assuming fire
In their fifth assessment report, the IPCC highlighted

Information Box 4.2 (continued) several of well-established and ongoing global changes
(Hurtt et al. 2002). By focusing on the management (2014, Fig. 4.5). For example, with a greater level of retained
role of fire suppression, the authors present two radiant energy, our planet is experiencing enhanced rates of
scenarios which span a broad range of possible futures, evaporation and transpiration and creating, in many parts of
one in which current fire suppression efforts continue the world, more rapid and energetic hydrologic cycles.
to be effective at reducing fires throughout the twenty- Although precipitation is increasing in many parts of the
first century, and a second which assumes that fire world, historically dry areas may, in contrast, face even
suppression ceases (or ceases to be effective) given more extreme drought conditions (Fig. 4.5e). Even as the
the increased fire risk as ecosystems accumulate car- average global temperature is rising, regional temperature
bon. As highlighted in Information Box Fig. 4.2, the changes are non-uniform (Fig. 4.5b), with the least change
results reveal a dramatic difference in the impact of fire in equatorial regions and the greatest changes in polar areas,
management on the carbon sink. Even with continued resulting, at the poles, in reduced snow and ice cover and
fire suppression (Information Box Fig. 4.2a), the reduced areas of sea ice (Fig. 4.5c). Similarly, glaciers on all
US sink is expected to decline as forest recovery continents (except Australia, which has no glaciers) are
slows and begins to equilibrate. However, if fire sup- retreating, and attendant glacial and alpine environments are
pression efforts were to completely fail, the shrinking in size (Brown et al. 2007).
US carbon sink would be replaced by a source due to Water in the oceans is also warming, particularly from the
extensive burning from large-scale fires (Informa- surface to about 700 meters in depth. As it does, the ocean
tion Box Fig. 4.2b). The authors notes here that “with- increases in volume. This effect, known as thermal expan-
out substantially positive increases in the area of sion, is the main contributor to worldwide sea level rise
forests, without substantially positive changes in land- (Fig. 4.5d). Furthermore, as atmospheric concentrations of
use practices, without large net positive effects of CO2 CO2 rise, more CO2 will diffuse into the oceans, shifting the
or climate change in the future, or without some other chemistry of sea water so that its total carbonate alkalinity
new significant carbon storage mechanisms, the will decrease (i.e., seawater pH will increase, and the oceans
US carbon sink will decrease substantially over the will become more acidic). The global “conveyor belt” of
21st century. . .total US fossil fuel emissions would worldwide ocean currents, which transfers heat energy from
need to be reduced by an additional 7–30% to compen- tropical areas to colder temperate and polar oceans and their
sate for the declining sink and stabilize net emissions” adjacent seacoasts, as well as bringing nutrients up from
(Hurtt et al. 2002). This study highlights the important deeper waters into more productive, photosynthetically
role of forest carbon sequestration for climate stability active surface waters, will slow, and potentially stop, as the
and to carbon mitigation policies and planning. oceans warm (Hoegh-Guldberg 2005).
The sum of these and other changes, suggest that all living
things on Earth are facing reality of climate change, one to
which they must adapt for the foreseeable future. For those
Because overall CO2 concentrations, and human
humans engaged in the work of conservation, these changes
contributions to them, are still rising, and are expected to con-
have profound implications. As scientists we are motivated
tinue rising through at least the first half of the twenty-first
to continue documenting, quantifying, mapping, projecting,
century due to past emissions, the world appears to be, at
and responding to anticipated climate changes and their
minimum, decades from climate stability. Even the most
impacts. As global citizens, we must consider our role in
optimistic scenarios do not foresee a stabilization of CO2 levels
advancing climate mitigation strategies that seek to slow the
short of 450 ppm (Table 4.1). Even if human-induced CO2
changes our world is experiencing to safeguard the diversity
inputs begin to fall, the persistence time (also known as “resi-
of life on Earth. Although the degree of future change
dence time”) of carbon in the atmosphere and oceans, in a variety
humans will allow to happen is still uncertain, the over-
of forms, will prevent any significant change in atmospheric CO2
whelming advances in climate science and knowledge of
for decades or centuries. Human action, or perhaps more aptly
interactions between our atmosphere, ocean and terrestrial
inaction, is transforming life as we currently know it. The effects
biosphere provides a solid foundation from which to engage
of a warming world are truly global and showcase a range of
the challenges of climate-informed conservation.
responses among all components of our climate system.
Table 4.1 Assessments in this table are based on the probabilities calculated for the full ensemble of scenarios in the Intergovernmental Panel on
Climate Change’s (IPCC’s) Fifth Assessment Report
Additional Global mean temp. increase above Change in global
radiative CO2 CO2-eq pre-industrial at equilibrium, Peaking CO2 emissions in No. of
forcing concentration concentration using “best estimate” climate year for 2050 (% of 2000 assessed
Cat. (W/m2) (ppm) (ppm) sensitivitya,b ( C) emissions emissions) scenarios
I 2.5–3.0 350–400 445–490 2.0–2.4 2000–2015 85 to 50 6
II 3.0–3.5 400–440 490–535 2.4–2.8 2000–2020 60 to 30 18
III 3.5–4.0 440–485 535–590 2.8–3.2 2010–2030 30 to +5 21
IV 4.0–5.0 485–570 590–710 3.2–4.0 2020–2060 +10 to +60 118
V 5.0–6.0 570–660 710–855 4.0–4.9 2050–2080 +25 to +85 9
VI 6.0–7.5 660–790 855–1130 4.9–6.1 2060–2090 +90 to +140 5
Total 177
The vast majority of scenarios overshoot the category boundary of 480 ppm CO2-equivalent concentration
a
Note that global mean temperature at equilibrium is different from expected global mean temperatures in 2100 due to the inertia of the climate
system
b
Non-linearities in the feedbacks (including e.g., ice cover and carbon cycle) may cause time dependence of the effective climate sensitivity, as well
as leading to larger uncertainties for greater warming levels. The best-estimate climate sensitivity (3 C) refers to the most likely value
Reprinted from Climate Change 2007: Mitigation of Climate Change. Working Group III Contribution to the Fourth Assessment Report of the
Intergovernmental Panel on Climate Change [B. Metz, O.R. Davidson, P.R. Bosch, R. Dave, L.A. Meyer (eds.)]. Cambridge University Press,
Cambridge, United Kingdom and New York, NY, USA
while in other cases, it acts as a compounding factor by

4.2 The Global Fingerprint of Climate exacerbating existing vulnerabilities.
Change on Biodiversity In one of the first major syntheses of its kind, Camille
Parmesan and Gary Yohe undertook a meta-analysis of
4.2.1 Common Ecological Responses published studies, evaluating over 1700 species, to determine
to Current Climate Change if recent biological trends in range shifts and biological
timing events matched climate change predictions
Just as climate scientists have been able to identify the global (Table 4.2). Among these studies, 87% of those documenting
markers of climate change within the climate system, so have timing (phenological) events, 75–81% of those examining
biologists and ecologists been able to identify signs of a range boundaries, and 81% of those measuring community
changing climate among vegetation and wildlife. Over the abundance found changes consistent with predictions of cli-
past several decades, scientists have not only documented a mate change. Overall, Parmesan and Yohe documented pole-
range of species-specific responses, but also identified com- ward range shifts of 6.1 km per decade and advancement of
mon responses across broad geographic regions and diverse spring events by 2.3 days per decade in these studies.
taxonomic groups. These changes are having a profound Between 1899 and 2003, the publication date of the authors’
impact on conservation. As bioclimatologists Lee Hannah meta-analysis, 866 peer-reviewed papers had documented
and Laura Hansen point out, “Climate change affects selec- changes through time in species or systems that could be, in
tion of [conservation] targets in two fundamental ways. First, whole or in part, attributed to climate change. In 2013,
it will alter ecoregion boundaries. . .[resulting] in different ecologist Elvira Poloczanska and her colleagues, including
biome boundaries in the future, which may change the size Camille Parmesan, conducted a similar large-scale review of
or configuration of an ecoregion. . . .The second way in which published marine ecological observations. Reviewing
climate change can affect target selection is by exacerbating 279 observed shifts taken from 36 published studies, the
existing threats. . . Therefore, climate change is likely to authors found that 83% of observed changes in single species
increase the number of conservation targets, through both studies were in the direction expected under climate change.
changing the planning area and by increasing the number of Of all observations considered, 95% of these studies
species at threat” (Hannah and Hansen 2005:331). In some identified temperature as the primary climate change driver
cases, climate is the predominant or singular force of change, with the remainder relating biological change to changes
4.2 The Global Fingerprint of Climate Change on Biodiversity 135
Fig. 4.5 Multiple observed indicators of a changing global climate aligned to have the same value in 1993, the first year of satellite
system. (a) Observed globally averaged combined land and ocean altimetry data, and (e) Map of observed precipitation change, from
surface temperature anomalies (relative to the mean of 1986–2005 1951 to 2010. (Reprinted from Climate Change 2014: Synthesis Report.
period, as annual and decadal averages), (b) Map of the observed Contribution of Working Groups I, II and III to the Fifth Assessment
surface temperature change, from 1901 to 2012, derived from tempera- Report of the Intergovernmental Panel on Climate Change [Core
ture trends, (c) Arctic (July to September average) and Antarctic Writing Team, Pachauri, R.K. and Meyer, L. (eds.)]. IPCC, Geneva,
(February) sea ice extent, (d) Global mean sea level relative to the Switzerland)
1986–2005 mean of the longest running data set, and with all data sets
in pH, sea level rise, sea ice extent and climate oscillations coherent signal” represents a “diagnostic fingerprint” of the
(Poloczanska et al. 2013). impact of global climate change on biodiversity (Parmesan
These changes in species response are not random, but and Yohe 2003). This fingerprint “of temporal and spatial
consistently in the direction expected from climate change. ‘sign-switching’ responses” is uniquely predicted by twenti-
Further, the responding species are spread across diverse eth century climate trends (Parmesan and Yohe 2003). It
ecosystems, and come from a variety of taxonomic and provides compelling evidence for current widespread climate
functional groups. The authors submit that this “globally impacts and the potential reconfiguration of ecosystems into
Table 4.2 Summary of biological change events from a meta-analysis of studies of 1700 species showing change in timing of biological events
(phenology) or changes in distribution or abundance in relation to changes predicted by climate change theory
Type of change Changed as predicted Changed opposite to prediction P-value
Phenological (N ¼ 484/(678)) 87% (n ¼ 423) 13% (n ¼ 61) <0.1 1012
Distributional changes
At poleward/upper range boundaries 81% 19% –
At equatorial/lower range boundaries 75% 25% –
Community (abundance) changes
Cold-adapted species 74% 26% –
Warm-adapted species 91% 9% –
N ¼ 460/(920) 81% (n ¼ 372) 19% (n ¼ 88) <0.1 1012
Meta-analyses
Range-boundaries (N ¼ 99) 6.1 km m1 per decade northward/upward shifta 0.013
Phenologies (N ¼ 172) 2.3 days per decade advancementa <0.05
Reprinted by permission from Springer Nature, from Parmesan, C., Yohe, G., 2003. A globally coherent fingerprint of climate change impacts across
natural systems. Nature 421, 37–42. # 2003 Nature Publishing Group
Notes: Data points represent species, functional groups or biogeographic groups. N, number of statistically or biologically significant changes/(total
number species with data reported for boundary, timing, or abundance processes). The no prediction category is not included here
a
Bootstrap 95% confidence limits for mean range boundary change are 1.26, 10.87; for mean phenological shift the limits are -1.74, -3.23
the future (Poloczanska et al. 2013). A 2017 study 4.2.2 Phenological Changes and Mismatched
emphasizes this point by estimating that 47% of terrestrial Interactions Across Trophic Levels
non-volant threatened mammals (out of 873 species) and
23.4% of threatened birds (out of 1272 species) may have 4.2.2.1 Timing Is Everything
already been negatively impacted by climate change in at Changes in species phenology, the seasonal timing of life
least part of their distribution (Pacifici et al. 2017). history events, is one of the most unambiguous consequences
of climate change. Life history traits such as fecundity,
development, and survivorship exhibit plastic (highly adap-
tive) responses to variation in temperature (Scranton and
What global fingerprints of climate change have you
Amarasekare 2017). This means that individuals can alter
seen where you live? Often to the observant, or by
(and have altered) their physiology, morphology and behav-
talking to those who have lived in a given area for a
ior in response to changes in climate. The degree of pheno-
long period of time, the changes happening around us
typic plasticity varies across species, both in the range of
becomes more obvious. How might taking a diagnostic
temperatures these species currently inhabit and the degree of
survey of these changes help to inform action to
future change to which a species may be able to adapt
address the negative consequences of them?
(Fig. 4.6). In addition to absolute changes in biological
events, there is a growing amount of evidence that such
changes in an individual species can disrupt the coordination
As summarized by Parmesan (2006), there are four general of activities between this species and other species in any
categories of species response to recent climate change: number of interactions, such as predator and prey, parasite
(1) phenological changes (advancement of spring events), and host, or pollinator and flower.
(2) interactions across trophic levels (increasing asynchrony For example, David Inouye and his co-workers at the
in predator-prey and insect-plant systems), (3) observed range Rocky Mountain Biological Laboratory in Colorado,
shifts and changing trends in local abundance (poleward range USA found that, from 1976 to 1999, yellow-bellied marmots
shifts, expansions of warm-adapted communities, complicated (Marmota flaviventris), large ground squirrels that live
trees and tree line shifts), and (4) extinctions (particularly for mainly in montane environments of the western United
range-restricted species). Greater attention to the ‘here and States, had advanced their emergence date by 38 days as a
now’ impacts of change is warranted and may be underappre- result of warmer local spring air temperatures. Such advance-
ciated in efforts to better predict the effects of future changes. ment has put them out of synchrony with most of the
Here, we review several examples of contemporary changes surrounding plant species, their major food source. At the
across both terrestrial and aquatic systems, before discussing same study site, American robins (Turdus migratorius) have
the tools being used to understand and project ecosystem advanced their arrival dates over about the same period
responses under a range of potential climate futures. (1974–1999) by 14 days, and this increasingly early return
Fig. 4.6 Thermal responses of

life history traits. (a–i)
Temperature responses of the per
capita rates of adult mortality (a–
c), birth (d–f), and maturation (g–
i) for three Hemipteran insect
species from temperate (Apolygus
lucorum), Mediterranean
(Murgantia histrionica), and
tropical (Clavigralla shadabi)
latitudes. Circles represent
response data from experiments.
Curves are the fits of mechanistic
response functions to the data.
The vertical line represents mean
habitat temperature and shaded
regions show the range of
seasonal temperature variation
experienced by each species. An
increase in the mean
environmental temperature of
3–6 C over 100 years decreases
peak abundance and mean annual
abundance in the tropical species.
The temperate species, by
contrast, benefit from a moderate
increase in the mean temperature
(3–6 C) with an increase in mean
annual abundance and peak
abundance. (Reprinted by
permission from the National
Academy of Sciences, from
Scranton, K., & Amarasekare,
P. (2017). Predicting phenological
shifts in a changing climate.
Proceedings of the National
Academy of Sciences, 114
(50), 13212–13217. # 2017
National Academy of Sciences)
has spread the difference between their arrival and the aver- pollinators and disrupt the plant-pollinator interaction net-
age first appearance of bare ground (which robins require for work structure (Devoto et al. 2007).
foraging) by an additional 18 days (Inouye et al. 2000). Ecologists Jessica Forrest and James Thomson undertook
a study within the Rocky Mountains of Colorado (USA) to
4.2.2.2 New Patterns of Pollination assess whether or not plants and insects in the subalpine
Climate change is affecting the flower date of many plants, habitats within this region were advancing their phenology
advancing their first flowerings earlier in the Spring than in in response to springtime warming similarly or asynchro-
the past (Calinger et al. 2013). Although less documented nously (2011). Over 3 years, Forrest and Thomson collected
over long-term records, some pollinators, such as bees and data on air temperature, insect emergence (specifically, the
butterflies, are also now appearing earlier in the year (Forister cavity nesting Hymenoptera) and flowering phenology. With
and Shapiro 2003). When phenological changes in plants and the use of experimental trap nests across a range of
pollinators occur at the same time in response to climate elevations, they checked each site for insect emergence and
change, the degree of synchronization between them can be documented the number of open flowers per plant. Among
maintained. However, when the timing of one species shifts their results, the authors found that one plant species
faster than another, or when this shift is in the opposite (Lathyrus lanszwertii) and one bee species (Hoplitis fulgida)
direction, such mismatched timing can result in a depression showed broad overlap in phenology at most sites in 2008 but
of plants or the starvation of pollinators (Morton and Rafferty only slight overlap in 2009, such that, at all sites, Lathyrus
2017; Fig. 4.7). At the same time, a mismatch in the spatial had finished or almost finished flowering by the time of peak
shift of pollinators and plants in response to change, regard- H. fulgida emergence (Forrest and Thomson 2011). This was
less of timing, could affect the visitation of plants by due, in part, to generally earlier flowering by this plant
Fig. 4.7 Conceptual depiction of

phenological and distributional
shifts in populations of plants and
pollinators under climate change,
with populations depicted as
single icons. The timing of
flowering for plants and activity
for pollinators is represented as
shades along a color scale. Pre–
climate change distributions are
shown in gray. (a) Positions
occupied by populations pre–
climate change (historical
baseline). (b) Phenological/
temporal shifts alone. (c)
Distributional/spatial shifts alone.
(d) Joint phenological/temporal
and distributional/spatial shifts.
(b–d) Each panel illustrates
possible outcomes of shifts:
(i) maintenance of historical
interactions (indicated by O),
(ii) loss of historical interactions
(indicated by –), and (iii) gain of
novel interactions (indicated by
+). The severity of the outcome
will vary depending on the pre–
climate change situation, the
degree of shifting of each
population, and many other
factors. (Reprinted by permission
from Wiley, from Morton, E. M.,
& Rafferty, N. E. (2017). Plant–
pollinator interactions under
climate change: The use of spatial
and temporal transplants.
Applications in Plant Sciences, 5
(6). # 2017 The Authors)
species at warmer, low-elevation sites in 2009 compared to 4.2.2.3 Shifting Predator-Prey Dynamics
2008, without a corresponding advance in phenology of the A change in phenology can also create new dynamics in
bees. However, when considering another plant-bee relation- predator-prey relationships. Using over 30 years of data,
ship, Potentilla hippiana and H. fulgida, the authors found scientists from the US Geological Survey and
consistent matching between insect emergence and peak US National Park Service evaluated the relationship
flowering days across all sites and years. between bald eagle habitat use and chum and coho salmon
In evaluating the implications of this work for plant and availability in the Skagit River (Washington, USA)
insect populations the author suggests that under novel cli- (Rubenstein et al. 2019). The Skagit River is a key geo-
mate conditions, some species might be able to co-adapt graphic and ecological feature in the western portion of the
together while other species will experience either a reduced North Cascades National Park Service Complex (NOCA),
or increased overlap. Although this contracted period of which connects remote montane ecosystems to the Puget
overlap does not amount to a complete decoupling of Sound and serves as a key wintering site for bald eagles
pollinator-flower interactions, it does suggest that a generalist (Haliaeetus leucocephalus), an apex predator that feeds in
bee species may be able to persist and outcompete specialist high densities on spawning chum (Oncorhynchus keta), and
pollinators as long as some plants within their diet are coho (Oncorhynchus kisutch) salmon. Here, long-term
flowering (for example, the H. fulgida maintained synchrony datasets are particularly helpful for understanding trends in
with Potentilla even though it missed Lathyrus flowering). species distributions, habitat use, and phenology, and
However, if plants consistently flower earlier than insect distinguishing climate change from natural variability
species, especially with more rapid snowmelt, plants are at (Rubenstein et al. 2019).
risk of reduced flowering over time which could negatively To evaluate the implications of change on the trophic
affect both the plant and the insect populations. interactions in this system, the authors specifically considered
trends in eagle habitat use, salmon escapement (the propor- this lag may be that suitable new conditions are hard to reach
tion of a salmon population that does not get caught by (like a mountain peak) or that the complex combinations of
fisheries and returns to their freshwater spawning habitat), topographic or microclimate requirements are hard to find
and the timing and intensity of flood events. They found that (for example, cooler locations on a poleward-facing slope).
peaks in chum salmon and bald eagle presence had advanced Although this work highlights a general directional pattern,
at remarkably similar rates (c. 0.45 days/year). Although species range shifts are individualistic and can be influenced
predators have been known to display slower phenological by non-climatic factors and species interactions.
responses than their prey (e.g., Thackeray et al. 2010), the
synchronous relationship observed here is likely a reflection 4.2.3.2 Rising Sea Temperatures and Deepening
of eagles’ ability to rapidly respond and adaptively take Fish Assemblages
advantage of ephemeral (temporary) resources across the The seas are warming faster than adjacent land and faster than
landscape (Knight and Knight 1983; Rubenstein et al. 2019). the global average (MacKenzie and Schiedek 2007). This
In contrast, the authors found an asynchronous temporal warming has a particular effect on the distributional
relationship between chum peaks and flood events. Over the responses of bottom-dwelling (demersal) fishes, with
30-year study period, the interval between chum peak and two-thirds of all North Sea fishes shifting mean latitude or
first flood event increased significantly, growing by nearly depth (Dulvy et al. 2008). This fish response is a marine
1 day per year. The authors note that as this interval grew analogue to the upward movement of terrestrial species to
over time, “a phenological paradigm shift occurred: histori- higher altitudes. In the span of 25 years, European scientists
cally, the earliest flood events occurred while chum were still documented an 1.6 C increase in the winter bottom temper-
alive and swimming, whereas increasingly, floods begin after ature of the North Sea with a corresponding deepening of all
chum have peaked, spawn, and are on gravel banks in the but one of 19 demersal fish assemblages by ~3.6 m per
form of carrion for eagles” (Rubenstein et al. 2019). decade (Dulvy et al. 2008). The only assemblage not
Although the ecological implications of this shift are not yet exhibiting a significant depth response was comprised of
fully understood, the authors suggest that salmon mortality species that are warm-tolerant, small bodied, have a northern
and consumption patterns could have wide-ranging range boundary, and are unexploited by fisheries (such as the
consequences for components of the riparian ecosystem that scaldfish, solenette and bib) (Fig. 4.8). This coherent deepen-
are affected by transfer of nutrients from marine to freshwater ing was accompanied by a heterogenous latitudinal response.
systems facilitated by salmon migration (Schindler et al. Assemblages shifting northward were comprised of abun-
2003). dant, widespread, and warm-tolerant species with narrow
thermal ranges, while the assemblage that showed a south-
ward shift of minimum latitude was the same as the one that
4.2.3 Observed Ranges Shifts and Changes did not exhibit a deepening response (Fig. 4.8b).
to Local Abundance Although the ecological significance of this deepening is
less clear than its terrestrial analogue of upward-shifting
4.2.3.1 Higher Latitudes and Higher Elevations alpine fauna, there could be biological consequences for
In a 2011 meta-analysis, biologist I-Ching Chen and geographically restricted species that cannot deepen or shift
colleagues reviewed the literature to investigate evidence their range to remain within their preferred temperature gra-
for a direct link between temperature change and species dient. For example, the eelpout (Zoarces viviparous) has
range shifts. Their latitudinal analysis revealed that species declined due to rapid warming in the shallow and enclosed
have moved away from the equator at a median rate of Wadden Sea (Pörtner and Knust 2007). Even without geo-
16.9 km per decade and made a shift to higher elevations at graphical barriers, the comparatively smaller area of deeper
a median distance of 11 m per decade (Chen et al. 2011). To habitat may restrict availability (Dulvy et al. 2008). Recent
estimate expected shifts, the authors calculated the distance in studies have indicated that fish nurseries in particular are
latitude (km) and elevation (m) that species in a given region showing sensitivity to climate warming as they are depth-
would be required to move to track corresponding tempera- limited and likely to warm quickly with limited resistance to
ture changes. Observed range shifts were positively temperature fluctuations (McLean et al. 2018). In one study
correlated with predicted distances (r ¼ 0.65, P ¼ 0.002 for of the Bay of Somme, within the Eastern English Channel,
latitude; r ¼ 0.39, P ¼ 0.035 for elevation), and increased biologists saw a substantial change to the functional structure
with the level of warming experienced. The authors noted of the fish community within the nursery (from r-selected to
that, although latitudinal range shifts were not consistently k-selected dominance) due to increasing sea surface
lagging behind the shift in climate there was a notable lag in temperatures (McLean et al. 2018).
elevation shifts despite the shorter distances required to track
climate changes (Chen et al. 2011). One potential reason for
Fig. 4.8 (a) Trend in depth

anomaly of individual fishes over
time (m decade–1). Solid points
are significant at P < 001. (b)
Trend in geographic response of
different demersal fish
assemblages over time; (a) mean
depth, (b) mean latitude, (c) mean
minimum latitude and (d) mean
maximum latitude. Black and grey
points indicate statistical
significance at P 0001 and
P 001 respectively. The x-axis
represents the direction and
strength of geographic response
over time – the slope of a
regression of distribution measure
on year. Positive values indicate
shallower (panel a) or northerly
distribution (panels b–d), with
negative anomalies representing
deepening or a more southerly
distribution. (Reprinted by
permission from Wiley, from
Dulvy, N. K., Rogers, S. I.,
Jennings, S., Stelzenmüller, V.,
Dye, S. R. and Skjoldal, H. R.
(2008), Climate change and
deepening of the North Sea fish
assemblage: a biotic indicator of
warming seas. Journal of Applied
Ecology, 45: 1029–1039. # 2008
British Ecological Society)
4.2.3.3 Contracting Ranges and the Spread the fungus can spread more readily under moist conditions.
of Pathogens Whitebark pine, being the conifer adapted to the highest
The whitebark pine (also known as the white pine, Pinus elevations, cannot “shift up” like lower elevation species as
albicaulis) is a conifer of the western United States and the climate warms. Lodgepole pine (Pinus contorta), the
Canada adapted to conditions at high elevations, the last species historically occupying elevational zones immediately
tree species present before reaching the treeless alpine vege- below the whitebark pine, has moved upward in elevation in
tation and talus slopes of the high plateaus and peaks. response to climate warming. As the distribution of
Whitebark pine produces a heavy, nutritious, and tasty seed whitebark and lodgepole pine have increasingly overlapped,
(“nut”), with highly variable seed production in different lodgepole pine has proven the superior competitor,
years. Because of the high mass of the seed, it has, on its demonstrating an ability to displace whitebark pine over
own, very limited dispersal ability. However, because of the time where the two species become established on the same
pine’s mutualistic relationship with a bird, the Clark’s nut- site. In addition, warming trends now occurring in the
cracker (Nucifraga columbiana), the seed is spread great whitebark pine’s range make the landscape more vulnerable
distances throughout the mountains. Clark’s nutcrackers, to fire, a disturbance to which the whitebark pine is poorly
which prefer habitat near the upper edge of the tree line, not adapted, but one to which lodgepole pine is favorably
only use the seeds of the whitebark pine as a short-term food adapted.
source, but also “plant” the seeds in caches over many miles Because whitebark pine contributes to biodiversity in sev-
of landscape. Direct estimates indicate that a single Clark’s eral ways with respect to their ecological roles, the loss of this
nutcracker may plant 50,000–120,000 seeds in up to 8000 species jeopardizes the health of high-elevation forest
caches (Koteen 2002:346). It is the diligence of the bird, not communities as we currently know them (Tomback and
the dispersal ability of the seed, to which the whitebark pine Achuff 2010). In the United States, 98% of whitebark pine
probably owes its extensive distribution throughout western can be found within national parks and forests, and in Canada
North America. the majority of whitebark pine occurs on federal and provin-
Another species feeding on whitebark pine nuts is the cial public land (Keane 2000; Hamann et al. 2004). Given the
grizzly bear (Ursus arctos horribilis), a subspecies of overlap between current distribution and protected areas, it is,
brown bear, once widespread in North America but now as the authors describe, “paradoxical that whitebark pine is
confined mainly to the northwestern portions of Montana seriously threatened by anthropogenic problems – an
and Wyoming, the latter population living in Yellowstone introduced pathogen, fire suppression and climate change.”
National Park and its surrounding national forests. Grizzlies The losses of whitebark pine have become large enough on
obtain from 25 to 67% of their caloric energy from whitebark some severely affected sites to compromise their functional
pine seeds, depending on the availability of the crop (Mattson role, with resulting impacts on Clark’s nutcracker visitation.
et al. 2001). In years of high mast production by the Proposals to assist this threatened species in upward migra-
whitebark pine, grizzlies remain close to productive stands tion have caused significant controversy, but in a review of
of whitebark pine and greatly reduce their movements. In the benefits this keystone species provides to sub-alpine
years of poor mast production, they wander more widely in habitats, there may be good reasons for people to move
search of food. These wanderings result in a greater need for whitebark pine if rust-resistant strains can be clearly
and higher numbers of management trappings to remove identified (Palmer and Larson 2014).
grizzlies from situations of bear-human conflicts, with higher
rates of bear mortality associated with their encounters with 4.2.3.4 Competitive Advantage of Invasives
humans (Mattson et al. 2001). The problem of invasive species (Chap. 3), currently one of
Recent decades have seen a gradual but persistent reces- the greatest threats to biodiversity on Earth, may be amplified
sion of the range of the whitebark pine, up to 90% range by climate change if such changes favor invaders. Jeffrey
reduction in some areas. Various environmental stressors Dukes and Harold Mooney, two biologists who are experts
have contributed to such recession, but one of the most on endangered species, assessed the impacts of climate on
important is a pathogen. The whitebark pine is vulnerable this problem in these words. “We expect most aspects of
to a particular disease organism, the white pine blister rust. global change to favor invasive alien species and thus to
The blister rust, a type of fungus, spreads to the whitebark exacerbate the impacts of invasions on ecosystems. . .These
pine through an intermediate host, the gooseberry or currant impacts include competitive effects, whereby an invading
(Ribes spp.). Although Ribes was historically uncommon at species reduces resources available to other species, and
the high elevations used by whitebark pine, warming ecosystem effects, whereby an invader alters fundamental
temperatures and increasing precipitation in recent decades properties of the ecosystem. Either type of effect can threaten
are increasing overlap between the species and increasing native biodiversity, and some ecosystem effects feedback to
incidence of infection in whitebark pine, especially since elements of global change” (Dukes and Mooney 1999:135).
Every invasive species presents a unique case history of including through their distribution and local abundance. In
mechanisms through which the invader becomes established total they documented characteristics of 258 native and
and negatively affects native species. But Dukes and Mooney 177 exotic grass species in California, with both groups
believe their prediction is warranted in general terms because varying in their traits (height, width and length of leaves,
most invasive species share certain life history traits, traits seed mass, annual vs. perennial). Many of the characteristics
that confer increasing advantages under a regime of rapid were related to mean annual temperature, with warmer sites
climate change. First, among plants, many invasive species showing on average larger species with larger seeds, earlier
respond positively to elevated CO2 levels, such as cheatgrass flowering times, and longer flowering seasons.
(Bromus tectorum), kudzu (Pueria lobata), and Japanese On average, the authors found that exotic grasses were
honeysuckle (Lonicera japonica). As a consequence of ele- taller, had longer, wider leaves, and larger seed size than
vated CO2 levels, most plants, including a number of inva- native grasses. Exotics were also more likely to be annuals
sive species, use water more efficiently and reduce their rate and evidence more light-capturing ability than native species.
of transpiration. In areas where plant species abundance is Possessing these traits, the authors hypothesized that exotics
limited by water availability, such species will gain a com- and invasive species would be favored at higher
petitive advantage. It is also characteristic of many invasive temperatures, with the warmer parts of the state showing a
species that they possess wider environmental tolerances and higher proportion of exotic species. Through this work, they
thus occur over wide latitudinal gradients, pre-adapting them found that the proportion of exotic species within a given
to climate change. zone was positively related to mean annual temperature. Leaf
Many invasive species also possess high rates of effective width difference between native and exotics alone in relation
dispersal, as well as potential for rapid reproduction and to temperature, explained 71% of the variation in the native
population growth, contributing to more rapid adaptive total richness ratio in California (Sandel and Dangremond
responses. Species that are able to shift ranges quickly are 2012). The number of exotic species in the state continues to
at an advantage. As Dukes and Mooney note, “Rapid dis- increase (Fig. 4.9) and corresponding shifts in plant commu-
persal is characteristic of many biological invaders. Within nity composition may affect herbivores, resulting in further
the genus Pinus, species that are invasive tend to have traits changes to the plant community (Wookey et al.
that facilitate rapid range shifting, such as short juvenile 2009) (Chaps. 3 and 9).
period and low seed mass (which is associated with long-
distance wind dispersal). . .” (Dukes and Mooney
1999:137) (Chap. 3). Overall, “A rapid anthropogenic cli- 4.2.4 Increased Threat of Extinction
mate change might disadvantage species that cannot quickly
extend their ranges into newly suitable regions, such as plants 4.2.4.1 Local Climate-Driven Extinctions
with long generation times. A climate-driven decline of late- From 1901 to 1987, mean annual temperature in the deserts
successional plant species could lead to increased dominance of the southwestern United States has increased 0.12 C per
of early successional species or could leave ill-adapted plant decade (Lane et al. 1994), while, at the same time, annual
communities that are susceptible to invasion by species that precipitation has decreased by roughly 20% over the last
can thrive in the area’s new climate” (Dukes and Mooney century in southeastern California (Ball et al. 1998). Drought
1999:137). More troubling is that, as many invasive species can cause increased mortality among desert bighorn sheep
become established, they have shown the ability to alter basic (Ovis canadensis nelsoni) (Monson 1960) and affect recruit-
ecosystem properties in ways that feed back to affect many ment dynamics (Wehausen et al. 1987). C. W. Epps and his
components of global change, creating positive feedback colleagues used simple presence and absence data to examine
loops that facilitate their permanent establishment. the effects of climate and other factors in determining bighorn
Assessing the likelihood of these outcome within a partic- metapopulation structure. They first examined all mountain
ular system, biologists Brody Sandel and Emily Dangremond sheep ranges known to hold or to have held desert bighorn
evaluated a trait-climate relationship among natives and populations in California, and then scored these for variables
invasives in California’s grasslands (2012). With increased that described climate, metapopulation dynamics, human
mean annual temperature, they expected to see increased impacts, and other environmental factors (Fig. 4.10). Climate
potential evaporation, increased temperature seasonality and factors such as temperature and moisture affect annual nutri-
increased water deficits in areas without substantial increases ent availability in forage plants used by bighorns. However,
in precipitation (Sandel and Dangremond 2012). The authors Epps et al. used elevation as a surrogate variable for temper-
examined relationships between specific plant functional ature because, in this environment, the two are strongly
traits and climate variables (e.g., increasing leaf width with correlated, and elevation can be measured much more pre-
increasing temperatures) to derive predictions for interactions cisely at individual sites used by bighorns.
between climate change and invasives at a large scale,
Fig. 4.9 Changes in the exotic

grass flora of California over time.
For any year, each exotic species
was deemed present in the state if
there is an herbarium record of
that species before that year,
otherwise it was deemed absent.
The number of exotic species in
the state continues to increase into
the twenty-first century (a). Later
arrivals in California were more
likely to be perennial and C4,
leading to increasing proportions
of these groups within the exotic
flora over time (b and c). The
earliest detected exotic species in
California had small seed sizes.
Later introductions increased
mean seed size rapidly, until about
1920. (Reprinted by permission
from Wiley, from Sandel, B. and
Dangremond, E. M. (2012),
Climate change and the invasion
of California by grasses. Global
Change Biology, 18:
277–289. # 2011 Blackwell
Publishing Ltd)
Classic metapopulation theory predicts that extinction to become extinct. The best predictive models contained the
probability should decrease with increasing area of habitat parameters of maximum average annual precipitation, maxi-
patches available for population subunits (Hanski 1991, mum elevation, presence of domestic sheep grazing
1997) and with increasing immigration of individuals from allotments, and presence of dependable springs. Extinction
one population subunit to another, which in turn, depends on was, in the words of Epps and his colleagues, “negatively
interpatch distance. With these traditional assumptions in correlated with precipitation, elevation, and dependable
mind, Epps et al. tested two hypotheses to determine if springs, but positively correlated with the presence of
extinctions were linked more strongly to the processes of domestic-sheep grazing allotments” (Epps et al. 2004:108).
metapopulation dynamics than to overriding climate In other words, for a given population, the more precipitation,
conditions. Based on the previously stated predictions, Epps the greater the elevation, and the more dependable springs
et al.’s first hypothesis was that extinct populations inhabit present, the lower the probability of extinction. But the
ranges with smaller two-dimensional areas than ranges with greater the presence of domestic sheep, the greater the proba-
extant populations. Using similar reasoning, the second bility of extinction.
hypothesis was that extinct populations were more isolated Recognizing the implications of these findings in terms of
from other mountain ranges containing bighorn sheep than ongoing and projected future climate changes in this region,
are extant populations. (Epps et al. 2004). Thus, these Epps et al. wrote, “. . .in the maximum temperature-change
investigators also measured the size of the area used by scenario of +2.0 C in the next 60 years, average risk of
each population subunit (“patch size”) and its relative isola- extinction increased substantially to 0.26. Extinction risk
tion from other subunits (mean distance to other subunits), also increased drastically when precipitation was reduced,
along with parameters related to geology (which affects both such that a 0.7 C increase combined with a 12% decrease
vegetation and water availability) and various aspects of in precipitation elevated extinction probabilities to levels
potential human disturbance. All parameters were then observed with a 2.0 C increase with no change in precipita-
incorporated in logistic-regression models that evaluated the tion. Average extinction risk increased from 0.21 (no change)
strength of each parameter in correctly predicting population to 0.30 when a 2.0 C increase was combined with a 12%
persistence. precipitation decrease” (Epps et al. 2004:108–109). Further,
In the models that resulted, all parameters that were “Populations in mountain ranges of lower elevation were
climate-related or climate-dependent (elevation, precipita- much more likely to become extinct, particularly at
tion, and the presence of dependable springs) were strongly <1500 m. Populations in regions with the lowest annual
correlated with population persistence. Bighorn populations precipitation, especially <200 mm annual precipitation,
that inhabited lower, drier mountain ranges were more likely were also more likely to become extinct, as were populations
Fig. 4.10 Native, relocated, and

extinct populations of desert
bighorn sheep (Ovis canadensis
nelsoni) in California. Using
presence and absence data, the
authors demonstrated that
population extinctions of desert
bighorn sheep in the twentieth
century are consistent with a range
contraction to areas of higher
elevation and greater
precipitation. (Reprinted by
from Epps, C.W., McCullough,
D.R., Wehausen, J.D., Bleich, V.
C., Rechel, J.L., 2004. Effects of
Climate Change on Population
Persistence of Desert-Dwelling
Mountain Sheep in California.
18, 102–113. # 2004 Society for
Conservation Biology)
without dependable springs and populations in which expected correlation with population size if populations are
domestic-sheep grazing allotments formerly overlapped or strongly regulated by density dependence. If populations are
abutted desert bighorn habitat. This suggests not only that regulated by environmental factors, however, one can expect
desert bighorn sheep are vulnerable to climate warming, but a much weaker relationship between patch size and popula-
that climate warming has already affected their distribution in tion size. . .Our findings that precipitation and elevation, but
California” (Epps et al. 2004:109). not patch size, were correlated with population extinction are
Recall that Epps and his colleagues had hypothesized, consistent with strong environmental regulation of desert
based on predictions of metapopulation theory, that popula- bighorn sheep populations.” (Epps et al. 2004:110).
tion subunits using smaller areas and that were more isolated
from one another should be more prone to extinction. This 4.2.4.2 Declining Health and Local Disappearance
was not the case. Population extinction was not sensitive to of Coral Reefs
patch size (two-dimensional area of the inhabited mountain Climate change in oceans causes: (1) changes in calcium
ranges) or to the degree of population isolation. But why, carbonate saturation state (pH), (2) changes in sea level,
exactly, was a climate effect able to override a fundamental and (3) changes in temperature of ocean water. As Ova
population process that has been demonstrated to be such a Hoegh-Guldberg, a leading expert on marine biodiversity
strong influence on metapopulation persistence in other stud- puts it, “. . .not all of these changes (in isolation) are likely
ies? Epps et al. give this explanation, “The strong effect of to have a negative impact on marine biodiversity. The com-
patch size on persistence is thought to result from the bination of these changes, however, is expected to drive
major changes on the distribution and abundance of marine 4.2.4.3 Disappearing Ecosystems and the Species
organisms” (Hoegh-Guldberg 2005:258). The damage and That Rely on Them
death of coral reefs is significant to world marine C. A. Enquist examined the effect of climate change on
biodiversity (Chap. 8). The biodiversity of coral reefs world- elevational life zones (“Holdridge Life Zones”) in Costa
wide is at estimated 1 million species of plants, animals and Rica, estimating the species richness and number of endemic
protists, all of these living within a total of only 400,000 km2 species in every Holdridge Life Zone by direct field
of coral reefs (Hoegh-Guldberg 2005:262). But coral reefs measurements (Enquist 2002:519). Of all the available
have undergone major changes in the past 35 years as a result types of ecosystem classification, Holdridge Life Zones are
of climate change, and all to the detriment of their uniquely suited to evaluations of climate change on
biodiversity. ecosystems. The Holdridge Life Zone system includes
The most common response in corals to increased ocean 38 life zone categories defined by plant moisture demand
water temperature is called coral bleaching. Although corals and biotemperature (Fig. 4.11). Biotemperature, in this clas-
have been studied for over a hundred years, bleaching was sification system, is defined as the mean value of daily
never observed until the mid-1970s. Hoegh-Guldberg gives a temperature above 0 C divided by 365. Thus, as Peterson
precise description of this phenomena: “coral bleaching et al. put it, “Biotemperature, which is closely related to
occurs when corals rapidly lose the cells and/or the pigments growing degree days, gives a measure of the heat available
of symbiotic dinoflagellates (zooxanthellae) that populate in during the growing season and is likely to be more directly
their tissues by the millions. Bleaching results in colonies related to plant growth than is mean annual temperature”
turning from brown to white, often with spectacular host (Peterson et al. 2005: 219).
pigments (pocilloporins. . .)” (Hoegh-Guldberg Using current and projected levels of temperature and
2005:263–264). The most recent bleaching events have precipitation from different climate models and Holdridge
been extreme with 45% of the coals bleached in some life zone criteria, Enquist predicted how such zones would
locations in Hawaii and 93% of surveyed reefs on the Great change elevationally under different climate scenarios, and
Barrier Reef with >50% coral mortality observed (Van how this might affect species richness and numbers of
Oppen et al. 2017). Although some scientific observations endemic species in each zone. He noted that “High elevation
of reefs in the Indo-Pacific have seen a small increase in life zones were shown to be more sensitive to changes in
bleaching tolerance (Penin et al. 2013), the >40% loss of temperature, while lower elevation life zones tended to be
the world’s coral reefs over the past four decades has more sensitive to changes in precipitation. Regional life zone
indicated that the rate of temperature increase is outpacing diversity was greatly reduced in an extreme wet and warm
the natural rate of evolution of thermal tolerance in corals climate scenario. Three elevation-associated life zones (lower
(Van Oppen et al. 2017). montane rainforest, montane rainforest, and premontane
Warming of ocean temperatures might suggest that the rainforest) ranked in the top four in percentage number of
distribution of corals will simply shift northward, as has been endemic species. The lowland seasonally dry forest life zone
the case with many terrestrial species. But corals are ulti- ranked second in this group, suggesting that this life zone has
mately light-limited because of their symbiotic relationship a unique species composition in comparison with other low-
with photosynthetic algae, zooxanthellae. Thus, “however land Holdridge life zones. Of the 19 life zones, these four life
wonderful this scenario may sound (coral reefs off zones displayed particular sensitivity to the climate changes
New York or Sydney), corals are ultimately limited by light modeled here” (Enquist 2002:519).
levels and possibly carbonate alkalinity (which decreases in a The most sensitive of these life zones may be the Central
poleward direction). These factors are likely to limit coral American cloud forests (Fig 4.11b). Cloud forests, such as
reefs to small changes in their latitudinal range” (Hoegh- those in Costa Rica, are distinctive not only in the amount of
Guldberg 2005:268). Analyses of current coral bleaching rainfall they receive, but in the historically “misty” environ-
reveal that “thermal events that exceed 8 degree-heating ment that predominates in them. Usually found at the upper
weeks have almost always (99% of the time) resulted in elevations of high tropical mountains, it was historically rare
coral bleaching. If conditions improve, bleached corals will for such systems to experience consecutive mist-free days.
recover their symbionts and hence their brown color. If Amphibians, the one class of vertebrates that depends on
heating continues, however, and the degree-heating-weeks direct respiration through their skin as much as on gills or
attains values of 13 or more, the event is likely to result in lungs, must keep their skin moist in order for such respiration
large-scale coral mortality. Coral dominated ecosystems may to occur. Cloud forests provide an environment of relatively
become remnants of the past if sea temperatures continue to constant high humidity that could be uniquely beneficial to
climb” (Hoegh-Guldberg 2005:269–270). amphibians through its enhancement of this respiration path-
way. Amphibian species richness in cloud forests has histori-
cally been large and contained such charismatic endemic
Fig. 4.11 (a) First published by Leslie Holdridge in 1947, and updated (PET). (b) Monteverde Cloud Forest in Costa Rica. The Holdridge life
in 1967, the Holdridge life zones system is a bioclimate scheme for zone system was originally developed for tropical and subtropical
identifying regions along the three axes of precipitation, biotemperature, regions. (Part A courtesy of P. Halasz, reprinted under CC-BY-SA-
and potential evapotranspiration ratio to mean total annual precipitation 2.5. Part B courtesy of R. Lamb)
amphibians as the golden toad (Bufo periglenes) among over decades of field work and described mechanisms
others. through which climate change alters species composition.
Alan Pounds and his colleagues, who have long studied Based on direct measurements, they noted, “Dry days have
cloud forest amphibians, reported changes they have seen increased in frequency since the 1970s and have increasingly
coalesced into dry periods. Whereas mist free periods in the its ecosystem structure. Many marine mammals such as seals,
1970s rarely exceeded two days, they have recently lasted up sea lions, and walruses need access to open water to feed. At
to three weeks” (Pounds et al. 2005:71). The outcomes of the same time, they require ice in proximity to such open
such changes were predictable. “Massive declines of frogs water of sufficient thickness to support adults and young,
and toads were apparent by 1990. A multispecies population who are weaned in such ice areas close to open water. The
crash in 1987 led to the disappearance of the endemic golden primary predator of such mammals in the Arctic, the polar
toad (Bufo periglenes) and many other species. . .Twenty of bear (Ursus maritimus), also requires large areas offering this
the 50 [species] were missing throughout the surveys of a juxtaposition of ice and open water in order to find sufficient
30-km2 area during 1990–1994, and there is still little sign of numbers of prey. In the Arctic’s western Hudson Bay area,
recovery 15 years after this crash” (Pounds et al. the ringed seal (Phoca hispida) has shown a pattern of
2005:70–71). declining recruitment from 1990 to 2001 coincident with a
Pounds et al. evaluated the probability that such climate pattern of declining snowfall, lower snow depth,
disappearances might be random events. Summarizing their warmer April–May temperatures and earlier breakup of
analysis, they noted, “Tests of null models based on long- spring sea ice (Ferguson et al. 2005).
term studies of other amphibian assemblages suggest that the If this pattern of decreased recruitment continues, the
number of disappearances is improbable in the context of ringed seal, a principle prey of the polar bear, could see
normal demographic variability. Moreover, surviving significantly smaller populations in the coming decades.
populations for which baseline data exist have fluctuated far The polar bear is remarkable among all mammals for its
below crash levels, undergoing simultaneous downturns in ability to fast for extended periods, not only during its winter
1994 and again in 1998” (Pounds et al. 2005:71). denning period (during which all bears fast), but during
As the humidity of cloud forests declined, other species of non-denning periods when it is active. In Hudson Bay, for
amphibians and other animals, traditionally found on lower example, polar bears can prey effectively on marine
slopes, colonized higher elevations. But the cloud forests mammals like the ringed seal only while ice is present adja-
could go no higher. As Noss concluded, summarizing other cent to open water. Ice break up in Hudson Bay has histori-
studies, “Simulations of changes in temperature and moisture cally occurred in late June, with the entire ice melt completed
under doubled CO2 show an upward shift in the cloud layer by late July, and polar bears then fast for 4 months until the
of hundreds of meters during the winter dry season, coupled fall ice freeze up in early November. In recent years
with increased evapotranspiration... Cloud forests have (1962–2000), spring ice break up has advanced from
nowhere to shift and are expected to be lost, along with 23 June to 30 May, with all ice gone by mid- to late June,
their endemic species...meanwhile, species from lower adding 4–6 weeks of fasting to the polar bear’s already
elevations have invaded these forests... In situations such as challenging energetic regime.
these, ex situ preservation of species in zoos and botanical The loss of sea ice has serious consequences for all polar
gardens until global warming is reversed may be the only bears, but it is particularly detrimental to females with cubs.
way to avoid extinction” (Noss 2001:586). Although female polar bears regularly make winter dens on
land where they give birth to cubs, they also commonly make
4.2.4.4 Loss of Arctic Sea Ice: The Case such dens on ice adjoining land (so-called “land-fast ice”) as
of the Endangered Polar Bear well as on drifting sea (pack) ice. Historically, pregnant
Based on predictions of global climate models, the IPCC has females often choose to den on ice where the risk of human
stated that temperature increases associated with global cli- disturbance of the den is less, but unstable ice will often lead
mate change will be disproportionately higher in polar areas, on-ice denning attempts to fail. Additionally, the ice must be
particularly in northern latitudes (IPCC 2001, Räisänen covered with sufficient snow depth so that the female polar
2001). The IPCC posits that, by the end of the twenty-first bear can construct an adequate den to nurse and protect
century, northern high latitude regions will experience her cubs.
temperatures at least 40% greater than the global mean, and Anthony Fischbach and his colleagues at the US Geologi-
the central Arctic area will experience temperatures 100% cal Service Alaska Science Center monitored 99 radio-
higher than increases in the global mean, an average increase collared female polar bears and documented 142 denning
of 3–4 C (IPCC 2001). One impact of increasing events in northern Alaska and the Beaufort Sea from 1985
temperatures in the Arctic is the loss of sea ice, with at least to 2013 using satellite telemetry (Olson et al. 2017,
one sea-ice-free Arctic summer expected every 10 years with Fig. 4.12). In the 1980s and 1990s, maternal denning
a global warming of 2 C (IPCC 2018). occurred primarily on sea ice (67% of the time). Between
These predicted events are now being observed. 1985 and 2005, denning substrates shifted such that 67% of
Reductions in extent and thickness of sea ice are significant maternal denning occurred not on ice but on land by the end
to polar ecosystems because sea ice is a critical component of of this period (Fischbach et al. 2007). This decline parallels
Fig. 4.12 Denning locations of

female polar bears (Ursus
maritimus) in the southern
Beaufort Sea between 1985 and
2013. Denning was identified
based on temperature data
collected by satellite radio collars.
The polar bear has a range that
extends from Icy Cape, Alaska,
USA (159 W), to Tuktoyaktuk,
Northwest Territories, Canada
(133 W), with a northern
boundary of approximately 74 .
The Beaufort Sea is nearly 100%
ice- covered from November to
June. Since 1996, declines in
summer sea ice have caused the
ice edge to retreat north beyond
the narrow (~100 km) continental
shelf and into the deep waters of
the Canada Basin. Most SB bears
follow the retreating ice north,
while a proportion (17.5% in
2000–2013, 27% in 2007–2010)
of the subpopulation moves to
land. (Reprinted by permission
from US Geological Survey
from Olson, J., and others. (2017).
Collar temperature sensor data
reveal long-term patterns in
southern Beaufort Sea polar bear
den distribution on pack ice and
land. Marine Ecology Progress
Series, 564, 211–224)
declines in multi-year ice, and a reduction in the extent of body condition and reproduction due to fasting. The Beaufort
summer pack ice (Olson et al. 2017). Over the final 8 years of Sea population of polar bears has previously been able to
the study, researchers found that while land denning spend their entire summers on sea ice, which historically
continued to increase across years, this trend was only mar- never completely melted. Under these conditions, bears
ginally significant. However, looking at the relationship could forage efficiently for long periods and store large
between land denning and increasing distance between the amounts of fat in preparation for winter. But recent recessions
coast and suitable sea ice concentrations, researchers found and losses of sea ice in the Beaufort and other Arctic seas
that for every 100 km increase in the distance to 50% sea-ice force polar bears out over much deeper waters with lower
concentration, the frequency of denning on land increased on productivity, forcing them to swim much longer distances to
average by 32% (Olson et al. 2017). Furthermore, bears that adjacent ice flows. These events can contribute to a negative
spent substantial time (>25 days) on land before denning energy balance in polar bears or leave them stranded on shore
were highly likely to den on land whereas only 29% of where there is less prey available. Under these conditions, the
those that remained on ice, came back to land for denning, authors concluded that “most Beaufort Sea bears probably
perhaps suggesting that polar bears may have had a difficult are not fat enough in mid-summer to fast for months and then
time reaching land for denning due to the distance between den successfully without first returning to sea ice to feed. . .
summer sea ice and land (Olson et al. 2017). bears occupying the polar basin may be less capable of
Assessing these results and others, the authors conclude maintaining long-term reproductive viability by denning on
that an increase in bears summering and denning on land in land after a prolonged summer and autumn fast. Whether
response to sea-ice declines could have implications for they are forced onto land or far offshore by the recent changes
human-bear interactions, substantially increase energetic in sea ice, polar bear foraging opportunities in the Alaska
costs with long-distance swims, and result in declines in region appear reduced from earlier times.” (Fischbach et al.
4.3 Foundational Tools for Assessing Future Climate Impacts 149
2007:1403). Thus, it is now not surprising that biologists and used within ecological niche models, also referred to as
other observers are seeing reduced physical size in adult polar climate-envelope models, which utilize statistical
bears and reduced fat deposits in both adult polar bears relationships between climate variables and patterns of spe-
and cubs. cies distribution to map how a species’ preferred “environ-
mental space” moves across the landscape given anticipated
climate changes. Although an important tool for
4.3 Foundational Tools for Assessing Future characterizing climate exposure, they are limited in their
Climate Impacts ability to estimate extinction risk or delineate suitable future
habitats (Botkin et al. 2007).
4.3.1 Integrated Vulnerability Assessments The second component of vulnerability is sensitivity,
for Biodiversity defined as “the degree to which the survival, persistence,
fitness, performance, or regeneration of a species or popula-
Accurately anticipating the ongoing consequences of climate tion is dependent on the prevailing climate, particularly on
change on biodiversity requires a range of scientific tools and climate variables that are likely to undergo change in the near
techniques. In an “integrated science of climate-change bio- future” (Dawson et al. 2011:53). Here, the focus is on
diversity assessment” authors Dawson et al. draw from mul- species-specific factors that may indicate a higher level of
tiple lines of evidence to better characterize species risk relative to expected climate changes due to intrinsic
vulnerability as a prerequisite for developing effective attributes or traits that either moderate or exacerbate the
climate-informed conservation strategies (2011). Reviewing impact of climate pressures (Foden et al. 2019). As examples,
paleoecological observations, recent phenological and micro- particular life histories, ecophysiology or microhabitat
evolutionary responses, experiments and computation preferences can influence how well a species may tolerate
models, the authors submit that “heavy reliance on a single anticipated changes, with the most sensitive species showing
scientific approach creates risks of policy or management the greatest reductions in survival or fecundity under small
failures” (Dawson et al. 2011:54) (Fig. 4.13). As a mecha- changes in climate conditions. This second factor of vulnera-
nism for describing how different scientific approaches can bility is important for management since species across the
work together to address multiple attributes of climate- same spatial domain may exhibit varying degrees of sensitiv-
change assessment, the authors define overall vulnerability, ity to this same degree of exposure. Furthermore, climate
“the extent to which a species or population is threatened change pressures may operate inconsistently across a species’
with decline, reduced fitness, genetic loss, or extinction range and evidence differential effects within both
owing to climate change,” (Dawson et al. 2011:53) as a individuals (changes in life history, physiology, morphology)
function of three primary components: exposure, sensitivity and subpopulations (altered abundance and metapopulation
or adaptive capacity. This definition follows the one offered dynamics) (Foden et al. 2019).
by the IPCC in their fourth climate assessment (2007) and has Finally, vulnerability should also be understood as a func-
been widely adopted in the conservation literature despite a tion of adaptive capacity. Dawson et al. define adaptive
more recent change in terms offered in the fifth IPCC assess- capacity as “the capacity of a species or constituent
ment (2014) (Foden et al. 2019). populations to cope with climate change by persisting in
Recent climate change assessments have largely focused situ, but shifting to more suitable local microhabitats, or by
on clarifying a single component of vulnerability, exposure. migrating to more suitable regions” (2011:53). Utilizing
Dawson and his colleagues define exposure as “the extent of empirical, observation, and modeling studies, scientists
climate change likely to be experienced by a species or have worked to characterize the adaptive capacity of a
locale” (Dawson et al. 2011:53). Here, the rate or magnitude given population or species based on a range of intrinsic
of change provides an indication of how disruptive this factors including phenotypic plasticity, dispersal ability, col-
change will be to an ecosystem. The drivers of these climate onization ability or “evolvability” associated with genetic
changes are societal demands and needs (social, economic diversity (Foden et al. 2019). In practice, this work is chal-
and political) which result in associated changes to green- lenging to project given interactions between intrinsic and
house gas emissions (Foden et al. 2019). A range of potential extrinsic factors (i.e., a species with intrinsic high dispersal
future changes, or climate scenarios, are often developed via capacity surrounded by a landscape with inhospitable
Global Climate Models (GCMs) and can be downscaled to conditions). Given the diversity and number of species
highlight abiotic pressures or resulting effects on the physical under threat, it is important to both mine existing literature
environment such as increased temperatures or altered for clues about inherent capacities as well as focus on creat-
drought frequencies. These climate scenarios can also be ing empirical models that can estimate a range of ecological
Fig. 4.13 An integrated science of climate-change biodiversity assess- conservation and adaptation. (Reprinted by permission from The Amer-
ment will draw from multiple sources and approaches. Each provides ican Association for the Advancement of Science, from Dawson, T.P.,
useful but incomplete information on exposure, sensitivity, and adaptive Jackson, S.T., House, J.I., Prentice, I.C., Mace, G.M., 2011. Beyond
capacity. Integration of these approaches will provide a more robust Predictions: Biodiversity Conservation in a Changing Climate. Science
basis for vulnerability assessment and allocation of resources for 332, 53–58. # 2011 AAAS)
and evolutionary responses to forecasted environmental etc.)? What is timeframe for this assessment (given needs of
change. site managers, species generation times, intervals of available
Given that there is no one-size fits all approach to climate projections, etc.) (Table 4.3)? Once defined, these
mitigating risk, a combination of approaches crossing all goals can help scientists generate a list of pressures to which a
three components of vulnerability is important. Furthermore, species might be exposed, and identify likely impacts at
as we discuss throughout this chapter, conservation managers individual and subpopulation levels in consultation with
will have to actively work to adapt traditional conservation experts and the literature (Foden et al. 2019). Those
tools to meet the new demands of climate-change assessment conducting the assessment should consider both the full
strategies. However, the result of this effort is not only to range of climate change pressures as well as interactions
better identify and address unique climate threats, but to between climate change and other forms of global environ-
inform management decisions, which will, as Dawson et al., mental change such as habitat loss and fragmentation
write “depend on judgements of potential risks and benefits (Chap. 7). The final step in the process often requires increas-
balanced against costs and available or anticipated ing data complexity and higher resource requirements to
resources. . . the assessment should aim to maximize the evaluate the species’ sensitivity and adaptive capacity using
likelihood of the desired management outcome, minimized trait-based, correlative and mechanistic approaches. Here, the
the financial costs, and assess associated risks” (2011). IUCN Red List (Chap. 2) may provide valuable information
The first step in any vulnerability assessment should be to on demographic and behavioral characteristics. Utilizing a
define goals and objectives by answering a series of decision matrix, Dawson et al. (2011) highlight how a bal-
questions: What is the taxonomic focus (species, subspecies, ance of consideration among all three components of vulner-
metapopulations, etc.)? What is the spatial focus of analysis ability in service of such an assessment could directly
(single site, network or sites, a political unit such as a state, influence the intensity of management response (Fig. 4.14).
Table 4.3 Framework for describing the objectives of a climate change vulnerability assessment in clear and certain terms
Select an objective Select a time
category Select a taxonomic focus Select a spatial focus frame
Which? Subpopulation Single site Present
How much? Species Network of sites 5 years
Who? Higher taxonomic group Range of a subpopulation 20 years
Where? Multiple higher taxonomic Entire range of taxon/taxonomic group 50 years
When? groups Politically defined geographical area (e.g., national, continental, 100 years
What’s missing? global, etc.)
Reprinted by permission from Wiley, from Foden, W.B., Young, B.E., Akçakaya, H.R., Garcia, R.A., Hoffmann, A.A., Stein, B.A., Thomas, C.D.,
Wheatley, C.J., Bickford, D., Carr, J.A., Hole, D.G., Martin, T.G., Pacifici, M., Pearce‐Higgins, J.W., Platts, P.J., Visconti, P., Watson, J.E.M.,
Huntley, B., 2019. Climate change vulnerability assessment of species. Wiley Interdisciplinary Reviews: Climate Change 10:e551. # 2018 Wiley
Periodicals, Inc
Fig. 4.14 The vulnerability of a species or ecosystem is based on its axis. This axis is primarily determined by biological characteristics of
exposure to climate change, its sensitivity, and its inherent capacity to species that influence their mobility, specificity, and sensitivity (intrinsic
adapt to change. The relative balance of these different components of factors). These include, for example, physiological constraints, pheno-
vulnerability would lead to different management interventions. The x typic plasticity, evolutionary potential, dispersal and growth capacity,
axis represents the degree of exposure to climate change faced by and biotic interactions critical to persistence. The relative position of
species and communities. This axis is largely determined by the species’ species and ecosystems along the axes can inform decisions on appro-
or population’s geographical location, the rate and magnitude of climate priate research, monitoring, and management strategies. (Reprinted by
change anticipated for that region, and the size, cohesiveness, and permission from The American Association for the Advancement of
connectivity of the species’ habitat within and beyond that region Science, from Dawson, T.P., Jackson, S.T., House, J.I., Prentice, I.C.,
(extrinsic factors). The other two measures from the vulnerability frame- Mace, G.M., 2011. Beyond Predictions: Biodiversity Conservation in a
work, adaptive capacity and sensitivity, are plotted together on the y Changing Climate. Science 332, 53–58. # 2011 AAAS)
Climate and ecosystem modeling are central tools within 4.3.2 The Bioclimate Envelope: The Correlative
many vulnerability assessments. Although empirical and Approach to Modeling Climate Effects
observational data, and experimental manipulations provide on Individual Species
important calibration and validation for such models, conser-
vation and land-use planning requires information on the Half a century ago ecologist G. Evelyn Hutchinson defined
future. In the following sections we review a range of the fundamental ecological niche as consisting of those envi-
modeling tools used by scientists to better understand the ronmental conditions under which a species can survive and
implications of ongoing climate change for species, grow (Hutchinson 1957). This so-called fundamental, or
populations, and evaluate the conservation strategies we are “Hutchinsonian niche” has been contrasted, in every text-
using to protect them. book on general ecology and elsewhere, with the so-called
“realized niche,” that part of the fundamental niche the spe- biodiversity: the use of field experiments to determine
cies actually occupies when constrained by the presence of correlations, physiological tolerances, and effects on
and interactions with other species. biological timing associated with climate change that could
Part of the tolerable environmental conditions that com- then be used to estimate the climatic, especially thermal,
prise the fundamental niche, as shown earlier in the work of tolerances of a given species. Studies like Saavedra’s form
Joseph Grinnell, are climatic in nature. Thus, every species the bricks and mortar for building an understanding of a
has a “climate component” of its fundamental niche, or what species’ climate niche. The second step is to couple this
could be referred to more directly as its climate niche. His- work with atmospheric models that can project future tem-
torically, biologists have seen, and some have carefully stud- perature changes within the species’ range. If one can pre-
ied, the tendency of every species to track its climate niche cisely define the climate niche of a species and predict how
through space and time in the face of changing climate the climate will change over time within its historic range,
conditions. For example, bird migration is one form of cli- one should be able to project what will happen to the distri-
mate niche-tracking that is employed to cope with seasonal bution of a species as climate changes, and whether a species
climatic variation. would have to migrate in order to remain within its climate
Initially, interest in a species’ climate niche, and the effect tolerances (Sanford 2002).
of climate change on a species, was investigated through Scientists first integrated these data and concepts into what
small scale experiments that permitted investigators to make has come to be called the bioclimate envelope of a particular
direct measurements to determine a species thermal toler- species (Pearson and Dawson 2003). Bioclimate envelope
ance, or how its biological rhythms and events might be models (also known as ecological niche models, habitat
altered in altered climatic conditions. For example, biolo- suitability models, or species distribution models) generally
gist Francisca Saavedra demonstrated how the frequency work by discriminating between the climates of locations
and date of flowering could be advanced in an alpine wild- inside and outside the ranges of a given species based on
flower, the two-lobe larkspur (Delphinium nuttallianum) by particular climate variables, such as minimum and maximum
removing the snow from the soil surface where it germinated temperature (Table 4.4). This strategy of modeling is techni-
(Saavedra 2002). cally known as a “correlative approach,” to characterize the
Saavedra’s results are intuitive, but these and many other statistical relationship between between observed species
such studies have formed the foundation for studying and distributions and environmental variables (derived primarily
modeling the effects of climate change on the Earth’s from climatic data but including a measure of soil-type) (e.g.,
Table 4.4 Several designations in the literature about correlative models exploring the relationship between species occurrences and environmental
predictors, and their implications for interpretation of the results
Designation Implications
Bioclimate or species-climate Term expressing that a multivariate space of climatic variables (the envelope) best matching observed
envelope models species’ distributions is being estimated. The term “envelope” also has been used to refer to multivariate
approaches using presence-only data, but all models constructing a multivariate space of predictor variables
can be said to generate an envelope. The words “climate” or “bioclimatic” can be limited as species may
relate to other environmental covariates; more general terms, such as “abiotic” or “environmental envelope”
overcome this limitation.
Ecological or climatic niche Expresses ideas analogous to those of “environmental envelope models,” but instead of providing
models descriptive terminology, an attempt is made to link the envelope to elements of ecological niche theory
rooted in the early work of Grinnell and Hutchinson. The link between description of pattern and niche
theory has raised controversy, but progress is being made toward consensus.
Habitat-suitability models Related to “envelope” or “niche” because it refers to the suitability of areas for species rather than actual
distributions. “Habitat” emphasizes the physical space where species live and the resources it can use. As
such, it opens the door for incorporation of resource variables and biotic factors that are often absent from
models. The term is mostly used by researchers working at landscape scales because their resolutions are
those at which such connections are more easily achieved.
Species-distributions models Term implying that the geographic distribution of the species is the quantity modeled. Most such
applications, nonetheless, characterize the multivariate environmental space delimiting species’
distributions, and then project this subset of environmental space back onto geography. Important
mechanisms, such as species dispersal, establishment, and biotic interactions are not accounted for. If only
suitability is modeled, a species-distribution model (termed as such) will estimate something more closely
related to the potential distribution and will be exposed to criticism of being based on implausible
assumptions and often contradicted by empirical evidence
Reprinted by permission from Wiley, from Araújo, M.B., Peterson, A.T., 2012. Uses and misuses of bioclimatic envelope modeling. Ecology
93, 1527–1539. # 2012 Ecological Society of America
Fig. 4.15 A hypothetical example of how different data layers could be (green bands). (Reprinted by permission from Natural Areas Associa-
combined to generate composite maps for different conservation tion, from Schmitz, O.J., Lawler, J.J., Beier, P., Groves, C., Knight, G.,
approaches. The composite maps along the bottom depict biodiversity Boyce, D.A., Bulluck, J., Johnston, K.M., Klein, M.L., Muller, K.,
hotspots, large intact natural landscapes (area surrounded by dashed Pierce, D.J., Singleton, W.R., Strittholt, J.R., Theobald, D.M.,
white line), geophysical settings that provide the greatest heterogeneity Trombulak, S.C., Trainor, A., 2015. Conserving Biodiversity: Practical
(area outlined by dashed black line), climate refugia (red areas), and Guidance about Climate Change Adaptation Approaches in Support of
conservation corridors that maintain and restore ecological connectivity Land-use Planning. Natural Areas Journal 35, 190–203)
Pearson and Dawson 2003). Conceptually, the model is cre- range limits of the European holly (Ilex aquifolium) were
ated through a series of spatial overlay maps, each with consistently associated with the 0 C degree winter isotherm,
different attribute data. For example, when precipitation and and this species’ range were shifting distributions northwards
temperature data are combined across space, these geo- over time as they were tracking increasing temperatures at
graphic domains can begin to shape distinct climatic higher latitudes.
ecoregions across the landscape and become coupled with Establishing clear associations between climate and spe-
other input data layers to identify spatial areas of interest cies distribution can be challenging due to incomplete data or
(Fig. 4.15). spatial data utilized at inappropriate resolution and extent.
The most important theoretical assumption of these For example, at a continental scale, climate considerations
models is that species distribution in its broadest sense is will be dominant, such that only those areas where the
determined wholly or partly by climate. Two pieces of sup- species’ climate tolerances match actual climate conditions
port for this assumption are that: (1) distributional limits of can be considered potential occupiable range. However, that
species match particular combinations of climate variables, consideration alone may not produce accurate predictions of
and (2) these limits shift through time in synchrony with future species occurrence. At regional and landscape scales,
changes in climate (Araújo and Peterson 2012). For example, topography and vegetation (habitat) may become a more
a study by Walther et al. (2005) showed that the northern critical detector of presence or absence. At even finer scale
resolution, one might examine biotic interactions and micro- 4.3.3 Dynamic Global Vegetation Models:
climate conditions, such as slope aspects (south or north), Process-Based Approaches to Modeling
soil conditions (dry or hydric), or geologic factors (such as Species Response to Climate
limestone, serpentine, or granitic rock bases) that could limit
the occurrence of the species. Accurate modeling of land- Models of global climate change provide broad resolutions of
scape and topography is essential because “the ability to future climate change worldwide. Several General Circula-
migrate is a function not only of individual species’ tion Models (GCM) are now accessible through the internet,
characteristics, but also the structure of the landscape over including downscaled climate projections such as the NASA
which dispersal is occurring, including the presence of natu- Earth Exchange (NEX) based off of the IPCC’s greenhouse
ral barriers (such as mountain ranges) or the artificial frag- gas emissions scenarios known as Representative Concentra-
mentation of habitats (through, for example, the growth of tion Pathways (RCPs). With the right skills and software, a
urban areas or deforestation)” (Pearson and Dawson single scientist, working at a personal computer with suffi-
2003:3765). cient computing power, can now compare simulated results
Using models of bioclimate envelopes to predict future from multiple models. Regional climate models are often
species ranges carries additional risks. If the model’s only embedded within GCMs to provide higher resolution results
information is that of climate projections and species climate for particular geographic areas. These models focus on the
tolerances, it will track the potential future range of the transfer of energy between the atmosphere, ocean, land-
species, which will almost certainly be larger than its actual surface and ice.
future range, just as a species’ fundamental niche is almost Bidirectional interactions between terrestrial vegetation
always larger than its realized niche. Araújo and Peterson and atmospheric climate conditions determine the dynamic
discuss potential methods to address this challenge via equilibrium state of the system (Quillet et al. 2010). How-
experiments (e.g., Kearney et al. 2010; Buckley et al. ever, most global and regional climate models represent
2011), but note that this approach is not practical for most vegetation as a static component of the climate system. For
species, which will “frequently be too poorly known to the past two decades there has been a considerable scientific
permit first-principles approaches or will prove intractable effort to better represent these interactions dynamically
for experimentation” (2012). Assumptions and extrapolation through the construction of dynamic global vegetation
must be handled carefully and named explicitly in all models (DGVM). DGVMs provide insights into potential
modelling efforts. A second critique is that bioclimate enve- vegetation response to future climate change, either through
lope models assume that a species would either fail to dis- the effect of climate change on successional processes (e.g.,
perse or disperse without limitations, and essentially ignore forest-gap models) or by assessing tolerances of plant
other biotic interactions that shape current and future species communities or individual species to different climate
distributions (Pacala and Hurtt 1993; Hurtt and Pacala 1995). conditions. Foundationally, these models assess the impact
Although an important known limitation, some scientists of variables such as photosynthesis, respiration, surface
argue that these assumptions are not integral to model func- energy fluxes, and carbon and nutrient allocation within the
tionality as they aim to estimate suitable conditions across plant (Foley et al. 2000; Peng 2000). To facilitate model
landscapes, and not population processes such as dispersal parametrization, DGVMs typically use a number of plant
(Brook et al. 2009; Araújo and Peterson 2012). functional types (PFTs) to describe vegetation. Although
Despite these challenges, bioclimate envelope models this limited number of PFTs may not be representative of
remain one of the principal forecasting tools used for conser- the high diversity of tree species found in the world’s tropical
vation planning. Using knowledge of the climatic tolerances forests, it allows for global spatial coverage (Powell et al.
of individual species, such models try to predict where certain 2013; Schimel et al. 2015).
species, especially plants, could occur in the future under With dynamic coupling, scientists can more accurately
projected changes in temperature and moisture, or other represent earth system processes and improve climate change
climate variables. However, moving from static bioclimate planning. One key factor in these models is the role of
envelope models to a dynamic representation of climate disturbance (Shevliakova et al. 2009). Disturbance can be
change impacts, requires models that include a species poten- defined as either natural hazards or the result of anthropo-
tial range shifts, population and dispersal processes, as well genic influence (e.g., fires, insects, diseases, land-use
as ecological processes that influence habitat suitability, such change). These impacts can have a strong effect on energy
as disturbance (Huntley et al. 2010). Integrated and dynamic fluxes, carbon storage, the nitrogen cycle and the water cycle
modeling has sought to strike a balance between a desire for (e.g., Ellis and Ramankutty 2008). Although difficult to
“realistic process representation” and the heavy data forecast, the representation of agricultural grazing, forest
requirements and computational demands to model such harvest, or other forms of land cover conversions in
complexity. DGVMs is supported by land-use change models which try
up or down the mountain slope over small distances. In these

areas, relatively small climate velocities are required to keep
pace with the rate of temperature change (1e) (Loarie et al.
2009). In contrast, flatter topographies, evidenced in flooded
grasslands and mangroves, will require higher climate
velocities. Here, we see a strong negative correlation between
topographic slope, and velocity from temperature change
(correlation coefficient ¼ 0.92).
Based on these calculations, the authors related the con-
cept of climate velocity to the size of existing global
protected areas. Loarie et al. found that only 8% of these
areas have residence times exceeding 100 years, meaning that
the vast majority of protected areas would experience entirely
novel climates within the century. The size of existing
protected areas did influence overall residence times. For
example, Mediterranean-type temperate broadleaf forest
Fig. 4.16 Annual net emissions of carbon from land-use change, biomes which are small in spatial extent, show small resi-
according to different DGVMs (fine lines) and bookkeeping models dence times despite relatively small (slow) velocities. Con-
(heavy lines). The shaded portion represents one standard deviation of versely, large protected areas with high velocities may still be
all estimates. (Reprinted by permission from Springer Nature,
from Houghton, R.A., 2018. Interactions Between Land-Use Change
able to retain some of the original protected biome within its
and Climate-Carbon Cycle Feedbacks. Current Climate Change Reports borders over this same period due to their spatial extent.
4, 115–127. # 2018 Springer International Publishing AG) Loarie and colleagues suggest that these results have impor-
tant implications: (1) resident species will have to be able to
to quantify both historical patterns of land-use change as well migrate with equivalent speed (km/yr) to keep up with
as project future land-use transitions due to economic and anticipated changes and, (2) existing protected areas may
social factors (e.g., Hurtt et al. 2011; De Rosa et al. 2016). no longer be sufficient in their current extent to protect
Furthermore, land-use models and DGVMs can together assemblages that rely on contemporary climate conditions.
offer a more precise estimate on the emissions contribution Although initially focusing on climate exposure, without
of land-use change (Fig. 4.16). As the suite of global and any comment on specific species tolerance, this concept has
regional DGVMs is developed, the resulting linkages been advanced over the last decade to showcase the
between climate and vegetation change provide a foundation applications of a simple climate metric for conservation.
of support for understanding species response to environmen- Recent studies have also described climate velocity as a
tal change. vector, that not only describes the speed of change but also
its direction. Here, distance is measured by considering a
climate at a single point in time and its future climate analog
4.3.4 Tracking Climate Velocity: Calculating (like drawing an arrow to the climate’s destination point into
the Pace of Climate Change the future) (Brito-Morales et al. 2018). This climate-analog
version of velocity often includes more variables than just
In 2009, Scott Loarie and his colleagues posited that species temperature. These additions lend to greater ecological real-
survival may depend as much on keeping pace with moving ism but can also come with greater uncertainty depending on
climates as the climate’s ultimate persistence. Creating a new scale of future climate projections for these variables.
index, climate velocity, the authors use a simple equation Raquel Garcia and her colleagues evaluated existing
( Cyr-1/ Ckm-1 ¼ kmyr-1) to map the pace of climate change metrics for local and regional climate change and how appro-
across space. Utilizing an ensemble of GCM scenarios which priate uses of each can enhance understanding about demo-
estimate the rate of mean annual temperature increase over graphic and phenological changes, species range
time, and a topographic relief map to show change in eleva- displacements, species range sizes, and novel species
tion, climate velocity can be visualized at fairly high spatial assemblages (2014, Fig. 4.18). Garcia et al. offer four
resolutions (Fig. 4.17). Given the weight of these two factors, categories of climate change metrics which measure the
the magnitude of effect depends both on the emissions sce- magnitude, timing, position and availability of future
nario used and the temporal gradient of change. As shown in climates across local and regional scales. For instance, rela-
the example of Central California in Fig. 4.17, the spatial tive to climate positioning, the metrics of climate velocity,
gradient of temperature change is highest in mountainous and corresponding changes to the distance between analo-
areas (1c) resulting in large changes in temperature moving gous climates in the future, highlight both threats and
Fig. 4.17 (A) (a) Current (1950–2000) mean annual temperature at This would manifest as a high climate velocity directed towards the nearest
800 m resolution globally. The black rectangle indicates the Central occurrence of the original temperature (red arrow). (Part A reprinted by
California (USA) inset in b. (c) The spatial gradient of temperature change permission from Springer Nature, from Loarie, S.R., Duffy, P.B.,
(9-pixel kernel). (d) The temporal gradient of climate change from 2000 to Hamilton, H., Asner, G.P., Field, C.B., Ackerly, D.D., 2009. The velocity
2099 using GCM ensemble projection with an IPCC A1B emission of climate change. Nature 462, 1052–1055. # 2009 Macmillan Publishers
scenario; (e) The velocity of climate change determined from the quotient Limited.; Part B reprinted by permission from Elsevier, from Brito-
of (d) and (c). (B) Slow (low) climate velocities (directed uphill or to the Morales, I., García Molinos, J., Schoeman, D. S., Burrows, M. T.,
closest climate analog areas) due to limited movement required of an Poloczanska, E. S., Brown, C. J., Ferrier, S., Harwood, T. D., Klein,
organism to maintain its thermal environment (blue arrow). Flat C. J., McDonald-Madden, E., Moore, P. J., Pandolfi, J. M., Watson,
landscapes are more homogenous thermal environments, and an organism J. E. M., Wenger, A. S., & Richardson, A. J., 2018. Climate Velocity
experiencing a warming landscape might need to migrate a long distance Can Inform Conservation in a Warming World. Trends in Ecology &
to remain in its original thermal environment (i.e., longdistance dispersal). Evolution, 33(6), 441–457. # 2018 Elsevier Ltd)
Fig. 4.18 Metrics of climate change and associated threats and and species assemblages. (Reprinted by permission from The American
opportunities for species. Metrics are grouped into four dimensions of Association for the Advancement of Science, from Garcia, R.A.,
change, and either quantify changes at a local (locality) or regional (set Cabeza, M., Rahbek, C., Araujo, M.B., 2014. Multiple Dimensions of
of localities) level. Links are established between metrics and potential Climate Change and Their Implications for Biodiversity. Science 344,
threats and opportunities for population dynamics, species occurrence, 486-+. # 2014 AAAS)
opportunities for species movements (2014). Where low (GVMs) in an attempt to predict how vegetation communities
velocities exist relative to a species’ dispersal abilities, cli- worldwide would respond to projected climate changes. They
mate velocity can assist managers in tracking suitable found that, among locally endemic species, extinctions
climates over the region’s topography and identify areas for ranged from <1% to 43% (average 11.6%). The degree of
enhanced habitat connectivity. biome specificity of a species had the greatest effect on
In principle, threats and opportunities for biodiversity extinction rate, with the most specialized species suffering
increase where a species or population is exposed to several the highest losses. Among the world’s designated “hotspots”
dimensions of climate change, often interacting to create a where biodiversity is most concentrated (Chap. 2), the most
different set of ecosystem impacts than might otherwise be vulnerable were the Cape Floristic Region (South Africa), the
expected by looking at any single metric in isolation. Alter- Caribbean, Indo-Burma, the Mediterranean Basin, Southwest
native climate metrics also reveal contrasting spatial patterns, Australia and Tropical Andes. In these areas, projected plant
with potential implications for management. For example, extinctions per hotspot sometimes exceeded 2000 species
the authors note that while mapping climate anomalies (Malcolm et al. 2006).
reveals where populations may potentially be most Working under the assumption that projected habitat
threatened with demographic changes, these maps by them- changes would be realized within 100 years, species
selves do not identify spatial opportunities for organisms to extinctions associated with climate change in hotspots in
track moving climates (Garcia et al. 2014). In sum, while some cases exceeded the number of extinctions caused by
climate exposure is only one component of assessing vulner- deforestation, making climate change an even more powerful
ability, as highlighted at the beginning of this section, teasing threat to biodiversity than habitat destruction. In the model,
apart alternative dimensions of change, or integrating them endemic species with the most restricted ranges were the
within models, can provide valuable information for biodi- most susceptible to the effects of climate change.
versity conservation. Extinction in hotspots was generally no worse than in
other areas, according to the models, but, as Malcolm et al.
noted “Although it is encouraging. . .that these species rich
4.3.5 Decision-Making with Imperfect regions did not appear to be unusually vulnerable to climate
Estimates of the Future change compared with other areas,. . .it suggests that these
high extinction rates can be extended to non-hotspot areas
Model predictions have suggested that many species will lose with similar collections of biome types (mostly tropical and
a substantial proportion of their current distribution, or even subtropical in this case) and where species have similarly
become extinct globally as a result of predicted future climate restricted ranges” (Malcolm et al. 2006).
change (Warren et al. 2013; Thomas et al. 2004). Simulta- One of the reasons for such wide estimates of potential
neously, models have suggested that the diversity of ecologi- extinction rates resulting from climate change is that
cal assemblages may be expected to increase, if the species Malcolm et al. incorporated no species-specific dispersal
that benefit from climate change become much more wide- abilities into the models they constructed. Instead, they sim-
spread and move into new habitats (Thomas 2013, Newbold ply used two contrasting alternatives, “perfect migration,” in
2018). No model can perfectly estimate future conditions, which every species was assumed to be able to track its
even with increasing knowledge about complex interactions bioclimate envelope regardless of the distance or rate of
between components of the climate system. One reason for movement required, and “zero migration,” a scenario which
this is because the drivers of contemporary climate change assumed that species had no ability at all to move to new
are fundamentally social, with human choices influencing locations to adapt to climate change. Neither scenario is
greenhouse gas emissions from all sectors such as land-use realistic for most species, and, when both are used in alterna-
change, transportation and fossil fuel combustion. Despite tive model outputs, estimated extinction rates will show
the uncertainties regarding future human choices, models enormous variation. However, based on paleontological
can help us understand the ecological implications of certain data and theoretical work, these assumptions provide a help-
human choices before we make them. As conservation ful set of bounding conditions that with increasing complex-
biologists utilizing model results to inform decision-making, ity, may be able to improve recommendations for particular
it is important to understand some of the challenges embed- geographic regions or species.
ded in every modeling effort. For example, the implications
of assumptions and the boundaries they can place around
potential futures.
Over 10 years ago, Jay Malcolm and his associates exam-
ined the possible effects of global climate change by
integrating an array of GCMs with global vegetation models
4.4 Conservation Strategies in a Time of Climate Change 159
4.4 Conservation Strategies in a Time other global environmental change. Spatial prioritization of
of Climate Change land-areas for conservation given multiple social goals will
also be necessary, with direct (e.g., temperature change) and
4.4.1 Beyond Traditional Conservation indirect (e.g., shifting agricultural production) changes
Approaches affecting current and future land conservation over time and
space (Jones et al. 2016). Here, we will review several over-
In a 2009 review, biologists Nicole Heller and Erika Zavaleta arching strategies for climate-informed conservation in an
found at least 524 distinct recommendations, from effort to support strategic planning.
113 papers, for conservation actions that could be undertaken
to mitigate the threat of climate change to biodiversity (Heller
and Zavaleta 2009). Although the authors note that these 4.4.2 Fine- and Coarse-Filter Strategies
recommendations were biased towards North America and
Europe, the diverse geographic, biomic and evidence base of Conservation biologist, Morgan Tingley, and his colleagues
these articles signal an overwhelming appraisal of the new understand that climate change presents challenges to
challenge that climate change poses to conservation existing conservation strategies, and differentially affects
(Fig. 4.19). With the availability of spatial tools, modeling the efficacy of past and ongoing conservation planning.
and increasing data collection, it is not a question of whether In an effort to support a cohesive and unified framework
we should engage in climate-informed conservation but of for conservation responses to climate change, the authors
which tools to apply under what conditions to which species classify conservation actions as either “fine-filter” or
conditions (Tingley et al. 2014). The degree to which climate “coarse-filter” and utilize a metaphor of a theater to describe
change adaptation planning is being actively integrated into the interactions and differences between the two strategies
existing management plans or is reflected in conservation (2014). Figure 4.20 provides a summary of this classifica-
policy is more uncertain. tion using the case of Bwindi Impenetrable National Park
In this new era of global change, it is increasingly impor- in Uganda which is situated across the Albertine Rift
tant to consider the range of climate change conservation montane forest in east Africa. The portfolio of strategies
tactics available and recognize that some of the best conser- that can be implemented depend on a manager’s answers
vation options may already be in the conservation biologist’s to three key management questions: where? when? and
“existing toolbox” (Tingley et al. 2014). Improved targeting why?
of resources and planning could make all the difference for First, fine-filter strategies are those that work to “save the
long-term ecosystem resilience under threat from climate and actor.” These strategies seek to understand the impacts of
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Fig. 4.19 The (a) geographic focus, (b) biome focus, and (c) evidence academic or non-governmental institutions. (Reprinted by permis-
basis for recommendations addressing climate change adaptation sion from Elsevier, from Heller, N. E., & Zavaleta, E. S., 2009. Biodi-
strategies for biodiversity management. Recommendations address var- versity management in the face of climate change: A review of 22 years
ious stages in an adaptation process, from research needs to methods for of recommendations. Biological Conservation 142, 14–32.
impact assessments to large-scale changes in policies by governmental, # 2008 Elsevier Ltd)
Fig. 4.20 (a) Conservation units. Fine-filter and coarse-filter actions Conservation strategies for climate change. Depending on the approach,
conserve the “actors” and the “stage” of the ecological theater. To there is a portfolio of conservation strategies that can be implemented in
illustrate, large protected areas, such as Bwindi Impenetrable National the face of climate change. (Reprinted by permission from Wiley, from
Park in Uganda, can help to protect unique communities such as the Tingley, M.W., Darling, E.S., Wilcove, D.S., 2014. Fine- and coarse-
Albertine Rift montane forest of Africa, yet individual species with filter conservation strategies in a time of climate change: Conservation
restricted ranges and species-specific responses to climate change, with climate change. Annals of the New York Academy of Sciences 1322,
such as mountain gorillas, may fall through the wide mesh of a coarse- 92–109. # 2014 New York Academy of Sciences)
filter approach and require single-species conservation management. (b)
climate change on a single species and mitigate its impact. In further note that “protected areas previously established for
a general sense, actions at this scale are either in situ (protect single-species management may no longer be able to fulfil
the species within its current range) and ex situ (include their fine-filter purpose but have the potential to be managed
conservation within a zoo or botanical garden). Although successfully as movement facilitators for a larger suite of
these actions are part of the bedrock of traditional conserva- species” (Tingley et al. 2014).
tion, they are made more challenging as current ranges shift, Another potential fine-filter strategy is to identify climate
and the static protected areas enveloping or intersecting core refugia at very fine spatial scales. Species that are highly
habitats fail to permanently protect these species as they sensitive to a changing climate would benefit from spatial
move. One planning response to this challenge is to increase areas that are likely to change more slowly relative to
connectivity between existing patches of suitable habitat, and surrounding areas. Refugia could be as expansive as a moun-
even work to restore habitats along pathways of future cli- tain range and as small as a microclimate, created by slope,
mate change trajectories, For example, a recent study by aspect, and surrounding topography (Tingley et al. 2014).
Patrick Jantz and colleagues shows that, using a high- Although this strategy is not likely to be permanent given
resolution data set of vegetation carbon stocks, there are ongoing climate change, it could help buy time for species
currently more than 16,000 unprotected habitat corridors in behavioral adaptation or for conservation managers looking
the tropics that, if protected, could connect 5,600 protected to protect more suitable land area in the future. As an exam-
areas in the region with co-benefits for climate mitigation and ple, California has adopted new Thermal Refugia Protection
species movement (Jantz et al. 2014). Tingley and colleagues Policy to identify and protect cold-water refugia for salmon
within the Klamath watershed (Stein et al. 2013).Where there (Hunter et al. 1988). Tingley et al. note that by focusing on
is no refugia, increasing habitat connectivity may be the only variables such as soils, geology, topography and elevation,
remedy for helping species move from one point to another coarse-filter approaches are “technically considered
on their own, if they can keep pace with and track their scenopoetic, sensu Hutchinson, from the Greek for ‘setting
climate niche. the scene.’ Not surprisingly then, coarse-filter conservation is
In the most extreme cases, translocation, otherwise often described metaphorically as ‘conserving the stage’ of
referred to as assisted migration, might be necessary to Nature’s theater” (2014).
help imperiled species establish new populations in areas As fine-filter conservation can be broadly classified as
with more suitable climate (Thomas 2011; Tingley et al. either in-situ or ex-situ, coarse-filter approaches involve
2014). Over the past decade, there has been considerable either conserving land facets or conserving areas with slower
discussion within the scientific community about the ethics, climate change velocities. Land-facets are broadly described
feasibility and risks of assisted migration. Hoegh-Guldberg as geographic areas across a landscape that have similar
and colleagues argue that translocation should become topographic and soil characteristics and can act as surrogates
an important part of integrated conservation strategies for ecological and evolutionary processes that support
as long as each case is undertaken with independent consid- gamma (or landscape) diversity (Tingley et al. 2014). Current
eration of extinction risk, technical capabilities, and the global conservation priorities are not often based on land
biological and socioeconomic costs and benefits (2008). For facets, and if considered in spatial planning, land facets
example, it is likely to be unacceptable to move a toxic plant may provide more connectivity for a suite of focal species
or large carnivore into regions currently prioritized for than a corridor plan based on the individual species (Brost
grazing livestock even if it is technologically feasible and Beier 2012; Tingley et al. 2014). The theoretical and
(Hoegh-Guldberg et al. 2008). One expected benefit from paleoecological foundations for this approach also suggest
translocation may be increased gene flow, with increased that using land facets as the primary point of consideration
genetic diversity aiding otherwise isolated populations with may help protect both current and future biodiversity, includ-
adaptation to new climates (Aitken and Whitlock 2013). ing species that are still unknown to scientists (Hunter 1990).
There are clearly many negative examples of transloca- A second coarse-filter approach is the protection of suit-
tion, where introduced species create unintended able climate space. As discussed earlier, climate velocity and
consequences. However, with intentional decision-making other climate metrics have been used extensively to estimate
frameworks, low-risk climate-informed translocation may the distance a species would have to travel to keep pace with
be the best changes a species has for survival if their current their shifting climate niche. One approach for maximizing
range is fragmented and they are otherwise unable to shift long-term persistence is to identify and protect areas with the
with climate change on their own. As was discussed in this smallest expected climate velocities and use climate trajec-
chapter’s section on modeling, uncertainty regarding tory to improve future reserve planning given expected direc-
specific-specific (fine-filter) strategies can be high. However, tional movements (Loarie et al. 2009, Dobrowski et al. 2013).
a range of conservation scenarios with clear estimates of One effort called “Spatial Planning for Protected Areas in
uncertainty and the propagation of error through the model, Response to Climate Change (SPARC)”, led by bioclima-
can enable decision-making and support conservation action tologist Lee Hannah and colleagues at Conservation Interna-
(Anderson 2013). tional, utilizes a big data approach to identify new regions
Parallel strategies designed to increase the resiliency of and ecosystems that, if protected, will preserve biodiversity
protected area networks, even with potential risks to individ- and critical ecosystems over the coming decades (SPARC
ual species, are what Tingley et al. refer to as coarse-filter 2019). Although topographic heterogeneity is an identifying
approaches, or “conserving the stage.” Although, the original characteristic of areas with slow climate velocities and
goal of this ecosystem-based approach was to conserve targeted land-facets, Tingley et al. note that diverse
assemblages of species rather than any particular species, topographies are already overrepresented in protected area
the authors stress that “coarse-filter conservation should not networks and present certain “conundrums” for future terres-
be based on contemporary communities – or any biological trial conservation, and so ask, “Do we invest in climatically
assemblage – but rather on conserving locations with unique resilient sites, even though they are already overrepresented
environmental characteristics unchanged by species’ pres- in protected area networks, or do we invest in underprotected
ence, such as calcareous shaded substrates or rocky reefs” landscapes, notably lowland areas with high soil productiv-
(Tingley et al. 2014). This change in focus follows important ity, with the risk that such sites may offer fewer refugia to
work by Malcom Hunter and colleagues, which emphasizes species in the face of climate change?” (Tingley et al.
that biological communities are “ephemeral aggregations of 2014:101).
species rather than intact units moving together through time” The framework we employed for global prioritization
and essentially “temporary occupants of those arenas” strategies for land use (Chap. 2, Sect. 2.10.1, Fig. 2.24)
(Kilpatrick 1983; Brooks et al. 2006) can also be applied to

climate change strategies in conservation. Takuya Iwamura
and his colleagues note that “while global conservation
priorities affect the spending of an approximately US$ 1.5
billion from international organizations (Halpern et al. 2006),
none of the schemes explicitly consider the impacts of cli-
mate change. . . . and that there are substantial differences
among existing global conservation priorities in terms of their
robustness with respect to climate change” (Iwamura et al.
2013). As a function of cost, some strategies, and the land-
prioritization processes that accompany them (Fig. 4.21), are
more robust to future climate change, and may show a greater
return on investment (upper right), while other may require
enhanced funding for climate adaptation activities to navigate
a larger degree of climate instability and change (lower left).
Fig. 4.21 Categorical mapping of global conservation priority schemes
Sometimes fine- and coarse-filter approaches can result in
based on endemism, irreplaceability, and vulnerability. The figure
expresses the three-dimensional plot for: X-axis: irreplaceability; similar solutions. Figure 4.22 compares two approaches to
Y-axis: endemism, and Z-axis (depth): vulnerability. Each of the conservation management in the context of climate change,
schemes are as follows: Biodiversity Hotspots (BH); Endemic Bird one fine-filter approach focusing on the vulnerabilities of
Area (EBA); Global 200 (G200); High Biodiversity Wilderness Areas
Indo-Pacific reef coral and fish species given a parallel con-
(HBW); Crisis Ecoregions (CE); Megadiverse Countries (MC); Frontier
Forests (FF); Last of the Wild (LW). (Reprinted by permission from cern of fisheries exploitation (A), and a second coarse-filter
Elsevier, from Iwamura, T., Guisan, A., Wilson, K. A., & Possingham, approach considering the relative climate vulnerability of
H. P., 2013. How robust are global conservation priorities to climate different terrestrial ecoregions given pre-existing levels of
change? Global Environmental Change, 23(5), 1277–1284.
vegetation intactness (B). Under either approach, each of
# 2013 Elsevier Ltd)
the four quadrants highlight a different management strategy
Fig. 4.22 Fine-filter (a) and coarse-filter (b) ecological units are that require different types of conservation actions, as outlined in
distributed across four scenarios of climate and local stressor impacts. Numerals I–IV indicate four distinct quadrants that require different
(a) Species-specific sensitivities of 163 Indo-Pacific reef coral and fish types of conservation actions, as outlined in Table 4.5. (Reprinted by
species to climate impacts and fisheries exploitation based on trait and permission from Wiley, from Tingley, M.W., Darling, E.S., Wilcove, D.
population responses to these two stressors. (b) The distribution of S., 2014. Fine- and coarse-filter conservation strategies in a time of
803 global terrestrial ecoregions across gradients of climate vulnerabil- climate change: Conservation with climate change. Annals of the New
ity (reverse of climate stability in Watson et al. 2013) and natural York Academy of Sciences 1322, 92–109. # 2014 New York Academy
vegetation intactness. Numerals I–IV indicate four distinct quadrants of Sciences)
Table 4.5 Fine- and coarse-filter approaches propose similar conservation actions for climate change adaptation to four distinct threat scenarios
Priority conservation actions that
Future ecological trends under threat can be applied to fine- and coarse-
Threat scenarioa Current ecological community regime scale managementb
I. Low climate vulnerability, Intact habitat, high diversity of Low species turnover, sensitive Identify and manage local
high risk to local stressors sensitive and tolerant species species remain in community stressors (land-use, exploitation,
(e.g., fisheries exploitation, invasive species)
habitat loss) Best chance for functioning Priority investments for protected
ecosystem processes areas
Areas may act as climate refugia;
expect incoming climate migrants
Monitoring and adaptive
management critical to maintain
intact ecosystems
II. High climate vulnerability, Intact habitat, high diversity of Loss of sensitive species, stress- Identify and protect climate
high risk to local stressors sensitive and tolerant species tolerant and weedy life histories refugia
(e.g., fisheries exploitation, favored
habitat loss) High turnover as species track climate Manage local stressors (land use,
niches exploitation) within refugia
Promote connectivity as species
shift with climate niches, and
retreat into refugia
Climate-sensitive species may
require translocation
III. Low climate vulnerability, Degraded habitat; low species Relatively little change Habitat restoration a good
low risk of local stressors diversity; community investment for a stable climate
composed of generalist and Small turnover of species within Experiment with creative
opportunistic species habitat approaches to habitat
reengineering: goal is species
recovery within stable climates
Some recovery of more sensitive Identify and manage local
species stressors to promote recovery of
sensitive species
Species are not likely to require
immediate translocation
IV. High climate Degraded habitat; low species Short-term change will depend on Increase connectivity, dispersal
vulnerability, low risk of local diversity; community co tolerance of species and habitats to potential
stressors composed of generalist and climate impacts
opportunistic species Over the long term, climate impacts Identify and protect refugia
may exceed tolerance of generalist,
opportunistic species, resulting in
high turnover
Climate-sensitive species may
require translocation
Reprinted by permission from Wiley, from Tingley, M.W., Darling, E.S., Wilcove, D.S., 2014. Fine- and coarse-filter conservation strategies in a
time of climate change: Conservation with climate change. Annals of the New York Academy of Sciences 1322, 92–109. # 2014 New York
Academy of Sciences
a
There are four common “threat scenarios” under the combined impacts of climate and local stressors (see Figure 4.22). Each of these scenarios is
associated with a current ecological community that can be described by diversity, species turnover, and the composition of sensitive and tolerance
species. With the continued pressure of these threats and predicted future ecological trends, a combination of fine-scale and coarse-scale priority
conservation actions can be identified for each scenario
b
Conservation actions include both traditional approaches and novel climate-relevant approaches. Critically, while actions have philosophical ties to
either fine- or coarse-filter approaches, the actions themselves can be equally applied to conservation at either the species- or landscape-level scale
given the degree of climate or other stressors. For a species maximize climate refugia. This approach could parallel one
with high vulnerability to climate and high risk of fisheries for an ecoregion that has both high land-use change and
exploitation, new reserves may need to be created which climatic instability and require either ecosystem restoration
or the translocation of sensitive species (Tingley et al. 2014, these efforts over time to achieve a balance between anthro-
Table 4.5). pogenic emissions by sources and removals by sinks in the
Many climate-informed strategies have not distinguished second half of this century (UNFCCC 2019).
between scale of management and have been unclear as to Complementing this agreement are a range of country-level
whether the strategy targeted species, assemblages, or and regional climate initiatives which work to actualize emis-
surrounding physical environments. In practice, the best sion reductions.
management strategies will be those most sensitive to place Starting in 2023, and then every 5 years after, the Parties
(particular patch, park, region, or country). And, as has will come together to evaluate their collective progress
already been discussed, climate change does not act in isola- against the goals of the Agreement, sometimes called a
tion from other anthropogenic (human-caused) stressors and “global stocktake.” Some of the central goals of the agree-
must be taken into account when investing resources. For ment include: pursuing efforts to further limit the global
example, in a vulnerability assessment of 300 plant and temperature increase to 1.5 C, reaching the peak of global
animal species in Florida, certain species of highest vulnera- greenhouse gas emissions as soon as possible, conserving
bility were remarkably consistent across a range of stressors and protecting carbon sinks (like forests), establishing market
such as climate change, land-use change, and sea-level rise and non-market approaches to support voluntary cooperation,
(Reece et al. 2013). Although climate change should be and investing in adaptation that will improve global resilience
considered within future management actions, it should not and support those already feeling the effects of climate
replace actions targeted at other stressors. change (Hoegh-Guldberg et al. 2018). Although one of the
Even though the “stage is on fire,” the “show must go on” strengths of the Paris Agreement is the enhanced flexibility
(Tingley et al. 2014). Utilizing a diversified portfolio strat- individual countries have to set their own emissions goals,
egy, conservation biologists should include climate change this flexibility has also caused some to question whether
considerations into their management planning and recognize current ambition levels are really enough to keep stated
that the past investments they have made in protected areas temperature goals. For example, in a recent Nature publica-
and species conservation plans may not hold up “as is” in the tion, Rogelj et al. argue that two thirds of the global GHG
face of ongoing global climate change. As information budget for keeping warming to below 2 C has already been
improves, and uncertainties and risks are clarified, managers emitted, and that the window for limiting warming to below
will have to adapt their own behavior to protect our diversity 1.5 C already seems to have closed (2016). Although sum-
of life. ming emissions reduction pledges across all Parties may
seem like simple arithmetic, global decision-makers must
reckon with a range of potential reduction scenarios for
4.5 Policy Initiatives for Climate Change each participating Party (or country) (Fig. 4.23) that are
and Conservation often dependent on the availability of financial or technolog-
ical support, as well as a common measurement, verification
Conservation responses to climate change require two and reporting system (Rogelj et al. 2016).
components. The first is to adapt conservation strategies to
manage dynamic biodiversity and mitigate the overall effects
of climate change. The second is to engage policy makers to
What has your county pledged to do to keep warming
take action to reduce greenhouse gas emissions to levels that
at or below 1.5 C? How might your understanding of
will keep biological changes manageable.
climate change impacts on conservation efforts better
The second strategy is manifested in the Paris Agreement
inform your personal and professional advocacy for
(2015), the primary global instrument to attempt to regulate
climate change solutions?
carbon emissions, now signed by 195 countries and ratified
by 185 worldwide. The Agreement aims to limit carbon
emissions sufficiently to keep global warming below 2 C.
The Paris Agreement builds on the United Nations Frame- Shared indicators of success will be required to track
work Convention on Climate Change (UNFCCC) and works global progress through time. Some of these include the
to overcomes some of the challenges presented in previous fuel switch from coal to gas, and the relative share of fossil
agreements such as the Kyoto Protocol. All Parties to the fuels in energy production relative to renewables. Although
Paris Agreement must submit their own individualized plan several measures of progress are still broadly consistent with
for emissions reductions, called Nationally Determined emission scenarios to keep temperatures below 2 C, the
Contributions (NDCs), putting forward their “best effort” to continued lack of large-scale carbon capture and storage
reduce emissions economy-wide, gradually strengthening threaten agreement targets through 2030 (Peters et al.
4.5 Policy Initiatives for Climate Change and Conservation 165
Fig. 4.23 Global greenhouse gas emissions as implied by Intended given. Symbols represent single studies and are offset slightly to
Nationally Determined Contributions (INDCs) compared to no-policy increase readability. Dashed brown lines connect data points for each
baseline, current-policy and 2 C scenarios. White lines show the study. (Reprinted by permission from Springer Nature, from Rogelj, J.,
median of each range. The white dashed line shows the median estimate den Elzen, M., Höhne, N., Fransen, T., Fekete, H., Winkler, H.,
of what the INDCs would deliver if all conditions are met. The 20th– Schaeffer, R., Sha, F., Riahi, K., & Meinshausen, M., 2016. Paris
80th-percentile ranges are shown for the no-policy baseline and 2 C Agreement climate proposals need a boost to keep warming well
scenarios. For current-policy and INDC scenarios, the minimum–maxi- below 2 C. Nature, 534(7609), 631–639. # 2016 Macmillan
mum and 10th–90th-percentile range across all assessed studies are Publishers Limited)
2017). In particular, ecosystem restoration and the conserva- says that greenhouse gas concentrations should be stabilized
tion of existing forests represent some of our best options for at a level at which ecosystems will be able to “adapt natu-
increased carbon sequestration. Although sometimes seen as rally” within a predictable time frame. However, as global
a secondary mitigation option within NDCs, forest conserva- temperatures increase, the designs of nature are not predict-
tion and restoration through intentional land-use can turn a able; once disaggregated, they do not and will not reassemble
net anthropogenic source during 1990–2010 into a net sink of in previous forms” (Lovejoy and Hannah 2018). Citing a
carbon by 2030 (Grassi et al. 2017). Of particular importance study by Erb et al. 2018, Lovejoy and Hannah argue given
here are improved carbon monitoring technologies which can the historical contribution of ecosystem destruction and deg-
support consistent global planning, such as those being radation to atmospheric CO2 (450–500 Gton), intentional and
offered through the recent NASA Global Ecosystem Dynam- purposeful ecosystem restoration could help us keep global
ics Investigation (GEDI) mission, providing a 3-D view of temperature rises at 1.5 C, with a reduction of atmospheric
the world’s forests with the aid of LiDAR (NASA and UMD loading by 1 ppm of CO2 for every 7.7 Gton of ecosystem
2019), and NASA Carbon Monitoring System products services restored (Lovejoy and Hannah 2018).
which can now report forest carbon sequestration potentials Given ongoing opportunities for synchronized climate
at very high spatial resolutions (90 m) (Hurtt et al. 2019). change mitigation and adaptation, conservation biologists
In a recent editorial, Thomas Lovejoy and Lee Hannah are poised to advance solutions that both slow climate change
note that many biological systems around the planet will not and protect biodiversity. Many international initiatives such
be able to tolerate temperature rise much beyond 1.5 C, as the recently launched Intergovernmental Science-Policy
further emphasizing the need for ecosystem restoration. Platform on Biodiversity and Ecosystem Services (IPBES),
They argue, “The essential unpredictability of nature in its modelled after the IPCC, provide a platform for scientific
response to climate change runs counter to the language of engagement on a global stage. The benefits that biodiversity
the UN Framework Convention on Climate Change, which and nature provide to human development also underpin the
success of UN Sustainable Development Goals and must be

considered when envisioning the future of the planet through Some species cannot move. Neither the top of the
policies and planning. Every country must be encouraged to world nor the top of a mountain provides room for
participate in this process, and offer their best science, poleward migration or upward mobility, except into
technologies and policies. When individual countries, partic- the mists of extinction, a place for the ghosts of biodi-
ularly large GHG emitters, remove themselves from solution- versity past who can no longer dwell on solid ground.
building conversations on the global stage, they put both For these species, their conservation ex situ, in the zoos
people and ecosystems at risk. and conservatories of the world, will take on new
significance, as humans strive to perfect the skills to
keep their kind alive, until we have made the world
Synthesis
good enough for them to live in again. But in a world of
The long-predicted effects of climate change have,
climate change, these are the elements that must form
until recently, been thought to be in the distant future.
our climate of hope for the work of conservation biol-
Significant future events of climate change are
ogy. And, there is much we can yet do.
undoubtedly ahead for planet Earth, but climate change
is no longer a future scenario. It is here, it is because of
us and it is serious. If the best laid local, regional, and
global conservation plans can be swamped by unman-
ageable planetary forces of temperature, precipitation,
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Conservation Genetics
5
In the past two decades, a new field of conservation genetics has emerged with two general goals: (a) the precise
description of genetic changes affecting population survival that occur during range and population contrac-
tion; and (b) of genetic insight to successful management of threatened populations.
Stephen J. O’Brien (1996)
Keywords (McNeely et al. 1990). Genetic diversity is essential for

Genetic diversity · Genetic drift · Population viability adaptation to environmental change, and such adaptation is
analysis · Inbreeding · Genetic technology · Phylogenetic key to long-term survival in any species (Schemske et al.
analysis · Landscape genetics · Genetic management · 1994). Taxonomy is critical in conservation for defining
Captive breeding management · Genetic applications components of biodiversity, and genetics and genetic tech-
nology have become increasingly dominant means of
identifying taxa (Haig et al. 2011). Finally, there is a direct
correlation between population genetic diversity and popu-
Overview lation fitness, a relative or absolute measure of reproductive
In this chapter you will learn about: efficiency or success (Reed and Frankham 2003).
Conservation biologists have two primary goals in their
1. How genetic processes and events can lead to popu- study and application of genetics. One is to preserve heritable
lation decline. genetic variation, particularly in small populations threatened
2. How genetic diversity is measured. with extinction. Genetic variation is related to the allelic
3. How inbreeding can lead to population decline and diversity in a population. Alleles are different forms of the
extinction. same gene, and the number of alleles present for a given gene
4. How landscape characteristics affect genetic is one measure of the genetic variation of a population. A
diversity. second goal is to prevent fixation of alleles, which can con-
5. Practical means of managing genetic diversity to tribute to reduced fitness and accumulation of harmful
achieve conservation goals. mutations (Lynch 1996). Preserving high levels of heritable
6. Applications of genetic theory and technology to variation helps to retain a population’s evolutionary
solve conservation problems. potential – its capacity to adapt to environmental change
over the long term. Preventing fixation of deleterious alleles
can prevent declines in survivorship and fecundity that often
occur in small populations as a result of the expression of
traits associated with such alleles.
In the 1960s and early 1970s, even before conservation
5.1 Conservation Genetics biology emerged as a distinct discipline, genetic concerns
and Conservation Biology about small populations were growing. Moore (1962) and
Hooper (1971) considered problems associated with
In 1990, IUCN identified the conservation of genetic diver- inbreeding depression that could arise in populations con-
sity, along with species and ecosystem diversity, as one of the fined to refuges. At about the same time, Michael Soulé, in
three foundational elements of biodiversity conservation

172 5 Conservation Genetics
his classic review of potential genetic liabilities for small 5.2 Bottlenecks, Inbreeding, and Population
populations, “The Epistasis Cycle: A Theory of Marginal Decline
Populations” (Soulé 1973), demonstrated that, in populations
of fruit flies (Drosophila), marginal populations rarely pos- 5.2.1 The Theoretical Foundation
sessed unique gene arrangements (haplotypes), and many had
reduced allelic diversity. “Marginal populations,” wrote The problems Soulé described are exacerbated in a demo-
Soulé, “are. . .prone to severe reduction in numbers and can graphic event known as a bottleneck (Frankel and Soulé
experience intermittent drift. Just a trickle of gene flow can, 1981, Fig. 5.1) in which a population declines to very few
however, restore lost alleles; but, if the organism has poor individuals, inevitably accompanied by a loss of genetic
dispersal powers, then marginal, isolated demes are expected diversity. After a bottleneck, the remaining individuals repre-
to be allelically depauperate” (Soulé 1973). The problem sent only a sample of the original population. The smaller the
Soulé was referring to was genetic drift, the random sample, the more likely it is to be unrepresentative of its
fluctuations in allele frequencies that occur as a result of source, and the more certain that some alleles, especially
non-representative combinations of gametes created during rarer ones, will be lost. Such loss of genetic variation may
breeding. Genetic drift refers to the effect of chance genetic potentially permit a greater proportion of recessive alleles to
events in populations relative to selective forces, making occur in a homozygous condition, so that previously masked
small populations more vulnerable to outcomes of individual phenotypes are expressed. Many of these recessive alleles
matings instead of overall probabilities of inheritance. have deleterious or even lethal effects on organisms. The
Since the 1980s, many studies followed showing the rela- longer the bottleneck persists, the more alleles are likely to
tionship of genetic diversity to population size and fitness, be lost (Frankel and Soulé 1981, Table 5.1).
but proponents of the so-called “No Genetic Impact Hypoth- The reduced numbers of individuals associated with a
esis” (NGIH) (Lande 1988) claimed that most endangered bottleneck creates conditions that can provide increased
and recently extinct populations were reduced or opportunity for inbreeding, the mating of genetically-related
exterminated by demographic processes long before loss of individuals. Inbreeding in a population can result in
genetic diversity had negative effects. Such voices asserted inbreeding depression, a pattern of reduced reproduction
that genetic concerns were less important than historically and survival frequently observed in small, inbred populations
believed because “. . .loss of diversity is more likely to be a (Frankham et al. 2002:24). Based on theoretical
symptom of endangerment than its cause. . .[and] those considerations alone, Frankel and Soulé derived “the basic
alleles most likely to be lost as a result of genetic drift – rule of conservation genetics,” expressed as a percent change
rare alleles – are also the least likely to contribute to any in the inbreeding coefficient (the probability that two alleles
immediate response to natural selection” (Holsinger et al. at the same locus in an individual are identical by descent).
1999). The “rule” is that selection for performance and fertility can
The NGIH has not been supported in detailed genetic balance inbreeding depression if change in the inbreeding
studies of endangered populations. In an extensive meta- coefficient (ΔF) is no more than 1% per generation
analysis of published literature completed in 1996, conserva- (Fig. 5.2). The significance of the rule is provided by the
tion geneticist Richard Frankham (1996) demonstrated that authors themselves: “We refer to the 1% rule as the basic rule
genetic variation declined with declining population size, and of conservation genetics because it serves as the basis for
was greater in species with larger geographic ranges and
in mainland populations than in island populations of the
same or similar species (Frankham 1996). Subsequent
studies have shown that genetic factors did contribute to
endangerment and extinction, and that genetic diversity is
correlated with a population’s adaptive capacity Fitness is
consistently higher in populations with greater genetic diver-
sity (heterozygosity) and in larger populations compared to
smaller ones (Reed and Frankham 2003). Conversely, lower
genetic diversity increases risk of extinction, with threatened
species evidencing lower genetic diversity than
non-threatened species in the same taxonomic groups
(Spielman et al. 2004). Small populations suffer loss of
genetic diversity through multiple mechanisms, but two that
deserve our attention first are population bottlenecks and Fig. 5.1 Graphical representation of population size before (a), during (b),
inbreeding. and after (c) a population bottleneck. (Design by M. J. Bigelow)
5.2 Bottlenecks, Inbreeding, and Population Decline 173
Table 5.1 Percent change in genetic variation and proportion of rare alleles lost from a population at bottlenecks (minimum sizes) of different
magnitudes
Percent change in genetic Proportion of rare alleles lost
Number of individuals in sample (N ) variation p1 ¼ 0.70, p2 ¼ p3 ¼ p4 ¼ 0.10 p1 ¼ 0.94, p2 ¼ p3 ¼ p4 ¼ 0.02
1 50.0 0.6300 0.7200
2 25.0 0.4950 0.6925
6 8.3 0.2125 0.5900
10 5.0 0.0925 0.5000
50 1.0 0.0025 0.1000
1 0.0 0.0000 0.0000
Developed based on equations from Frankel and Soulé 1981. Table design by M. J. Bigelow
Values of p represent proportions of each of 4 alleles
United States, most (98.5%) have been, at some point, cross

bred with domestic cattle and show evidence of cattle alleles
in their chromosomes. The YNP bison do not. Blessed with
good representation of ancestral genetic diversity, this popu-
lation has increased from 46 individuals in 1900 to over 3500
today without showing signs of abnormal physical
characteristics or reduced juvenile or adult survivorship
(Herman et al. 2014).
Another US population, the Texas State Bison Herd
(TSBH), provides a dramatic contrast to the YNP bison in
illustrating negative effects of inbreeding. Established in
1897 with 36 individuals, the TSBH, despite complete pro-
tection, has suffered multiple population bottlenecks over the
Fig. 5.2 Percent change in the inbreeding coefficient (ΔF) at different
last 125 years, as well as chronically small population size,
population sizes. Note that the value of the inbreeding coefficient low levels of genetic diversity, low recruitment, and high calf
increases as population size declines. (Design by M. J. Bigelow) mortality rates. Population viability analysis based on cur-
rent population demography posited a 99% probability of
calculating the irreducible minimum population size consis- extinction of the TSBH in less than 40 years. Why are there
tent with the short-term preservation of fitness” (Frankel and such pronounced contrasts between two populations formed
Soulé 1981:73, emphasis mine). Such short-term fitness pres- at about the same time with similar numbers of individuals?
ervation was considered safely achieved in most populations The TSBH was established by a rancher, Charles
with an effective size of 50 individuals (Franklin 1980; Goodnight, who bred the bison to Angus domestic cattle
Frankel and Soulé 1981). (Bos taurus) in an effort to produce a more robust and hardier
breed of cattle. Effects of those early cross-breeding efforts
are still present in the Texas herd. Six of the original
5.2.2 Of Bottlenecks and Bison 36 members of the TSBH contained domestic cattle-type
DNA. From 1998 to 2004, the Texas herd only increased
Whether or not past population reductions actually created a from 36 to 40 individuals. Natalie Halbert, a veterinary
genetic bottleneck, and associated inbreeding depression, is biologist, and her colleagues examined the fitness of the
influenced by the genetic diversity of individuals who survive Texas bison by testing all twelve adult males for fertility.
or “pass through” the bottleneck. In North America, bison Of these, noted Halbert, only four “exhibited normal sperm
(Bison bison) historically had a continental population of motility and morphology, but the remainder had
somewhere between 30 and 60 million individuals. abnormalities outside acceptable baseline ranges including
Decimated by hunting, only two wild herds survived to the low motility, bent tails, and detached heads” (Halbert et al.
mid 1880’s, one in Canada in what is now Wood Buffalo 2004). Halbert et al. (2004) also examined the 17 adult
National Park, and the other in the United States in females in the population, finding that, in 2001, “15 bison
Yellowstone National Park (YNP). The YNP bison are the (~83%) were pregnant. From these apparent pregnancies, five
only US population descended from a continuously free- calves were born and only one survived to 2003. . .,
ranging wild herd and have maintained a high level of genetic confirming the trend of poor recruitment in this herd. As
diversity. Of the estimated half million bison now in the such, it is probable that male infertility and the inability of
Fig. 5.3 Distribution of alleles from three bison (Bison bison) Oxford University Press, from Halbert, N.D., Raudsepp, T.,
populations examined at 54 microsatellite loci. Note that the Texas Chowdhary, B.P., Derr, J.N., 2004. Conservation Genetic Analysis Of
State Bison Herd, the most inbred population, has no genetic locus The Texas State Bison Herd. Journal of Mammalogy 85, 924–931. #
with more than four alleles, compared to as many as 7–10 in the 2004 American Society of Mammalogists)
comparatively less inbred populations. (Reprinted by permission from
females to carry pregnancies to term are negatively affecting have quantitative means of measuring it if one is to have any
the recruitment and population growth rates in the herd over hope of managing it. Some measures of genetic diversity are
the past six years” (Halbert et al. 2004). Halbert and her identical to measures of community diversity described in
co-workers also found that, compared to the YNP population Chap. 2. For example, the Shannon Index (a measure of
and the bison population in another national park (Theodore species diversity in a community) can be used just as effec-
Roosevelt National Park in North Dakota, USA), the Texas tively as a measure of genetic diversity, if proportional abun-
bison had lower genetic diversity, as measured by the number dance of alleles is substituted for proportional abundance of
of alleles at different genetic locations (loci) (Fig. 5.3). Inter- species. Three commonly used measures of genetic diversity
species hybrids often experience poor fertility due to sexual are polymorphism, average heterozygosity, and allelic diver-
incompatibility. Such low fertility can lead to reduced popu- sity (Appendix 5.1).
lation size which can precipitate a population bottleneck and Polymorphism refers to a genetic locus that has two or
resultant loss of genetic diversity. In the TSBH herd, more forms (alleles). In a population or population subunit,
hybridization and consequent loss of fertility set the popula- polymorphism is expressed as the probability (P) of encoun-
tion on a trajectory toward increasingly low genetic diversity, tering a polymorphic locus among all loci in the population.
all of which combined to reduce population growth. More generally, P (polymorphism), for an individual or
any larger group or unit, can be determined from the
expression
5.3 Measuring Genetic Diversity
in Populations P ¼ number of polymorphic loci=total number of loci
5.3.1 Foundational Measures of Genetic Although polymorphism is an easy concept to understand,

Diversity it is not the most frequently used measure of population
genetic diversity. That measure is average heterozygosity
As the saying goes in conservation (as well as in politics), (H) which refers to the average proportion of individuals in
“What gets measured, gets managed.” Because genetic diver- a population that are heterozygous (carrying two different
sity is so important to population conservation, it is vital to alleles) for a particular trait. This metric reflects the proportion
5.3 Measuring Genetic Diversity in Populations 175
of heterozygous individuals in a population measured across neither the average nor the range of genetic characteristics
several loci. We can calculate average heterozygosity as in the population. Genetic drift, which can be indexed by the
change in frequency of a randomly selected allele, q, in one
H ¼ ΣH i =N generation, can be estimated as
Where H is average heterozygosity at locus i and N is the total qð 1 qÞ

Δq ¼ ,
number of loci used in the estimate. Suppose there are four 2N e
loci in a population. We will call them (for lack of imagina-
tion) 1, 2, 3, and 4. Suppose the frequency of heterozygotes where Ne is the variance effective population size, a concept
for locus 1 is 0 (all individuals are homozygous for this we will examine shortly. If the frequency of q is 0.2 and the
locus), 0.3 for 2, 0.5 for 3, and 1.0 for 4. Then effective population size is 100, expected change in q (i.e.,
Δq) is 0.2(1–0.2)/2 (100) ¼ 0.16/200 ¼ 0.0008, or eight
H ¼ ð0 þ 0:3 þ 0:5 þ 1Þ=4 ¼ 1:8=4 ¼ 0:45: one-hundredths of 1%. In contrast, if effective population
size is 10, the effect on the same allele would be equal to
The third measure, allelic diversity (A) refers to the aver- 0.16/20 ¼ 0.008 (eight tenths of 1%). Both results represent
age number of alleles per locus. It can be calculated at very small effects but notice the order of magnitude increase
in the smaller population. If the population remains small and
A ¼ Σ ½A1 þ A2 . . . þ An =N, this relationship is reiterated for many generations, the effect
of genetic drift on gene frequencies can become large.
where A1 is the number of alleles at locus 1, A2 the number of Genetic drift can produce loss of heterozygosity or fixa-
alleles at locus 2, and so on through all N loci. We can put tion of deleterious alleles. As a result, there is a decrease in
concepts of heterozygosity (H ) and allelic diversity (A) to heterozygosity and in the number of different alleles at a
immediate use in understanding two processes affecting single locus in the population (Caughley 1994). The degree
genetic diversity in small populations: bottlenecks and of decline in heterozygosity is a function of population size,
genetic drift. N, over the number of generations, t (Wright 1931).
Generalizing the equation for any number of t generations,
heterozygosity declines as
5.3.2 Loss of Genetic Diversity over Time:
Bottlenecks and Genetic Drift H t ¼ H 0 ½1 1=ð2N Þt
After a population bottleneck, low genetic diversity can per- where H0 represents the original level of heterozygosity
sist for generations even after the population recovers to a (usually expressed as a proportion) and Ht represents the
larger size. Thus, current population sizes do not always new level of heterozygosity after t generations. Over one
correlate positively with the genetic diversity of a population generation (t ¼ 1), the amount of heterozygosity, H, changes
if it has suffered one or more bottlenecks in the past. This loss this way:
of genetic variation can lead to a loss of heterozygosity,
which, in some studies, has been correlated with a loss in H 1 ¼ H 0 ½1 1=ð2N Þ,
overall fitness (Frankel and Soulé 1981). Recessive traits
previously masked may now be expressed. Many of these The smaller the value of N, the greater the decline in
recessive alleles have deleterious or even lethal effects on an heterozygosity. A population of 50 individuals that began
organism. with a 0.5 level of heterozygosity would lose 1% of its
Small populations, including but not limited to those that heterozygosity in each generation (from 0.5 to 0.495). In
suffer a prolonged bottleneck, also experience genetic drift, contrast, a population of 10 individuals with the same initial
an effect that can occur in populations of any size as a result heterozygosity would lose 5% of its heterozygosity (from 0.5
of non-representative mating. Genetic drift occurs in all to 0.475) (Table 5.2). Mutations increase genetic variability
populations but has a greater effect in small populations and heterozygosity, but change in heterozygosity, ΔH, is also
because the proportion of non-representative mating affected by population size:
increases when the total number of matings is low. The

smaller the population, the greater the probability that the 1
ΔH ¼ H=ð2N Þ þ mH ¼ H m ,
sample (random matings of individuals) may represent 2N
Table 5.2 Heterozygosity and the effect of population size

Ht Population A (50) Population B (10)
H0 0.500 0.500
H1 0.495 0.475
H2 0.490 0.451
H3 0.485 0.429
H4 0.480 0.407
H5 0.475 0.387
Developed based on data from Caughley 1994. Table design by M. J.
Bigelow
Heterozygosity (H ) of smaller populations declines at a faster rate than
that of larger populations. Shown here are two populations: Population
A with a starting size of 50 and Population B with a starting size of 10.
Within 1 year, Population B has declined to a level of 0.475. In contrast,
it takes Population A 5 years to decline to that level
Fig. 5.4 The Mauritius kestrel (Falco punctatus), a species found only
where m is the addition of heterozygosity through mutation, on the island of Mauritius in the Indian Ocean, displays classic evidence
of loss of genetic diversity following a severe reduction in population
typically expressed as a rate. Populations reach an equilib- size (population bottleneck) during the 1960s and 1970s. (Photo cour-
rium level of heterozygosity (H, where ΔH ¼ 0) at tesy of M. Williams)
H ¼ 2Nm:
The smaller the population, the lower its equilibrium

heterozygosity.
Genetic drift and population bottlenecks can combine to
produce long-lasting effects on populations, even after
numerical recovery. Consider the Mauritius kestrel (Falco
punctatus) (Fig. 5.4), a small, rare falcon found only on the
island of Mauritius in the southwestern part of the Indian
Ocean. Following declines associated with use of pesticides
and destruction of its habitat from deforestation in the 1960s
and early 1970s, this population was believed to have been
reduced to a single breeding pair by 1974. Through careful
protection, habitat restoration and managed breeding, the
population had been restored to over 200 individuals by
1994 (Fig. 5.5a), and to 400–500 individuals, and over
200 breeding pairs, by the late 1990s (Groombridge et al.
2000). By examining DNA from living kestrels and compar-
ing specific loci from these to DNA obtained from museum
skins of kestrels from historical populations, Jim
Groombridge and his colleagues documented the loss of
genetic diversity that accompanied this bottleneck. Overall
allelic diversity declined by 57%. Several unique alleles
found in the ancestral population were not present in living
kestrels (Fig. 5.5b) and the restored population showed a
Fig. 5.5 (a) The population size of the Mauritius kestrel (Falco
57% reduction in heterozygosity (Table 5.3). punctatus) from 1940 through 1994. Note the severe reductions begin-
In every population, there are some alleles, known as ning in the 1960s when the population declined and remained at less
recessive lethal alleles, which, although unexpressed in a than 50 individuals. (b) DNA fingerprints (microsatellite genotypes)
from Mauritius kestrel museum skins (top) compared to DNA from
heterozygous state, will, in a homozygous condition, result
the same region in birds from the restored population (bottom). “b.p.”
in the death of the individual, which usually occurs in the refers to specific DNA base pairs. Dark bands represent the presence of
zygote, but for evolutionary biology purposes death only specific alleles. Note the reduction in the number of bands in the restored
needs to occur before reproduction. The proportion of such population, indicating reduction in allelic (genetic) diversity. (Reprinted
by permission from Springer Nature, from Groombridge, J.J., Jones, C.
alleles in a population, its lethal load, rises when heterozy-
G., Bruford, M.W., Nichols, R.A., 2000. “Ghost” alleles of the
gosity is lost during a period of population reduction. As Mauritius kestrel. Nature 403, 616. # 2000 Macmillan Magazines Ltd)
Table 5.3 Genetic diversity of the Mauritius kestrel and other kestrel this assumption. Instead, we must be able to estimate the
populations effective population size, Ne, the size of a randomly mating
Species A He Sample size (2N) population subject to the same degree of genetic drift as a
Endangered particular “real” population. Put another way, effective popu-
Mauritius kestrel lation size of a real population is equal to the size of an ideal
Restored 1.41 0.10 150 population that has the same amount of variance in allelic
Ancestral 3.10 0.23 52 frequencies. Hence, the effective population size is more
Seychelles kestrel 1.25 0.12 8
correctly called the “variance effective size.” Effective popu-
Non-endangered
lation size of a real population can also be defined, and
European kestrel 5.50 0.68 20
estimated, as size of an ideal population that has the same
Canary island kestrel 4.41 0.64 16
level of inbreeding, which is then more precisely referred to as
South African rock kestrel 5.00 0.63 20
Greater kestrel 4.50 0.59 20
“inbreeding effective size” (Loew 2002:242). Regardless of
Lesser kestrel 5.41 0.70 16 which measure is used, effective population size is affected by
Reprinted by permission from Springer Nature, from Groombridge, J.J.,
a number of variables, including variance in progeny number
Jones, C.G., Bruford, M.W., Nichols, R.A., 2000. “Ghost” alleles of the (brood or litter size), differential sex ratios, fluctuations in total
Mauritius kestrel. Nature 403, 616. # 2000 Macmillan Magazines Ltd numbers, and deviations from random mating systems
Mean numbers of alleles and average heterozygosity of the restored (Frankham 1980), and is inevitably lower than the real popu-
(post-bottleneck) Mauritius kestrel population compared with those of
the ancestral (pre-bottleneck) population and with other kestrel
lation size. To examine the effect on Ne of variation in the
populations. Note that with the exception of the Seychelles population, number of progeny, let N equal the population’s actual size
which also suffered severe population reduction, the ancestral Mauritius (census size) and σ2 the variance in progeny number. Then
kestrel population had less genetic diversity than other African and
European kestrel populations. With further reductions in the restored
4N
population, differences are now even greater Ne ¼ :
2 þ σ2
alleles are lost, heterozygosity is reduced, as in the Mauritius If the size of the population is 100, brood size ranges from
kestrel, and the probability of homozygous expression of 0 to 8 and the variance is 4, effective population size (Ne) is
lethal genes is higher if they are present. Genetic material 400 divided by 6, or 67, one-third less than the census
present in the founders represents a sample of the population size. An equalization of family size in a popula-
population’s historical levels of genetic variation, but, in tion should lead to an approximate doubling of effective
small founder groups, the genetic constituency of the population size. This prediction has proven true in experi-
founders may not be representative of the population. If, for mental tests. In Drosophila, populations subjected to equali-
example, only one of the members of the last remaining pair zation of family size had greater genetic variation and greater
of Mauritius kestrels in 1974 were heterozygous for a reces- reproductive fitness than populations in which family size
sive lethal allele, its expression would likely emerge in future was not equalized (Boriase et al. 1993).
generations and depress population survival rates. But, if For populations with unequal sex ratios, effective popula-
such lethal alleles were not present in these founders, the tion size is
population is not exposed to this risk.
The case of the Mauritius kestrel suggests that wild 4Nm N f
Ne ¼
populations may be resilient to genetic loss, so conservation Nm þ Nf
efforts can be successful even after severe reductions in
numbers and loss of genetic diversity. Nevertheless, the where Nm is the number of males and Nf the number of
Mauritius kestrel reveals that population reductions can females. Consider a population of 100 elk (Cervus elaphus)
have effects on genetic diversity, which do not rapidly disap- (Fig. 5.6). If there are 50 reproductive males and 50 reproduc-
pear with increases in population size alone. tive females, each male mates with one female, and each pair
represents a unique association of individuals, then effective
population size is 10,000 divided by 100, or 100. But there is
5.3.3 Genetic Drift and Effective no wild elk population anywhere with such a sex ratio, nor
Population Size are any that use such a mating system. Wild elk populations
have more females than males. In autumn, during the breed-
Theoretical consequences of genetic drift are normally calcu- ing period or “rut,” males gather groups of females
lated for an “ideal” population in which each individual (“harems”) that they defend against other males for exclusive
contributes gametes equally to a genetic pool from which the breeding privileges. Suppose a breeding population of
next generation is formed. Real populations rarely conform to 100 elk is composed of 10 males and 90 females. Each
Fig. 5.6 Elk (Cervus elaphus)

are an example of a species with a
harem mating system that reduces
effective population size. (Photo
courtesy of US National Park
Service)
male takes a harem of 9 females and defends it from other

males. In harem-mating systems like this, relatedness of Point of Engagement Question
offspring of females within a harem is higher than relatedness Work out the mathematics of the effective population
of offspring from females of different harems. In this revised size for 10 generations with a population of 100 in
scenario, the effective population size is 41090 divided every generation, then repeat the calculation a second
by 10 + 90, or 3600 divided by 100, or 36. time, letting one generation “crash” to 10 individuals.
Thus, sampling in a population of 100 like the one What happens to the effective population size?
just described would be equivalent to sampling in a popula-
tion of 36 that experienced random mating with equal sex
ratios. In general, the effective population size of a popula-
Like the effects of unequal family size and unequal sex
tion with a biased sex ratio is about one-third that of a
ratios, unequal population sizes should lead to increased
monogamous population with a balanced sex ratio. As a
levels of genetic drift and loss of heterozygosity. These
result, sampling error (genetic drift) associated with random
predictions also have been verified experimentally
mating in a population of 36 individuals is equivalent to
(Woodworth et al. 1994). The formulas assume random mat-
sampling error associated with mating in a population
ing, but that is rare in real populations. More complex
of 100 individuals with the sex ratio and mating system
equations are needed to determine effective population sizes
just described (Briton et al. 1994). Polygamous mating
if mating is non-random. Even if random mating is
systems associated with unequal sex ratios increase rates of
approximated, most populations will have a lower genetically
inbreeding and loss of genetic variation, which is why harem
effective population size than their census size. The problem
breeding structures should be avoided in captive breeding
of genetic drift gets bigger as effective population size gets
programs.
smaller. When effective population size is large, expected
Effective population size also changes when populations
variation in a genetic character is determined mainly by
fluctuate. If population size varies from generation to genera-
strength of selection for or against that character. When
tion, then the effective number is the harmonic mean (the
effective population size is less than a few hundred
reciprocal of the arithmetic mean of the reciprocals of a finite
individuals, expected variation of the character becomes
set of numbers):
largely independent of strength of selection and is instead
determined primarily by the balance between mutation and
1 1 1 1 1
¼ þ þ ... , drift (i.e., variation is determined by random events).
Ne t N1 N2 Nt
where Nt is the effective size of the population at generation t. 5.3.4 Bottlenecks, Small Populations and Rare
It can also be expressed as: Alleles

N e ¼ t=Σ 1=N e i , In genetic bottlenecks like those previously described in
bison, rare alleles can be lost quickly. The expected number
where Nei is the effective population size in generation i. of alleles, E(n), remaining after a genetic bottleneck is
Table 5.4 Decreasing population size influences the average number of alleles
Average number of alleles retained, given the original frequency of allele
Effective number of individuals (Ne) p1 ¼ 0.70, p2 ¼ p3 ¼ p4 ¼ 0.10 p1 ¼ 0.94, p2 ¼ p3 ¼ p4 ¼ 0.02
1 4.00 4.00
50 3.99 3.60
10 3.63 2.00
6 3.15 1.64
2 2.02 1.23
1 1.48 1.12
Developed based on equations from Frankel and Soulé 1981. Table design by M. J. Bigelow
In this case, four alleles are observed – one with a high frequency and three with lower frequencies. Rare, less common alleles are more likely to be
lost during a bottleneck. These rare alleles are typically not essential in the initial environment. However, as the environment changes they might be
crucial for survival
X 2Ne
m 1 pj populations, but we will examine a few that are widely used
in conservation and prevalent in the literature of conservation
biology. Understanding capabilities and limitations of indi-
where m is the number of alleles prior to the bottleneck, p the
vidual techniques can not only help conservationists apply
frequency of the jth allele, and N the effective number of
these techniques appropriately, but also permit them to assess
individuals at the bottleneck. Suppose that m ¼ 4 and one
the value and validity of the results obtained from such
allele is common, but the other three are rare. Look what
techniques in studies reported in scientific literature.
happens to the average number of alleles (Table 5.4) as the
effective number of individuals drops from 50 to 1. The rarer
the allele, the more likely that it will be lost (Frankel and 5.3.5.2 The Polymerase Chain Reaction (PCR)
Soulé 1981). The development of the polymerase chain reaction (PCR) as
One of the earliest stated goals of conservation biology a standard genetic technique revolutionized conservation
was retention of 90% of a population’s genetic variability for biology beginning in the late 1980s. Today, this technique
200 years (Soulé et al. 1986). The loss of genetic variation is is widely used to determine genotypes of individual species
reduced (and the probability of meeting this goal improves) by employing a relatively simple reaction in which a short
as the effective population size grows to about 1000 region of a DNA molecule, even as little as a single gene or
individuals, although the precise number in a given popula- smaller, is copied repeatedly by a DNA polymerase enzyme.
tion will be influenced by the particular traits of the species’ In standard PCR, any region of a DNA molecule can be
biology. Above this level, further increases in effective pop- chosen as long as the borders (beginning and ending
ulation size do not usually increase genetic variability in the sequences) are known. To carry out the PCR, two
population (Lynch 1996). But effective population size is oligonucleotides (short pieces of DNA) hybridize to the
often only one-tenth to one-third the number of breeding DNA molecule, one to each strand of the double helix at
adults in the population for reasons noted previously, includ- opposite ends of the targeted region. These act as primers for
ing unequal family sizes, unequal sex ratios, and unequal the subsequent DNA synthesis and delimit the region to be
population sizes in different years. Thus, the Ne > 1000 crite- amplified. An enzyme, such as Taq polymerase from the
rion actually suggests the need for a stable population of bacterium Thermus aquaticus – an organism native to hot
3000–10,000 breeding adults in each generation to prevent springs – whose enzymes, including its DNA polymerase,
long-term loss of genetic variation. resist denaturization when exposed to heat, is added (Brown
1995). The polymerase is incubated in the solution and
facilitates production of new complementary DNA strands.
5.3.5 Measuring Genetic Change with Genetic The mixture is then heated so that new strands detach from
Technology original DNA. The strands are then cooled, allowing more
primers to attach at their respective positions, including on
5.3.5.1 General Considerations newly synthesized strands (Brown 1995). The polymerase
It is not just increased concern over genetic variation that has carries out a second round of DNA synthesis and the cycle
driven the science of genetics to prominence in conservation can be repeated many times, eventually resulting in synthesis
biology, but also the increasing precision of techniques of several hundred million copies of the amplified DNA
associated with genetic analysis, particularly at the molecular fragment which can then be analyzed in various ways
level. We will not review all of the genetic techniques cur- (Brown 1995) (Fig. 5.7). One of the most common and useful
rently used to assess genetic variation in individuals and analyses in conservation is the direct sequence analysis of the
Fig. 5.8 Schematic example of a DNA “fingerprint” produced by

fractionation of different bands of DNA associated with microsatellite
markers. Restriction enzymes cut the DNA at specific sequences. The
amount of DNA between sequences differs between individuals; there-
fore, unique patterns will appear. Because DNA has a slight negative
charge it will move towards the positive electrode. Smaller pieces move
faster and are found near the base of the gel. This process is referred to as
fractionation using gel electrophoresis. Following separation, the DNA
double strands are denatured to single strands using heat or chemical
treatment. A DNA print of the gel is transferred and fixed onto nitrocel-
lulose paper. The single stranded DNA is subjected to a probe specific
for a gene within the microsatellite. Unique binding patterns of the
radioactive probe can be observed using autoradiography. (Illustration
by M. J. Bigelow)
PCR products, resulting in identification of the genotype of

the organism from which material was obtained. The lengths
of the amplified regions can also be visualized by electropho-
resis in an agarose gel after staining with dyes (Fritsch and
Rieseberg 1996) (Fig. 5.8).
PCR analysis can be used as a postmortem technique, as
on museum specimens (Morin and Woodruff 1996), and only
minute amounts of DNA are required. The analysis can also
include genetic fingerprinting to identify particular
individuals. Using Reverse Transcription PCR (RT-PCR),
RNA can be amplified and the amount of messenger RNA
(mRNA) can be determined in the sampled tissues, providing
an accurate index of the activity of genes. From this index, it
is possible to infer many aspects of the physiological status of
the organism, including reproductive and nutritional
parameters (Morin and Woodruff 1996).
5.3.5.3 DNA Fingerprinting: Use of Satellite

Markers
Fig. 5.7 Schematic representation of the mechanism for a polymerase
chain reaction (PCR). PCR is a useful technique that is used to multiply
Many sections of an organism’s genome consist of short
(i.e., amplify) the amount of DNA of interest. (Reprinted by permission sequences of DNA that may be repeated up to one million
from McGraw-Hill Publishers) times. Such segments often have different sequences of bases
than other portions of DNA, and therefore different molecu-
lar densities. These repetitive sequences are known as satel-
lite DNA (Sudbery 1998).
Some types of satellite DNA are interspersed throughout

an organism’s genome. There are two classes of such DNA,
minisatellites and microsatellites. Larger minisatellites con-
sist of 10–100 base pairs repeated in tandem arrays that vary
in size from 0.5 to 40 kb (kb stands for “kilobase,” a unit of
1000 base pairs). Some loci on homologous chromosomes
within minisatellites are highly variable in length. Hence,
minisatellites also are referred to as variable number tandem
repeats (VNTR), and their variability forms the basis for one
type of DNA fingerprinting (Fig. 5.8). Differences in length
are unique to individuals, thus providing a basis for
identification.
Compared to minisatellites, microsatellites are smaller,
consisting of short tandem repeats (STRs) only two to four
nucleotides in length (Sudbery 1998). Like minisatellites,
microsatellites are polymorphic and provide valuable genetic
markers, but they have a more uniform distribution in the
genome. This makes them useful in determining pedigrees of Fig. 5.9 Schematic diagram of a loop of mitochondrial DNA. Unlike
individuals as well as in forensic applications to determine nuclear DNA, mitochondrial DNA is not enclosed within a membrane.
the origin of individual animals or wildlife products. Rather, it is found within the matrix of the organelle. (Figure courtesy of
National Human Genome Research Institute)
Microsatellites also can be evaluated with less expensive
and labor-intensive techniques, and their variability is often
species-specific, making them useful for many applications in Theoretically, phylogenies derived from mtDNA are not
conservation (Hedrick 1999). affected by historic changes in effective population sizes,
unequal sex ratios, or unequal family sizes, although it is
5.3.5.4 Mitochondrial DNA: Using mtDNA often possible to detect evidence of past genetic changes in
for Phylogenetic Analysis the population from local variations in mtDNA sequence
Mitochondria are organelles found in all eukaryotic cells that patterns. Many taxonomists consider phylogenetic analysis
function primarily in converting chemical energy into energy of mtDNA to be a superior, more objective method of deter-
that can be used directly by the cell. Unlike most other mining phylogeny because it is based entirely on quantitative
organelles, mitochondria have their own, non-nuclear com- characters. Because it is maternally inherited, mtDNA can be
plement of DNA. Each mitochondrion contains 1–15 copies used to determine a maternal lineage in an individual or
of a circular genome, much smaller than the corresponding group, but not a paternal one.
nuclear DNA genome. This mitochondrial DNA (mtDNA) The small size of the mtDNA genome means that its
(Fig. 5.9) is specific for particular cellular functions sequences have long been known, and it is easy to identify
associated with mitochondria, including synthesis of subunits polymorphisms within it. This trait is complemented by the
that function in cellular respiration, units that code for syn- fact that certain regions of mtDNA evolve faster than single-
thesis of the transfer RNA (tRNA) of each amino acid, and copy nuclear genes in mammals (Wilson et al. 1985), so they
DNA involved in synthesis of ribosomal RNA (rRNA) that can be especially useful for studying differences at the popu-
functions in protein synthesis. lation level. Overall, mtDNA evolves faster than nuclear
mtDNA also differs from nuclear DNA in that it is mater- DNA, permitting studies of recent evolution that nuclear
nally inherited as a linked set of genes, passed on to progeny DNA would not record.
in the cytoplasm of the egg cell (Cronin 1993; Sudbery Using mtDNA sequences for phylogenetic studies – for
1998). Thus, there is no recombination between maternal example to resolve questions of relatedness in similar
and paternal genomes. This mode of inheritance results in species – or in forensics to determine whether meat, pelts,
rapid mtDNA differentiation relative to nuclear genes and horns, or other animal parts come from an endangered spe-
makes the construction of phylogenetic trees more straight- cies is increasingly common, and part of a larger array of
forward, because without recombination, the number of genetic techniques that make use of DNA barcoding, as is the
nucleotide differences between mtDNA genomes of different increasing use of DNA that can be collected from the envi-
individuals, populations, or species can be assumed to be a ronment itself (environmental DNA or eDNA) (Information
direct measure of phylogenetic relatedness (Sudbery 1998). Box 5.1).
Information Box 5.1: Barcoding and eDNA – New Information Box 5.1 (continued)
Genetic Techniques with Conservation Applications (Barnes and Turner 2016). eDNA has also been used to
DNA barcodes are short standardized DNA sequences identify multiple species from the same sample (Barnes
that are species-specific and therefore can be used for and Turner 2016; Wilson et al. 2016). Finally, collec-
species identification. DNA barcodes can be especially tion of eDNA using remote and autonomous methods,
useful in identifying species found in biological including robotic sampling, can permit investigation of
inventories of sites or regions where many members organisms in places currently inaccessible to humans,
of a given taxon may lack formal description. For such as benthic environments of the deepest parts of the
example, Wilson et al. (2016) found that DNA ocean (Barnes and Turner 2016). These and other
barcodes could be used effectively for this purpose in applications of eDNA hold promise of greatly
arthropods where, in regions like southeast Asia, many expanding the effectiveness of conservation efforts on
site- and area-specific inventories have detected species multiple fronts.
not previously described. Such barcode data is now
being organized in a worldwide, internet-accessible
5.3.5.5 Genomics
library, the Barcode of Life Data (BOLD) System that
All the techniques described so far are based on analysis of
will have increasing use in studies of taxonomy, ecol-
small fragments of an organism’s total genomic complement,
ogy, evolution and conservation of life on Earth
usually one gene or non-coding region. Genetic analysis at a
(BoldSystems 2018).
particular locus is a powerful tool but limited in its capacity to
DNA barcoding can also be used in larger scale,
resolve the evolutionary history of some organisms as well as
minimal impact approaches to vertebrate monitoring,
other aspects of genetic structure. Today there are ways to
population assessments, and dietary analyses that can
move from analysis of a small fragment of DNA to the entire
make use of secondary sources of DNA, such as hair,
genome. Until recently, genomic approaches were impracti-
feces, or fragments of DNA found in the environment
cal for all but the largest research organizations, but today
itself, such as soil, water, or air, so-called environmen-
advanced technologies and declining costs have made geno-
tal DNA or eDNA (Barnes and Turner 2016; Wilson
mic investigation increasingly accessible, aided by the fact
et al. 2016; Panko 2017). The rapidly expanding study
that genomic sequence data are now available in public
of eDNA has generated unprecedented ability to detect
databases. As molecular geneticist Matthew Hudson noted,
species and conduct genetic analyses for conservation,
“The cost, labor and time required to perform sequencing on
management, and research, particularly where collec-
the scale needed for whole genome analysis has fallen signif-
tion of whole organisms is impractical or impossible
icantly as a result of the recent development of new
(Barnes and Turner 2016). There are multiple
technologies, and this trend is likely to continue” (Hudson
applications for eDNA analysis in conservation, espe-
2008).
cially in detection of target species, either those of
conservation interest, or as an “early warning system”
for the presence of potentially invasive species such as
5.4 Solving the Problem of Inbreeding
Asian carp (Hypophthalmichthys nobilis and
H. molitrix) into the US Great Lakes, New Zealand
5.4.1 What Do We Mean by “Inbreeding”
mud snail (Potamopyrgus antipodarum) in US fresh-
and How Would We Measure It?
water streams, or many species of aquatic invasive
organisms that can be transported and introduced
Genetic technology in conservation is only as good as the
worldwide through ballast water of ships.
problems it can solve, and the genetic problems of small and
Threatened and endangered species and other spe-
endangered populations are many. One problem is finding a
cies of conservation concern are another category of
mate, exacerbated in small populations because there is
“targets” that can benefit from eDNA analysis because
increased probability of finding the wrong mate. That is,
they are often difficult to observe and illegal to trap or
individuals in small populations are more likely to mate
handle. For example, eDNA assays have been used to
with close relatives with whom they share many alleles.
detect eastern hellbender (Cryptobranchus
This situation is known as inbreeding, a problem we have
a. alleganiensis), a large salamander of conservation
already identified as a threat to endangered species.
concern in the US states of Indiana and Missouri, while
Inbreeding is the production of offspring related by descent.
in the UK eDNA assays have been used for detection of
Consider three ways inbreeding can be measured. Inbreeding
the threatened great crested newt (Triturus cristatus)
can be assessed as a measure of: (1) shared ancestry in the
(continued)
5.4 Solving the Problem of Inbreeding 183
maternal and paternal lineages of an individual; (2) genetic “d” is meant to signify that this value of F is a measure of the
drift in a finite population; or (3) a system of mating in a averaged inbreeding by descent of all members of the local
reproducing population. Unless a species has strong barriers population, or deme (Templeton and Read 1994). Fd then
to inbreeding, each kind of inbreeding grows stronger as represents the average probability of inbreeding by descent, a
population size declines and must be carefully measured to measure of the effect of genetic drift on a population relative
make informed management decisions about breeding to an “ideal” population experiencing completely random
strategies for small populations (Templeton and Read 1994). mating.
The first concept of inbreeding, the measure of shared Remember that the first type of inbreeding, pedigree
ancestry of an individual in its maternal and paternal lines, inbreeding, increases in magnitude as population size
has been called pedigree inbreeding (Templeton and Read declines. In the second type of inbreeding, inbreeding by
1994), inbreeding that occurs when parents share ancestors descent, the value of Fd also increases as population size
(Fig. 5.10). This type of inbreeding can be quantified as the decreases and, for a given population size, also increases
inbreeding coefficient, symbolized by Fp (pedigree over time, expressed by the relationship
inbreeding). The value of Fp, which varies from 0 to 1, can
be calculated only for an individual of known pedigree. Fp FdðtÞ ¼ 1 ½1 1=2Nt ,
measures the amount of ancestry an individual shares with its
maternal and paternal lines, a form of inbreeding that where t is equal to time in generations. The larger the value of
intensifies as a population decreases in size. t, the closer Fd(t) comes to 1 (a completely inbred population).
The second concept of inbreeding is as a measure of Thus, Fd(t) will eventually reach a value of 1, and how fast it
genetic drift in a population, sometimes called inbreeding does so is a function of population size (Fig. 5.11). The
as nonrandom mating (Keller and Waller 2002) or smaller the population, the faster it will become inbred.
inbreeding by descent. If we knew individual values of Fp Finally, inbreeding can be used as a measure of a system
for every individual in a population, added these values of mating in a population, quantified as a value called the
together, and divided the sum by the number of individuals, panmictic index, f, also called inbreeding by population
the resulting quotient would be the average probability of subdivision (Keller and Waller 2002). The panmictic index
inbreeding by descent, symbolized as Fd. Here the subscript measures inbreeding as a deviation from a reference popula-
tion, which has a system of mating in which alleles at a locus
are paired in proportion to their frequencies in the overall
population (by definition, random mating). The panmictic
Fig. 5.10 Pedigree of male song sparrow (Melospiza melodia)

73543 from the song sparrow population of Mandarte Island (British
Columbia, Canada), a heavily inbred population. Note here, for exam-
ple, that 73543’s maternal grandfather (56916) is also his paternal uncle. Fig. 5.11 Relationship between the proportion of inbred individuals in
(Reprinted by permission from Elsevier, from Keller, L.F., and Waller, a population and population size. The smaller the population, the less
D.M., 2002. Inbreeding effects in wild populations. Trends in Ecology time it will take to become completely inbred. (Design by M. J.
& Evolution 17, 230–241. # 2002 Elsevier Science Ltd) Bigelow)
index thus evaluates deviations from heterozygosity

frequencies expected under random mating, so
f ¼ 1 H o =H e ,
where He is expected heterozygosity under random mating

and Ho is observed heterozygosity. In randomly mating
populations, frequency of heterozygosity is defined by the
Hardy-Weinberg equation. For two alleles, p and q, that is
ðp þ qÞ2 ¼ p2 þ 2pq þ q2 :
Thus, the expected frequency of the heterozygote is 2pq. For

example, if the frequency of allele p is 0.6 and the frequency
of allele q is 0.4, then the He is 2 0.6 0.4 ¼ 0.48.
Observed heterozygosity (Ho) can be calculated from genetic Fig. 5.12 Relationship between “inbreeding load” (the number of
measurements of sampled individuals. If observed heterozy- lethal equivalents, β) and the strength of inbreeding depression.
gosity is greater than expected, f < 0, and the population has a (Reprinted by permission from Elsevier, from Keller, L.F., and Waller,
reproductive system that avoids inbreeding. If observed het- D.M., 2002. Inbreeding effects in wild populations. Trends in Ecology
& Evolution 17, 230–241. # 2002 Elsevier Science Ltd)
erozygosity is less than expected, f > 0, and inbreeding is not
avoided. The value of the panmictic index is its ability to
quantify the degree of avoidance of inbreeding in a when historically large, outcrossing populations suddenly
population. decline to only a few members. Remaining individuals may
Measurement of inbreeding is especially important in or may not be related. However, with the limited mate
managing breeding in a captive population. Using values of choices now available to them, high average relatedness
Fp in a captive breeding program, for example, a manager can will result in just a few generations. The population then
determine which potential breeding pairs would produce experiences reduced survival and fecundity. As mate choice
inbred versus non-inbred offspring. To minimize inbreeding, is reduced to related individuals, inbreeding depression may
a manager could mate individuals least related to one another. increase (Fig. 5.12). When the degree of relatedness of
An animal which is itself inbred should not be excluded from individuals in the population (“inbreeding by descent”) is
the breeding pool but breeding pairs should be selected to regressed against one or more traits affecting fecundity or
avoid creation of inbred offspring. Using the value of Fd, a survival, the resulting relationship can be used to calculate
manager could determine the effect of genetic drift on a the degree to which increased mortality or lower fecundity is
population and the degree of heterozygosity present in that associated with increased relatedness.
population and, from such data, determine whether current Inbreeding depression is an especially well-documented
population size is sufficient to maintain an acceptable level of problem in captive populations of vertebrates (Frankham
heterozygosity. Using the value of f, a manager could also 1995a). In a classic early study by Ralls and Ballou (1983),
determine if current mating systems are likely to lead to 42 of 45 captive, inbred vertebrate populations showed
avoidance or encouragement of inbreeding, and then act reduced juvenile survival compared to outbreeding
accordingly. populations of the same species. In subsequent studies of
captive populations of fruit flies, Drosophila melanogaster,
geneticist Richard Frankham created experimental
5.4.2 The Problem of Inbreeding Depression populations in which he manipulated density, rates of
inbreeding, and levels of environmental stress. In these
When populations become inbred, genotypic frequencies are experiments: (1) inbreeding and consequent loss of genetic
skewed toward increased proportions of homozygous diversity reduced resistance of flies to disease; (2) in inbred
individuals and heterozygosity declines. As the proportion populations, extinction rates rose as environmental stress was
of homozygous individuals increases, so does the manifesta- increased by adding additional stress factors (Fig. 5.13);
tion of recessive traits. Recessive traits can only be expressed (3) adaptive evolutionary potential (capacity for long-term
in a homozygous condition but are maintained in the popula- genetic change) was reduced in small populations as environ-
tion by heterozygous carriers. Inbreeding depression, a pat- mental stress was increased; and (4) rates of inbreeding
tern of reduced reproduction and survival that occurs on increased under stressful conditions (Reed et al. 2002;
account of inbreeding (Frankham et al. 2002:24), can arise Frankham 2005). In other studies of populations of
5.4 Solving the Problem of Inbreeding 185
use the same relationship to make comparisons between

related populations. For example, suppose that levels of
heterozygosity of a small island population of individuals
and that of the larger mainland population they originated
from are both known, and are respectively Hisland ¼ 0.35 and
Hmainland ¼ 0.81. Then the effective inbreeding coefficient
indirectly estimated from these values is
F e ¼ 1 ð0:35=0:81Þ ¼ 1 0:43 ¼ 0:57:
Here inbreeding in the smaller island population reduced its

heterozygosity by 57% compared to the mainland population.
Alternatively, some effects of inbreeding can be evaluated
Fig. 5.13 Proportion of populations of Drosophila melanogaster sur- by measuring the rate of juvenile survival, which is calculated
viving at different inbreeding coefficients (F) for inbred (full-sib mat- as
ing) populations in benign (no stress), single stress factor, and variable
stress factor environments. (Reprinted by permission from Wiley, from
Frankham, R., 2005. Stress and adaptation in conservation genetics.
ln ðSÞ ¼ A þ BF
Journal of Evolutionary Biology 18, 750–755. # 2005 European Soci-
ety for Evolutionary Biology) where S is the juvenile survival rate, A is the instantaneous
rate of juvenile mortality in progeny of unrelated parents, B is
the rate when the line is completely inbred (H ¼ 0) and F is
deliberately inbred domestic animals and plants, up to 95% the inbreeding coefficient (Ralls et al. 1979). This equation
became extinct after eight generations of brother-sister can be expressed graphically as the line described by S ¼ A –
matings (animals) or three generations of self-fertilization BF to see how the rate of survival is affected by inbreeding.
(plants) (Frankel and Soulé 1981). In wild populations, A can then be estimated if B is known, and vice versa
inbreeding depression has been documented in fish, snails, (Caughley 1994).
lions, shrews, white-footed mice, and plants (Frankham If you know the F coefficient and a measure of the
1995b). individuals’ fitness in the population, you can estimate the
severity of inbreeding depression, δ, as the proportionate
decline in mean fitness due to a given amount of inbreeding.
5.4.3 Measuring the Inbreeding Coefficient The relationship is
The inbreeding coefficient, F, is a measure of loss of hetero- δ ¼ 1 ðfitness of inbred offspring=

zygosity in a population due to effects of inbreeding. There- fitness of outbred offspringÞ:
fore, one can estimate the inbreeding coefficient from
changes in heterozygosity over time. Earlier we defined the To be useful, this measurement must be more specific. For
concepts associated with this measure. Now we will calculate example, let’s replace “fitness” in the above equation with
it. This measure of the inbreeding coefficient, known as the “survival rate.” Conventionally, magnitude of inbreeding
“effective inbreeding coefficient” (Fe) is estimated as depression is expressed as average reduction in mean fitness
value per 10% increase in the F coefficient (Van Oosterhout
F e ¼ 1 ðH t =H 0 Þ, et al. 2000). Given this information, we could estimate the
value of inbreeding depression as
where, as you recall, Ht is the level of heterozygosity at time
or generation t and H0 is the level of heterozygosity in the Survival rate at F ¼ 0:25
1 :
population at a previous time. For example, if the level of Survival rate at F ¼ 0:0
heterozygosity in generation t is 0.4 and the level of hetero-
zygosity in a previous generation 0 was 0.8, then the value of Suppose that the survival rate at F0.25 ¼ 0.2 and at
Fe is F0.0 ¼ 0.8. Then the cost of inbreeding is 1–(0.2/0.8) or
1–0.25 ¼ 0.75. Conceptually, the cost of inbreeding or
1 ð0:4=0:8Þ ¼ 1 0:5 ¼ 0:5: inbreeding load is the proportional decline in survival that
can be attributed to inbreeding of a given magnitude
This means that inbreeding has reduced heterozygosity over (Fig. 5.14). Such declines are well documented. In 38 species
this period of time in this population by 50%. One can also of mammals, Ralls et al. (1988) estimated actual average cost
Fig. 5.14 Relationship of relatedness in mating to levels of inbreeding

depression, which is the decline in fitness (reduced survival and fecun- Fig. 5.15 Survivorship of inbred (dotted line) and non-inbred (solid
dity) associated with increased frequencies of mating among closely line) white-footed mice (Peromyscus leucopus noveboracensis) in a
related individuals. The probability of an individual mating with a mixed deciduous forest in Illinois (USA). Bars represent standard errors
relation increases as the population size decreases. (Design by M. J. of estimates. Non-inbred animals had higher survivorship than inbred
Bigelow) animals in all time intervals. Although none of the differences between
groups differ significantly for any one estimate, when estimates are used
as repeated measures of survivorship for groups the difference is statis-
of inbreeding to be 0.33. Can this inbreeding load cause tically significant. (Reprinted by permission from the American Associ-
ation for the Advancement of Science, from Jiménez, J.A., Hughes, K.
endangerment and extinction?
A., Alaks, G., Graham, L., Lacy, R.C., 1994. An Experimental Study of
Inbreeding Depression in a Natural Habitat. Science 266, 271–273. #
1994 AAAS)
5.5 Can Inbreeding Cause Extinction?
Van Oosterhout et al. (2000) executed a carefully
5.5.1 Experiments on Inbreeding designed laboratory experiment on the effects of inbreeding
in squinting bush brown butterfly (Bicyclus anyana),
In a controlled experiment involving both field and labora- manipulating levels of inbreeding, gene flow, and population
tory environments, Jiménez et al. (1994) estimated survivor- size in different populations while measuring various fitness
ship of inbred and non-inbred white-footed mice components in each group. One key fitness metric, lifetime
(Peromyscus leucopus noveboracensis) in a mixed deciduous female fecundity, was inversely proportional to the
forest in Illinois, USA. Mice in both treatments had been inbreeding coefficient of the female parent (Van Oosterhout
raised in a laboratory but were all descendants of wild mice. et al. 2000). Smaller and more inbred populations also had
Mice were released into the field and recaptured at regular more sterile egg clutches. Other fitness indices, including
intervals. Trapping was done on surrounding adjacent habitat zygote and juvenile mortality, adult male and female longev-
to estimate the proportions of mice that left the release site. ity, and male development were all detrimentally affected by
Most mice were captured within 50 m of their release sites, inbreeding (Van Oosterhout et al. 2000).
and emigration rates were not different between the two
treatments. Based on release-recapture ratios of marked
mice in the inbred and non-inbred treatments, Jiménez et al. 5.5.2 Inbreeding in Wild Populations
(1994) determined that, over a 10-week period, non-inbred
mice had consistently higher survivorship than inbred mice In addition to increased mortality and decreased fecundity
(Fig. 5.15). Some differences were sex-specific. For example, associated with inbreeding depression, inbreeding also
male mice in both treatments lost weight in the first few days affects the genetic structure of populations. For example,
after release. Non-inbred males regained their weight, but Sullivan (1996) found that among populations of Colorado
inbred males did not and lost weight throughout the experi- chipmunks (Tamias quadrivattus) in the southwestern United
ment. The results of this study are consistent with an analysis States, inbreeding depression was more serious in isolated
of 33 population data sets which showed that approximately populations with limited gene flow, even if the overall popu-
40% of variation in fitness could be explained by variance in lation was large and geographically widespread. The most
heterozygosity, a reliable index of inbreeding (Reed and isolated populations had little or no genetic variation at up to
Frankham 2001). 30 loci (Sullivan 1996).
5.5 Can Inbreeding Cause Extinction? 187
A conservation biologist concerned with the pressing studied the subunits from 1993 to 1995, examining effects
question of extinction might be justified, if not polite, in of area, past trends in population size, density of butterflies in
asking, “So what? Are these chipmunk populations really in nearest neighboring populations, incidence of cattle grazing,
any danger because of inbreeding?” A dramatic answer to and heterozygosity on various measures of fitness, including
this question was provided, in part, by a serendipitous natural larval survival, adult longevity, egg hatching rate, and risk of
experiment on Mandarte Island off the coast of British extinction of the population subunit. The level of heterozy-
Columbia, Canada. Here, researchers involved in a long- gosity found at seven polymorphic DNA sites on one micro-
term study on song sparrows (Melospiza melodia) had been satellite locus in the butterfly genome was used as an index of
marking individuals and documenting genealogies over sev- inbreeding. Heterozygosity is a potential and relative index of
eral generations, such that an extensive amount of population inbreeding, but not a direct measure of it. Nevertheless, het-
demographics had been determined, including inbreeding erozygosity and inbreeding should be directly correlated. As
coefficients of most individuals. The population crashed in inbreeding increases, heterozygosity should decline. As
1989, with 206 individuals dying and only 10 surviving. Saccheri et al. (1998) stated about these subgroups, “The
Every sparrow with an inbreeding coefficient of 0.0625 or variance in inbreeding among populations is expected to be
higher died. In contrast, the 10 survivors had an average high. . .because there is substantial gene flow in many dense
inbreeding coefficient of only 0.0065, and only three regional networks of local populations, but also close
of these had a known history of inbreeding (Keller et al. inbreeding in many local populations that are extremely
1994). small and quite isolated. Thus, differences in average hetero-
The most definitive results have come from the study of zygosity of local populations, even if based on a limited
butterfly populations in Finland where Ilik Saccheri and his number of polymorphic loci, should reflect real differences
colleagues examined 42 population subunits of the Glanville in the degree of inbreeding” (Saccheri et al. 1998:419).
fritillary butterfly (Melitaea cinxia) (Fig. 5.16), living on the As in Van Oosterhout’s study, Saccheri et al. (1998) found
Ǻland Islands off Finland’s southwest coast. Subunits that female lifespan, percent of eggs hatching (based on
consisted of numerous, small, more-or-less isolated average group size after hatching), and larval weight (taken
populations that lived in dry meadows (Saccheri et al. as an index of larval survival) were all correlated with
1998). Individual population subunits were often transient, increasing heterozygosity. More homozygous, and presum-
with overall turnover averaging 200 extinctions and ably more inbred females, had shorter lifespans, lower rates
114 colonizations per year. Saccheri and his colleagues of hatching success, and smaller, less viable larva. But the
difference between Saccheri’s findings and Van Oosterhout’s
was that Saccheri et al. also documented the fate of each of
the 42 population subunits. Using data from a larger sample
of 336 other (different) population subunits, Saccheri et al.
constructed a model that predicted the likelihood of extinc-
tion using both ecological factors and heterozygosity. When
the model was applied to the 42 studied subunits,
investigators found that risk of extinction was, to varying
degrees, related to ecological and habitat variables, but het-
erozygosity also was an important factor, explaining 26% of
variation in extinction risk. As the number of heterozygous
loci increased, the risk of extinction declined dramatically
(Saccheri et al. 1998).
The results of Saccheri et al.’s study are consistent with
findings of more recent reviews which have confirmed not
only that inbreeding depression reduced traits associated with
early fitness (e.g., juvenile survival), but also reduced overall
fitness in natural populations (Hedrick and Garcia-Dorado
Fig. 5.16 The Glanville fritillary butterfly (Melitaea cinxia), a species 2016). In an extensive survey of bird and mammal population
in which inbreeding was a causative factor in extinctions of populations studies, conservation geneticists Lukas Keller and Donald
in southern Finland. (Photo courtesy of H. Aarnio; reprinted by permis- Waller determined that effects of inbreeding were pervasive,
sion from Wiley, from Ojanen, S.P., Nieminen, M., Meyke, E., Pöyry,
J., Hanski, I., 2013. Long-term metapopulation study of the Glanville
affecting birth weight, survival, reproduction, resistance to
fritillary butterfly (Melitaea cinxia): survey methods, data management, disease, predation and environmental stress. For example, in
and long-term population trends. Ecology and Evolution 3, 3713–3737. the endangered red-cockaded woodpecker (Picoides
# 2013 The Authors) borealis) of the southeastern United States, they noted that
inbreeding “resulted in significantly reduced hatching rates, problem of outbreeding, especially in plants, is one that
fledgling survival and recruitment to the breeding popula- deserves further study and consideration in current conserva-
tion” (Keller and Waller 2002). Also reviewing numerous tion efforts.
plant studies, Keller and Waller found that, in plants,
inbreeding affected seed set, germination, survival, and resis-
tance to stress (Keller and Waller 2002). 5.6 Landscape Genetics and Habitat
Fragmentation
5.5.3 Outbreeding Depression 5.6.1 Habitat Fragmentation: A Genetic Threat

to Large and Small Populations
Although usually less significant in its effects (or perhaps
simply less studied) than inbreeding depression, the problem Habitat fragmentation is one driver of the current extinction
of outbreeding depression deserves mention, both to define crisis (Chap. 3), resulting in overall habitat loss, reduction in
the concept and to note the characteristics of populations that size of habitat patches, and increased isolation of the habitat
may suffer from its effects. As the external environment may patches as habitat “fragments” (Honnay and Jacquemyn
mold local adaptations by natural selection, the internal 2007). Populations of plants and animals remaining in such
genetic environment of a population may lead to production fragments have reduced and more isolated populations which
of local complexes of alleles that interact in a mutually are less genetically diverse than those in larger, more contig-
favorable way. Thus, population subunits highly adapted to uous habitat areas.
local conditions may evolve co-adapted allele complexes in In a meta-analysis of the effects of habitat fragmentation
which alleles must be inherited together to produce adaptive on plant genetic diversity, Olivier Honnay and Hans
effects. When individuals from such normally inbreeding Jacquemyn of Belgium’s University of Leuven examined
populations mate with individuals from other populations of the relationship between four measures of genetic diversity
the same species (“outbreed”), they may decline in fitness as including expected heterozygosity (He), percent polymorphic
their own genetic combinations are disrupted. loci (P), inbreeding coefficient (FIS), and allelic richness (A)
Outbreeding depression is most common in plants and plant population size. They found, after examining
(Frankham 1995a), especially in populations that have devel- relationships in 52 plant species, that “. . .small populations
oped high levels of self-pollination (“selfing”). In fact, selfing consistently contained significantly less genetic
has evolved repeatedly in plant taxa that were previously variation. . .than large populations. Population size had a
outcrossing (Stebbins 1957; Grant and Grant 1965; Raven lower effect on He (gene diversity, measured as expected
1979). Selfing is advantageous where pollinators are at low heterozygosity) than on P and A, suggesting that alleles lost
density and the accompanying probability of being pollinated through habitat fragmentation and population size reduction
is therefore also low, but selfing populations often show were mainly those initially present at low densities. . .”
reduced reproductive performance and vigor (outbreeding (Honnay and Jacquiemyn 2007:826). Such results follow
depression) when crossed with different populations. In the expectations predicted by genetic theory already reviewed.
state of Oregon (USA), for example, Pyrrocoma racemosa What was surprising was the discovery that “. . .population
var. racemosa – a rare, endemic wetland variety of clustered genetic diversity (He, A, and P) was also eroded in species
goldenweed found on only six sites in the state – showed that were considered common. Even when historically or
evidence of outbreeding depression that could limit the very naturally fragmented populations of rare species were omit-
conservation efforts intended to protect it. Botanist Paul ted from the analysis, no difference between rare and com-
Severns pursued the possibility of increasing numbers of mon species in population genetic response to habitat
P. racemosa var. racemosa by personally and manually fragmentation was found. These results are dramatic in their
cross-pollinating plants from different sites, but discovered indication that many more plant species than previously
that outcrossed plants showed reduced reproductive perfor- assumed may be vulnerable to genetic erosion and loss of
mance, suggesting incompatibility with plants from different genetic diversity as a result of ongoing fragmentation pro-
populations, even those in close proximity to one another. cesses” (Honnay and Jacquemyn 2007:828). Thus,
“These results,” noted Severns, “suggest that there are fragmented habitats might be unable to support plant
genetic incompatibilities between the. . .populations that neg- populations large enough to maintain a balance between
atively impact fitness at a level comparable to complete mutation (which contributes new genetic material) and
inbreeding” (Severns 2013:23). Although perhaps less com- genetic drift (which results in fixation and loss of genetic
mon and less pervasive in its effects than inbreeding, the material), and such fragments might be too isolated to allow
5.6 Landscape Genetics and Habitat Fragmentation 189
sufficient gene flow to replenish lost alleles. This implies that, known. . . After sampling, genetic and statistical tools are
according to Honnay and Jacquemyn, “the effect of popula- used to determine the spatial genetic pattern and to correlate
tion size on genetic diversity is as pronounced in common it with landscape or environmental features” (Manel et al.
species as in rare species. This means that in our fragmented 2003:190). Broadly, landscape genetics examines genetic
landscapes, even common species may have reached a criti- distances between individuals relative to landscape structure,
cal threshold in population size and patch occupancy; thus, genetic subdivision of populations resulting from landscape
measures mitigating habitat fragmentation are strongly features, and how rates of gene flow are affected by landscape
needed” (Honnay and Jacquemyn 2007:829). Such effects characteristics (Holderegger and Wagner 2008) (Fig. 5.17).
would be especially strong in species that are obligate Through its integration of genetic and habitat data with
outcrossers (self-incompatible) because habitat fragmenta- spatial analysis tools, landscape genetics can enable conser-
tion that reduces population size would also reduce numbers vation biologists to better understand habitat features as dis-
of available mates, making even common species susceptible persal barriers. Let’s inspect one example.
to genetic loss (Morgan et al. 2013). The breeding habitat of the endangered golden-cheeked
warbler (Setophaga chrysoparia) (Fig. 5.18) is fragmented
5.6.2 Landscape-Induced Genetic

Differentiation
The interaction between loss of genetic diversity and habitat

fragmentation has given rise to a new branch of genetic study
known as landscape genetics, with diverse applications to
many practical problems in conservation. Landscape genetics
addresses the measurement of gene flow, with attendant
applications to conservation problems, using two approaches.
As Stéphanie Manel, one of the pioneers of landscape genet-
ics, and her colleagues, express it, “The two key steps of
landscape genetics are the detection of genetic discontinuities
and the correlation of those discontinuities with landscape
and environmental features, such as barriers. . .” (Manel et al.
2003:190). Manel and her colleagues explain further, “The
Fig. 5.18 The golden-cheeked warbler (Setophaga chrysoparia), an
identification of these spatial genetic patterns requires the endangered species and nesting habitat specialist breeding in Texas
collection of genetic data from many individuals (USA), whose populations are differentiated by landscape features.
(or populations) whose exact geographical location is (Photo courtesy of US Fish and Wildlife Service, public domain)
Fig. 5.17 A visual representation of approaches used in landscape side of the landscape feature. The assignment test identifies one individ-
genetics. In all three parts (a, b, and c) of the figure, the black line refers ual (circle with a solid outer line) as a recent immigrant from the other
to a specific landscape feature, such as a river, that might impede gene side of the landscape feature. (c) An investigative approach of current
flow. Filled circles signify adult organisms, and circles of different gene flow assessed by parentage analysis of offspring (squares). The
shades of gray signify genetically similar genotypes. (a) An investiga- hatched offspring has one parent on each side of the landscape feature.
tive approach based on genetic distances, where genetic distances (b) and (c) are both examples of investigative approaches to recent gene
among sampled individuals are determined and correlated with land- flow across a landscape feature, one of the major applications of land-
scape structure. In this case, the largest genetic distances are found on scape genetics. (Reprinted by permission from Oxford University Press,
opposite sides of the landscape feature impeding gene flow. (b) An from Holderegger, R., Wagner, H.H., 2008. Landscape Genetics. Bio-
investigative approach based on “assignment tests,” in which Science 58, 199–207. # 2008 American Institute of Biological
individuals are grouped into two pre-described populations on either Sciences)
throughout its breeding range in the state of Texas (USA). As 5.7 Managing Genetic Diversity in Wild
a nesting-habitat specialist, the warbler uses only dense, Populations
mature stands of Ashe juniper (Juniperus ashei) mixed with
deciduous trees (mostly oak (Quercus spp.) species). In their 5.7.1 Importing Genetic Diversity: Genetic
study of this species, biologist Denise Lindsay and her Restoration of Inbred Populations
colleagues predicted that “habitat fragmentation would lead
to reduced gene flow between populations, and, combined The common European adder (Vipera berus, Fig. 5.20) is, as
with decreased population sizes, would result in reduced its name implies, common. The IUCN Red List lists it in its
allelic diversity and heterozygosity within populations” category of “Least Concern,” small wonder given its wide-
(Lindsay et al. 2008:2123). They chose to assess gene flow spread Eurasian distribution (southwards from Scandinavia
using a genetic index, FST, commonly called the fixation and the Baltic region to France and the United Kingdom,
index, which reflects the proportion of genetic variance eastwards through most of Europe to Russia, Mongolia,
contained in the various subpopulations (the “S” subscript) People’s Democratic Republic of Korea and northwestern
relative to the genetic variance in the total population (the “T” China). Although widespread, populations are fragmented in
subscript) (Appendix 5.1). More specifically, FST is a mea- many areas, including southern Sweden. One isolated popula-
sure of the correlation of randomly chosen alleles from within tion near Smygehuk, Sweden, has long been confined to a
subpopulations relative to the entire population, which coastal strip of meadow 1 km long and 50–200 m wide which
provides a measure of the genetic variance. has been isolated from other adder populations for over a
There is more than one way to calculate the value of FST, century (Madsen et al. 1999). This population declined in the
but one is to use the equation 1960s and then began to exhibit a high proportion of deformed
and stillborn offspring coupled with low genetic variability,
FT FS both signs of inbreeding depression (Madsen et al. 1999).
F ST ¼
FT In 1992, Thomas Madsen of the University of Sydney
(Australia) and his colleagues attempted a “genetic rescue”
where FT is equal to the heterozygosity of the total population of this population, capturing 20 young adult male adders
and FS represents the heterozygosity of the subpopulations. from larger and more genetically variable populations and
To start with an extreme case, if FT and FS are equal (both releasing them into the Symgehuk population. Because male
contain equal genetic variation), the value of FST will be zero adders mature at about 4 years of age, Madsen and his
(there is complete genetic similarity between all colleagues did not expect immediate effect. But after only
subpopulations (i.e., no genetic differentiation between any 4 years they observed “dramatic increases in the number of
subpopulations)). In contrast, if the subpopulations each con- recently matured males recruited to [i.e., born into] the
tain much less genetic variability than represented in the total population. . . . genetic variability within the population also
population, the value of FST will approximate 1 (there is very increased rapidly [and]. . . recently recruited males exhibited
little genetic similarity between different subpopulations). much more genetic variability. . ., confirming that most of
Thus, a high FST implies a considerable degree of differenti- them were sired by the introduced males. The proportion of
ation between any given subpopulation and another subpop- stillborn offspring also fell suddenly, indicating that the rapid
ulation, whereas a low FST indicates little differentiation increase in recruitment was due to increased survival of juve-
between subpopulations. nile adders.” (Madsen et al. 1999:34) (Fig. 5.21).
Lindsay and her colleagues found associations between Judging this isolated population to be on the verge of
FST and landscape connectivity and between FST and geo- extinction prior to the introduction of the new males, Madsen
graphic distance for all warbler subpopulations (Fig. 5.19). and his colleagues concluded “Introducing new genes from a
The greater the distance and the less landscape connectivity different population enabled the adders to make a dramatic
(more nonhabitat) between subpopulations, the greater the recovery. This result encourages genetic approaches to con-
differences in the genetic composition of the subpopulations servation and supports the importance of preserving genetic
(higher FST, implying lower gene flow between variability as a way of increasing the viability of wild
subpopulations). Lindsay et al. concluded that “. . .a lack of populations” (Madsen et al. 1999:34–35). Several years
connectivity among habitat patches, resulting from loss of later, Madsen and his colleagues published another article
natural landscapes, is likely playing a primary role in the citing continued growth of the original population, having
notable genetic differentiation of at least one collected 39 adult male adders, more than any other time in
population. . .and is of concern for other populations as the 23-year study (Madsen et al. 2004). This ongoing popu-
well” (Lindsay et al. 2008:2130). lation increase highlights the value of introducing novel
5.7 Managing Genetic Diversity in Wild Populations 191
a
r = –0.40, P = 0.011
0.04
0.03
Pairwise FST
0.02
0.01
0.00
0 5 10 15
Percent connectivity
b
0.04 r = 0.40, P = 0.037
0.03
Pairwise FST
0.02
0.01
0.00
0 100 200 300 400
Geographical distance (km)
c r = 0.55, P = 0.008
0.04
0.03
Pairwise FST
0.02
0.01
0.00
0 20 40 60 80
Percent agriculture
Fig. 5.19 Associations (r) between pairwise estimates of gene diversity warblers perceive as non-habitat) between population subunits, the
(FST) in the golden-cheeked warbler (Setophaga chrysoparia) and greater the difference in genetic composition (higher FST)) between
(a) percent connectivity of populations, (b) geographic distance, and subunits. ((Reprinted by permission from Wiley, from Lindsay, D.L.,
(c) percent agricultural land between populations. The more connected Barr, K.R., Lance, R.F., Tweddale, S.A., Hayden, T.J., Leberg, P.L.,
population subunits are, the less they differ genetically from one another 2008. Habitat fragmentation and genetic diversity of an endangered,
(lower FST), suggesting higher levels of gene flow between population migratory songbird, the golden-cheeked warbler (Dendroica
subunits having high habitat connectivity. In contrast, the greater the chrysoparia): Genetic Diversity of Dendroica Chrysoparia. Molecular
distance and amount of agricultural land (which Golden-cheeked Ecology 17, 2122–2133. # 2008 Blackwell Publishing Ltd)
alleles for enhancing the viability of inbred populations. A

similar genetic restoration effort was conducted on a remnant
population of greater prairie chickens (Tympanuchus cupido
pinnatus) in Illinois (USA) with similar results (Hedrick and
Kalinowski 2000). Likewise, the Florida (USA) panther
(Puma concolor coryi) population, a subspecies of mountain
lion, was helped by introductions of individuals of the same
species (but not subspecies) from the US state of Texas in the
1990s when the Florida population was on the verge of
extinction. After the addition of Texas lions, Florida panther
numbers increased, genetic heterozygosity doubled, survival
and fitness measures improved, and signs of inbreeding
declined (Johnson et al. 2010).
Fig. 5.20 The Common European adder (Vipera berus) is widespread

throughout its Eurasian range, but individual populations can be subject Point of Engagement Question
to isolation and resulting inbreeding depression. One isolated and Although introduction of mountain lions from Texas
declining population in southern Sweden was restored through addition produced positive results for the Florida panther, did it
of more genetically diverse male adders from a larger population, an
example of “genetic rescue” in conservation. (Photo courtesy of R. “exterminate” the panther as a genetically unique sub-
Segers; reprinted by permission from Wiley, from Bauwens, D., species? Consideration of the potential loss of genetic
Claus, K., Mergeay, J., 2018. Genotyping validates photo- uniqueness is always essential in any effort at “genetic
identificationo by the head scale pattern in a large population of the
rescue,” especially when the rescue involves a small
European adder (Vipera berus). Ecology and Evolution 8, 2985–2992.
# 2018 The Authors) population that is genetically unique. We will explore
these questions further in our next section.
a 35 b
30
Total number captured
Number of recruits
25
Before After
Number of snakes
kb Introduction Introduction
20 9.4
6.6
Introduction of
new males 4.4
15
10 2.3
2.0
5
0
81 82 83 84 85 86 87 88 89 90 91 92 93 94 95 96 97 98 99
Year
Fig. 5.21 Change in number (a) and genetic diversity (b) of an isolated before introduction of new males (left) and in seven recruited males
population of European common adders in southern Sweden following sampled in 1999 (right). Note the changes in allelic frequencies of MHC
introduction of more genetically diverse adult males from a different, genes before and after introduction of new males as indicated by
larger population. (a) indicates total number and number of recruited changes in band numbers and locations. (Reprinted by permission
male adders captured in the isolated (Smygehuk, Sweden) population from Springer Nature, from Madsen, T., Shine, R., Olsson, M., Wittzell,
from 1981 to 1999, an index of population size; (b) represents a H., 1999. Restoration of an inbred adder population: Conservation
southern-blot analysis (a common genetic test) of major histocompati- biology. Nature 402, 34–35. # 1999 Macmillan Magazines Ltd)
bility complex (MHC) class I genes in seven male adders sampled
5.7.2 Hybridization and Introgression: The

Case of the Red Wolf Information Box 5.2: Real Life Applications
of Genetic Techniques in Conservation – The Case
Hybrids are offspring of matings between individuals of of the Red Wolf
different species, subspecies, or populations. In animals, Attempts to re-establish populations of the red wolf,
hybrids typically suffer a number of disadvantages compared which became extinct in the wild about 1975 (Wayne
to non-hybrids. Animal hybrids may be infertile, or, even if 1996), began when the US Fish and Wildlife Service
fertile, experience reduced mating success because (USFWS) released captive-bred red wolves on the
individuals in both their parental species may not recognize Alligator River National Wildlife Refuge, which
them as potential mates. These and related problems can encompasses about 154,000 acres on the Albemarle
make hybridization in animals a waste of reproductive effort Peninsula in northeastern North Carolina. Released
by potential parents, and wasted reproduction is not some- wolves became established on the refuge and were
thing small populations can afford. Hybridization can be designated as the Red Wolf Experimental Population
especially threatening to rare species due to reduced fitness (RWEP) – and the areas they occupied named the Red
of hybrids or through destruction of unique genotypes of rare Wolf Experimental Population Area (RWEPA). The
species (genetic assimilation). And some conservation legis- greatest threat to the RWEP was not from “simple
lation, such as the US Endangered Species Act (ESA), does hybrids” produced by pairings between red wolves
not protect hybrids. Thus, hybrid animals can suffer two and coyotes (Canis latrans), but from backcrossing
disadvantages: lower fitness and no legal protection. between red wolves and red wolf-coyote hybrids, a
When two closely related species coexist and one is rare, path of introgression that could eventually eliminate
the rare species may be genetically swamped – and its unique all remaining “pure” red wolves. Fredrickson and
genome exterminated – by interbreeding and hybridization Hedrick (2006) estimated the risk posed by these threats
with the more common species. A related problem is intro- by constructing a simulation model to predict future
gression, the acquisition and incorporation of genetic mate- trajectory of the red wolf population. In model outcomes,
rial from one species, subspecies, or population into the the number of red wolf pairs increased quickly to carry-
genome of another, a common result of hybridization. Intro- ing capacity when coyotes were not present. When
gression can make it difficult to identify, establish, and main- coyotes were present and pairing among red wolves
tain the genetic integrity of a species in relation to other, and red wolf-coyote hybrids was random, 80% of
closely related species. One of the best and most complex simulated red wolf populations were exterminated
examples of problems arising from inbreeding, hybridization, within 50 years (Fredrickson and Hedrick 2006).
and introgression involves the red wolf (Canis rufus) of the Based on model results, the most effective manage-
south-central United States (Fig. 5.22) (Information Box 5.2). ment technique was to sterilize paired coyotes and
paired hybrids, although this can be difficult when
appearance is the only criterion one can use in the
field. Nevertheless, a sterilization strategy was feasible,
and Fredrickson and Hedrick modeled outcomes of
three strategies: no sterilization, low sterilization
(50% of hybrid and coyote pairs are sterilized when
proportion of mixed pairs in the red wolf population
exceeds 0.40) and high sterilization (75% of hybrid and
coyote pairs sterilized when proportion of mixed pairs
exceeds 0.10). They combined these strategies with
three mating scenarios: random mating (red wolves,
coyotes, and hybrids select mates without discrimina-
tion), weak assortative mating (red wolves do not mate
with coyotes, but randomly choose between other red
wolves and hybrids), and weak assortative mating plus
“red wolf challenges.” Single red wolves and red wolf
pairs “challenge” and displace hybrid pairs and single
Fig. 5.22 The red wolf (Canis rufus), a species threatened with genetic
hybrids from their territories, reducing the proportion
extinction through hybridization with and introgression from gray of hybrids in the breeding population. Fredrickson and
wolves (Canis lupus), coyotes (Canis latrans), and dogs (Canis Hedrick developed algorithms to predict occurrence
familiaris). (Photo courtesy of B. Bartel, US Fish and Wildlife Service
and Point Defiance Zoo and Aquarium) (continued)

and outcomes of such challenges and fed the results (50 pairs) and persisted at that level for the length of the
back into the model so that challenges had an ongoing simulation (50 years). In this scenario, only 0.3% of
effect on population composition. Note the dramatic colonizing populations formed were predicted to
effect of a high sterilization effort on red wolf persis- become extinct after 50 years. At the other extreme of
tence, especially where red wolf challenges are no sterilization and random mating, the population
assumed (Information Box Fig. 5.1). Where a high declined to a low and highly vulnerable number of
sterilization effort was used as a management strategy animals (10 pairs) after 50 years. In this case, the
and red wolf challenges occurred at predicted rates, the model predicted extinction of 79.9% of colonizing
red wolf population rose quickly to carrying capacity populations (Fredrickson and Hedrick 2006).
Information Box Fig. 5.1 Mean numbers of red wolf pairs in Broken lines represent 95% confidence intervals. (Reprinted by
nonextirpated wolf populations over time when there is high or permission from Wiley, from Fredrickson, R.J., Hedrick, P.W.,
low sterilization effort of coyote and hybrid pairs with (a) random 2006. Dynamics of Hybridization and Introgression in Red Wolves
mating among red wolves, coyotes, and hybrids, (b) weak assorta- and Coyotes. Conservation Biology 20, 1272–1283. # 2006 Soci-
tive mating, and (c) red wolf challenges that displace hybrid pairs. ety for Conservation Biology)

From 2001 to 2013, findings associated with model coyotes were sterilized assuming that sterile individuals
results were integrated into a management plan which would serve as territorial placeholders, keeping unde-
divided the Albemarle Peninsula into three zones, each tected coyotes from dispersing into the peninsula (Bartel
with different management protocols (Information Box and Rabon 2013). In Zone 3, many sections were man-
Fig. 5.2). Zone 1 covered the core red wolf population. aged similarly to Zone 2, but in others, coyotes were not
Here coyotes and wolf-coyote hybrids were euthanized. euthanized or sterilized (Bohling and Waits 2015).
In Zone 2, hybrid individuals were euthanized, but
Information Box Fig. 5.2 Map of the Red Wolf Experimental Management practices in Zone 3 varied. Many sections of Zone
Population Area (RWEPA) and spatial distribution of hybrid and 3 were managed similarly to Zone 2, but in others coyotes were
red wolf litters in the Alligator River Wildlife Refuge on the not sterilized or euthanized. (Reprinted by permission from Bohling,
Albemarle Peninsula in North Carolina (USA), 2001–2013. Thick J.H., Waits, L.P., 2015. Factors influencing red wolf–coyote
black lines indicate boundaries of management zones. In Zone hybridization in eastern North Carolina, USA. Biological Conserva-
1, coyotes and hybrids were euthanized. In Zone 2, hybrid tion 184, 108–116. # 2015 Elsevier Ltd)
individuals were euthanized but coyotes were sterilized.
The genetic management of red wolf achieved some mea- recaptured and returned to refuge property. . . were unrealistic
surable success, but also experienced unforeseen problems. and scientifically unsound” (WMI 2014:4) and that the
Over the 13-year management period (2001–2013), more red USFWS should have had “greater oversight and support for
wolf litters were born (126) than hybrid litters (26), and most a landmark recovery program involving one of the most
hybridization events followed disruption of a stable pair of imperiled canids in the world” (WMI 2014:4). In its evalua-
red wolves due to the death of one or both of the adults. In tion of demographic data, the WMI concluded that the red
these cases, most deaths (69%) were the result of anthropo- wolf population would require higher survival or productivity
genic disturbance, mainly from gunshot wounds and traps, as than coyotes and coyote-wolf hybrids to persist. But, in fact,
the USFWS had allowed coyote hunting and trapping, as well local populations of red wolves, coyotes, and coyote-wolf
as deer hunting, in parts of the reintroduction area (Bohling hybrids had similar demographic rates. Based on this and
and Waits 2015). Faced with growing criticism of its restora- subsequent reviews, the USFWS determined that recovery of
tion effort, the USFWS contracted, in 2014, with the Wildlife the red wolf was feasible, but that it would “shift the Red
Management Institute (WMI) to conduct an independent Wolf Recovery Program’s focus and resources toward secur-
evaluation of its work. The WMI evaluation was critical of ing the species by fully supporting the captive population
the program, noting assumptions that “red wolves would stay . . .”, de-emphasizing efforts at increasing the RWEP in the
on refuge property or that they would be immediately field and designating it as a “non-essential” population
(USFWS 2017). Fredrickson protested these changes in the valuable and harvested populations whose wild stocks are
journal Conservation Letters, noting that “coyote demogra- declining. The detrimental effects on reproductive perfor-
phy had little effect on population outcomes suggesting that mance in the wild are increasingly apparent. For example,
red wolves do not need to demographically outperform lifetime reproductive success and overall fitness of steelhead
coyotes and hybrids to be successful” and that “assessing trout (Oncorhychus mykiss) declined precipitously in just two
the strength of reproductive barriers and addressing factors generations of domestication (Araki et al. 2007; Christie et al.
weakening these barriers should be a major focus of the 2012). Rearing in a hatchery environment reduced reproduc-
Service’s review” rather than demographic data (Fredrickson tive capabilities by ~40% per captive-reared generation when
2016:209). Today the red wolf’s future remains uncertain. fish moved to natural environments. As investigators bluntly
But despite the many problems and setbacks associated with put it, “Our data suggests a sharp decline in reproductive
this restoration effort, it has identified, developed, and, in success follows a very short time in captivity. . . These data
some cases, implemented management strategies that could fit very well on an exponentially declining curve. . .,
be used for mitigating effects of hybridization and introgres- [showing] 37.5% fitness decline per captive-reared genera-
sion in an endangered species. tion, suggesting that the fitness decline of captive-reared fish
can be remarkably fast” (Araki et al. 2007:102).
These studies demonstrate the dangers of genetic adapta-
5.8 Managing Genetic Diversity in Captive tion (GA) to captivity, which can be described mathemati-
Populations cally as
5.8.1 Measuring the Cost of Adaptation ~2 X

L
y y
Sh 1 L
to Captivity GA ¼ 1 ð1 m i Þ
L i¼1 ð2N e Þ
In his analysis of a failed attempt to establish a population of
where S is the selection differential in captivity (i.e., the
hybrid domestic X wild turkey (Meleagris gallopavo) in
strength of selection for captive traits) and h2 is heritability
Missouri (USA), Aldo Leopold attributed the failure to spe-
(additive genetic variation for reproductive fitness). y/L is the
cific differences in behavior between the hybrid turkeys
number of generations in captivity (expressed in years( y) per
(raised in captivity) and wild turkeys. He noted that domestic
generation length (L )). Ne is the (by now familiar) effective
turkeys were more docile, bred earlier in the season, and, as
population size, and mi is the proportion of genetic material
chicks, scattered when the hen sounded a note of alarm.
derived from immigrants (new animals) in the ith generation.
Chicks of wild turkeys hide. Commenting on this failure,
Even though at first glance an intimidating equation, we
Leopold remarked, “Wild turkeys are wary and shy, which
can begin to understand the relationship and its application
are advantageous characteristics in eluding natural and
by concentrating first on the efforts of a conservation biolo-
human enemies. They breed at a favorable season of the
gist working as the captive breeding manager. The founda-
year. The hens and young automatically react to danger in
tional goal is to reduce the value of GA to the lowest possible
ways that are self-protective. Reproductive success is
level. Look at the equation again and consider what the
high. . .Birds of the domestic strain, on the other hand, are
biologist could do to minimize the value of GA. The biologist
differently adapted. Many of their physiological reactions
could
and psychological characteristics are favorable to existence
in the barnyard but may preclude success in the wild”
1. reduce the amount of time in captivity (reduce the number
(Leopold 1944:192). Attempts at turkey reintroduction
using captive-raised turkeys were repeated widely throughout P
y=L
of iterations of )
the United States. With the exception of northern Michigan, i¼1
where a domestic-raised population did establish itself in the 2. reduce the strength of selection in captivity (the value of
wild, all efforts were failures. Turkeys were later S)
re-established throughout areas in the US where they had 3. equalize family sizes, so that selection occurs only within
been extirpated, but by using individuals from wild families (also reducing strength of selection in captivity)
populations translocated from other areas, not captive stock. 4. add more immigrants (unrelated individuals) from wild or
The problem Leopold described in turkeys is present in captive populations to slow the rate of genetic adaptation
every captive population. It is the problem of adaptation to
captivity. The more accustomed wild animals become to their Action 3, equalizing family size, should reduce selection
captive environment, the less likely they are to survive if for captivity (a way of implementing Action 2). At an exper-
returned to a wild one. The problem is prevalent today in imental level, Frankham (2005) found that equalizing family
efforts to use captive bred stock to supplement commercially sizes in fruit flies did yield the expected reduction in the
5.8 Managing Genetic Diversity in Captive Populations 197
strength of selection for captivity, but “produced little Management Plan (PMP) programs in North America, and
improvement in reintroduction success” and did not prevent their counterparts in Europe, the European Endangered Spe-
“substantial adverse effects upon reintroduction into the cies Program (EEP) and European Studbook (ESB), all of
‘wild’ from benign captive conditions. . .” (Frankham which are cooperative management programs in which zoos
2005:753). coordinate captive breeding efforts for hundreds of species.
Although it is increasingly difficult to implement Action Detailed information on each species is kept in databases
4 by capturing more wild individuals to augment captive called “studbooks,” which contain all known information
populations, “immigrants” to a particular captive population about each individual in the captive population, including
need not come from wild stocks. With well-designed captive its relationships to other captive individuals. Studbooks pro-
breeding management plans that involve the transfer of vide data for pedigree analysis, which enables managers to
animals from one zoo to another, animals from one region determine the kinship of individuals in a captive population.
can act as founders to other regions to which they are Informed by this knowledge of kinship, managers can evalu-
exchanged. Understanding the intention, methods, and ate the breeding priority of individuals and plan mating
scope of such management plans is essential if we want to strategies that minimize inbreeding and conserve genetic
grasp the way in which captive populations are integrated diversity. To see how such analyses can inform a breeding
into worldwide conservation strategies today. management plan, consider one species and its management
in captivity.
5.8.2 Managed Breeding: Mitigating Effects

of Inbreeding in Captivity 5.8.3 The Okapi: Analyzing Parameters
of Captive Breeding Management
Modern captive breeding efforts are no longer the haphazard,
often fortuitous adventure that was characteristic in times 5.8.3.1 History and Context: The Significance
past. They now are carefully planned, precisely executed of the Captive Okapi Population
international scientific initiatives using careful records of The okapi (Okapia johnstoni) could easily be the poster
past mating events and high levels of scientific technology species for global captive breeding efforts. Inhabiting only
and expertise for planned present and future breeding. In mature, tropical moist forests in Africa, all wild populations
2016, the Association of Zoos and Aquarium was overseeing of okapi today live within a single African nation, the Demo-
breeding and management plans for more than 450 species, cratic Republic of Congo (DRC, formerly Zaire). Although
and more continue to be added. Many captive management several large reserves were established in the DRC, including
programs are in place simply to supply individuals of partic- four World Heritage sites, enforcement of reserve boundaries
ular species to existing zoos and other facilities without and protection of species within the reserves has never been
further impact on wild populations. “Maintenance” programs secure. The reserves were vulnerable when they were
like these do not always involve endangered species and may established, and the situation grew worse with increasing
never have a role in the recovery of wild populations, but we human population growth and immigration of refugees into
cannot predict with certainty where, when, and in what reserves and surrounding areas (Wilkie and Finn 1990; Hart
populations recovery might someday be needed. and Hall 1996; Wilkie et al. 1997). Okapi populations
We have already learned (from turkeys and trout) that declined as a result of slash-and-burn cultivation and
behavior adaptive to captivity is detrimental to survival and subsequent forest succession associated with such human
reproductive success in the wild. Managers of populations activities (Wilkie and Finn 1990). To make matters worse,
likely to remain in captivity for a long time must also manage the government of the DRC was destabilized by civil war and
genetic considerations, especially the reduction of inbreeding incursions of neighboring countries which backed rebel
effects. Members of captive populations must retain as much forces in the 1990s. During that time, rebels held eastern
genetic diversity as possible from their wild-caught ancestors portions of the DRC, where the reserves are located. In the
to cope with natural (and unfamiliar) environments, and late 1990s, the headquarters of three of the four World Heri-
reduction of inbreeding depression is critical to maintain tage sites were looted. All technical equipment and vehicles
high survivorship and vitality among captive populations were stolen. Some park guards were killed (Hart and Hart
for as long as they may remain in captivity. 1997). In the face of such instability, conservation funding
Globally coordinated strategies of captive breeding may from western organizations declined (Hart et al. 1996). Open
lead to re-introduction in the wild at some future time to hostilities in the civil war came to an end in 2003 and the
supplement existing wild populations or establish new ones country has moved toward political stability, but civil unrest
(Hedrick and Miller 1992). This strategy has taken the form still exists and conservation efforts in the DRC remain uncer-
of the Species Survival Plan® (SSP) and Population tain, especially regarding the protection of existing parks and
Fig. 5.23 The female okapi “Oni,” the 50th calf born to the Antwerp planned and recorded for use in ongoing pedigree analysis of captive
Zoo in Belgium. The honor is fitting for the Antwerp Zoo as it was the populations. Studbook data are used to identify the breeding priority of
world’s first zoo to successfully maintain the okapi in captivity. Stud- each animal, minimize inbreeding, and reduce loss of genetic diversity
book data, such as those kept at the Antwerp Zoo, are used to develop in the managed captive population. (Photo courtesy of J. Verhulst,
pedigree analyses for worldwide captive breeding programs for many Antwerp Zoo, Belgium)
species. Today matings of animals in captive populations, are carefully
okapi populations within them. In 2012, an armed militia = Female

group opened fire on the headquarters of the Okapi Conser- = Male
vation Project, located in eastern DRC, killing six people and
13 of the 14 “ambassador” okapi that lived on public display
at the center (Platt 2012).
In contrast to the perils of life in the DRC, okapi have
thrived in captivity elsewhere, enjoying longevity and high
fecundity, and are now well represented in zoos around the
world. The okapi studbook database is kept at the Antwerp
(Belgium) Zoo, one of the world’s oldest and most famous
zoological gardens. It was the Antwerp Zoo where okapis
were first bred in captivity (Fig. 5.23). Data from the okapi
studbook have been used to develop a worldwide okapi
managed breeding program that identifies the breeding prior-
ity of each individual, minimizes inbreeding, and mitigates
loss of genetic diversity in the captive population. Let’s Fig. 5.24 A generalized example of a simple pedigree diagram.
examine how this can be accomplished. Squares represent males and circles represent females. In this example,
the parents (wild-caught founders) produce a male and a female off-
spring. The male offspring does not mate, but the female mates with an
5.8.3.2 Pedigree Analysis and Kinship unrelated male (avoiding inbreeding) to produce four offspring – two
Using studbook data, pedigree analysis can track males and two females (Reprinted by permission from McGraw-Hill
relationships among family members, both living and histor- Publishers)
ical. A pedigree follows the individuals (founders) that form a
new population and their descendants in each generation
through individuals of the most recent generation. A simple Knowing the ancestry of all individuals in a population
pedigree analysis might look like Fig. 5.24. allows managers to calculate kinship. As a result of the nature
If you wished, you could draw a pedigree for the okapi of Mendelian genetics and the independent assortment of
group shown in Table 5.5. chromosomes during meiosis, we can measure the kinship
You would use circles to represent females (dams) and between two individuals with simple probabilities. A kinship
squares to represent males (sires), and then connect related coefficient (kij) between two individuals (i and j) is the
individuals with lines as in the previous figure. Individuals probability that alleles randomly selected from homologous
with “WILD” parents are founders originating from wild or loci in two individuals are identical by descent from a com-
other unrelated populations. Founders are assumed to be mon ancestor.
unrelated to one another, and the same is assumed of wild To determine what these kinship probabilities are, start
parents unless data, such as molecular data or capture from the fact that a diploid organism inherits one-half of its
records, suggest otherwise, in which case known alleles from its mother and the other half from its father, so
relationships could be inserted into the “founder” matrix. that is has two alleles for each gene locus (Fig. 5.25, alleles B
and C in offspring). Thus, the probability of selecting one PðCÞdam PðDÞoffspring ¼ 0:50 0:50 ¼ 0:25:
particular allele (for example, B) of the two possible in any
diploid organism is 50% (0.50), the same odds as getting Thus, you can see that sibling-offspring kinship is nor-
either heads or tails if flipping a coin. If one were randomly mally a value of 0.25. Other common kinships in a typical
sampling alleles from a single gene of an individual and one pedigree are shown in Table 5.6.
of its parents, the probability of selecting a particular allele .
(for example, B) would be equal to the product of the two
individual probabilities (0.50 chance of choosing B from the 5.8.3.3 Population Mean Kinship
offspring 0.50 chance of choosing B from a parent ¼ 0.25). Once we determine an individual’s kinship to all other
This calculation reveals that there is a 25% probability that individuals in a population, a mean kinship for that animal
alleles selected from an offspring and its parent would be can be calculated. Mean kinship (mk) is the average of the
identical by descent from a common ancestor. Therefore, the kinships between a single individual and all other individuals
kinship between a parent and offspring is 0.25. in the population. Mathematically, it can be expressed as
In this example, the offspring share one allele of a given
gene with the sire and one allele with the dam. The kinship X
N
(k) of the sire with the offspring is equal to the probability of mk i ¼ k ij =N,
selecting the B allele in the sire times the probability of the j¼1
selecting the B allele in the offspring or

where mki is the mean kinship of the ith individual, kij is the
PðBÞsire PðBÞoffspring ¼ 0:50 0:50 ¼ 0:25: kinship of individual i to individual j, and N is the number of
individuals in the population. Thus, every individual (in this
Similarly, the kinship of the dam with offspring is equal to case, every okapi) would have its own mk and its own k to
every other individual. To extend the value of the original
pedigree analysis, the values of mk for every pairwise combi-
nation of individuals in the captive population could be
Table 5.5 A captive 10-member okapi (Okapia johnstoni) group recorded in a matrix configuration like the one in Table 5.7.
consisting of four wild-born founders (okapi 47, 71, 85, and 100) and Just as we can determine the mean kinship of individuals
their six descendants
to one another, we can also find the average kinship value of
ID Sex Dam Sire
71 Male Wild Wild
85 Female Wild Wild Table 5.6 Common kinships in a typical pedigree
47 Male Wild Wild
Relationship Kinship
100 Female Wild Wild
Parent – Offspring 0.25
160 Female 85 71
Sibling – Sibling 0.25
181 Male 85 71
Grandparent – Grandoffspring 0.125
196 Female 85 71
Half-sibling – Half-sibling 0.125
198 Male 100 47
Uncle/aunt – Nephew/niece 0.125
253 Female 160 198
Cousin – Cousin 0.0625
258 Male 196 198
Second cousin – Second cousin 0.03125
Data courtesy of K. Leus Unrelated 0
Such information can be used as a basis to begin a pedigree analysis of a
captive breeding group Table design by K. DeVoss
Fig. 5.25 Diagrammatic

representation of kinship
(k) between parents and their
offspring. Letters in boxes
represent alleles of each
individual (Reprinted by
permission of McGraw-Hill
Publishers)
Table 5.7 A matrix of kinship for each individual okapi to all other individual okapi in a breeding group
ID 47 100 71 85 160 181 196 198 253 258 mk
47 0.5000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.2500 0.1250 0.1250 0.1000
100 0.0000 0.5000 0.0000 0.0000 0.0000 0.0000 0.0000 0.2500 0.1250 0.1250 0.1000
71 0.0000 0.0000 0.5000 0.0000 0.2500 0.2500 0.2500 0.0000 0.1250 0.1250 0.1500
85 0.0000 0.0000 0.0000 0.5000 0.2500 0.2500 0.2500 0.0000 0.1250 0.1250 0.1500
160 0.0000 0.0000 0.2500 0.2500 0.5000 0.2500 0.2500 0.0000 0.2500 0.1250 0.1875
181 0.0000 0.0000 0.2500 0.2500 0.2500 0.5000 0.2500 0.0000 0.1250 0.1250 0.1750
196 0.0000 0.0000 0.2500 0.2500 0.2500 0.2500 0.5000 0.0000 0.1250 0.2500 0.1875
198 0.2500 0.2500 0.0000 0.0000 0.0000 0.0000 0.0000 0.5000 0.2500 0.2500 0.1500
253 0.1250 0.1250 0.1250 0.1250 0.2500 0.1250 0.1250 0.2500 0.5000 0.1875 0.1938
258 0.1250 0.1250 0.1250 0.1250 0.1250 0.1250 0.2500 0.2500 0.1875 0.5000 0.1938
Original matrix design by S. Long
Assume that founding individual 47, 71, 85, and 100 are unrelated to one another
the population, or the population mean kinship. Population 0% probability of having both of its alleles identical by
mean kinship is simply the arithmetic mean of all the indi- descent, so its inbreeding coefficient would be 0.
vidual mean kinships, as calculated in the previous expres- We can use the same data to determine another valuable
sion. An individual whose mean kinship is less than the measure of population genetics. That measure is called gene
population mean kinship has fewer relatives in the captive diversity.
population than an individual whose mean kinship is greater Gene diversity (GD) (not to be confused with the general
than the population mean kinship. concept of “genetic diversity”) is the most commonly used
The population mean kinship (MK) can be calculated as measure of the level of genetic variability in a captive popu-
lation, especially in zoos, by population managers. By defi-
X
MK ¼ mk i =N, nition, gene diversity is the probability that two alleles from
the same locus sampled at random from the population will
where mki is the mean kinship of the ith individual. not be identical by descent. That is, gene diversity is really
another name, and another way of looking at, a concept we
5.8.3.4 Kinship and Inbreeding introduced earlier, the concept of expected heterozygosity
Another genetic metric that can be calculated in a pedigree (He). However, if a population manager is armed with pedi-
analyses is an individual’s inbreeding coefficient (F), defined gree analysis data, she need not return to the theoretical
and calculated previously. Inbreeding, or the mating of calculation of He to determine a measure of gene diversity.
relatives, is a concern to genetic management because it The estimate is much easier because there is a simple, direct
reduces variation. Whereas managers of wild populations relationship between gene diversity (GD) and the value of
must be primarily concerned about threats associated with MK, previously calculated. That relationship is
environmental variation and catastrophes, mangers of captive
populations must focus on avoiding genetic deterioration of GD ¼ 1 MK:
the population, which can be aggravated by inbreeding.
Although we have previously introduced various If population mean kinship (MK) is high, then most
equations to calculate the value of the inbreeding coefficient individuals are related and gene diversity is low. Alterna-
F, we can now use an easier way, made possible by the data tively, if MK is low, the population contains a larger propor-
we have already analyzed in pedigree analysis and the relation of unrelated individuals and higher gene diversity.
tionship between kinship and inbreeding, namely an
individual’s inbreeding coefficient is equal to the kinship
between its parents. Thus, 5.8.4 Managing Captive Populations to Retain
Genetic Diversity
F ¼ k ij , where i ¼ sire and j ¼ dam:
5.8.4.1 Goals and Constraints in Captive Breeding
If an individual’s parents are completely unrelated, their Management
kinship to one another is equal to 0; they have 0% probability Managers of captive populations, no matter how dedicated,
of sharing any alleles that are identical by descent. Conse- experienced, and skilled, cannot maintain a population’s
quently, any individual offspring they produce would have a overall genetic diversity (for example, its allelic diversity)
indefinitely in a captive environment. Despite the inevitable without offspring do not pass on their genetic variation to
loss of overall genetic diversity, managers can, by focusing future generations. Managers can reduce loss of genetic diver-
on maintaining the value of a single genetic metric, the sity by increasing effective population size. If the ratio of Ne/
previously defined gene diversity (GD), maintain other N can be maximized, the influences of genetic drift and
kinds of genetic variation collaterally through their attempts inbreeding depression can be decreased. Methods to maxi-
to maintain GD. Specifically, managers of captive mize this ratio are to (1) grow the population to its captive
populations can manipulate population characteristics that carrying capacity as fast as possible, (2) maximize the number
influence the rate at which GD is lost or retained. Among of breeders in each subsequent generation, (3) equalize family
these characteristics are initial population size (number of sizes, (4) equalize the sex ratios of breeders and (5) reduce
founders), effective population size, target population size, fluctuations in population size. Such strategies, if successful,
population growth rate, and generation time. not only maximize the Ne/N ratio, but also retain existing
Founders – Initial population size, or number of founders, heterozygosity in the population.
can be used to estimate the gene diversity at the time of a A further influence on gene diversity is mean generation
population’s founding (GDt ¼ 0). We can express the rela- time, the average age at which animals produce offspring.
tionship as The number of generations occurring in a given number of
years is proportional to the amount of diversity that will be
GDt¼0 ¼ 1 ½1=ð2 N Þ, lost over that time. A population with a short generation time
will have more opportunities for loss of gene diversity during
(where N ¼ number of wild-born founders, assumed to have a given time period than a population with longer generation
unique (unrelated) alleles). If a manager is starting a captive time. Thus, managers can reduce rates of loss in gene diver-
population from scratch, the larger the number of founders, the sity by increasing the average age at which animals breed. In
greater the initial gene diversity. A manager must realize that practice, however, generation time is rarely manipulated in
there is a relationship of diminishing return in initial founder actual captive populations. Although theoretically a good
acquisition because, as a population is resampled, the addi- idea, managers usually consider the cost of losing animals
tional individuals collected becomes less likely to yield more to attrition prior to recruitment too high to merit delay of
new alleles. However, it still may be important to establish breeding. Additionally, increasing generation time can lead
strategies to acquire subsequent founders to supplement varia- to creating a selective environment for longevity, a further
tion lost to genetic drift in the captive population over time. adaptation to captivity that managers seek to minimize, and is
Effective population size – Recall that effective population likely inconsistent with conditions experienced by natural
size, Ne, is the size of an idealized, randomly mating popula- populations.
tion that is subject to the same degree of genetic drift as the None of the above considerations can yield a captive
actual population under consideration. The relationship breeding strategy that can be applied under all conditions.
between gene diversity at time t and effective population Comprehensive strategies can be developed, but they require
size is an integration of our knowledge of kinship and inbreeding
that focuses on managed matings, not managed population
GDt ¼ 1 1=ð2 N e Þ: size. Mating selection is a population process over which
managers of captive populations have the greatest control
Although effective population size has multiple (but related) because managers can determine which male and female
meanings, managers of captive populations usually refer to individuals will be permitted to mate with one another.
the “variance effective size,” an indicator of the number of Thus, current strategies for genetic management of captive
breeding individuals in a population. To estimate effective populations focus on the process of breeding pair selection.
population size, managers of captive populations use the sum
of the number of living males with living offspring and the 5.8.4.2 Avoidance of Inbreeding Strategies
number of living females with living offspring. The ratio of Remember that inbreeding, F, increases homozygosity in a
effective population size to census size (Ne/N) typically population and reduces mean heterozygosity (gene diversity,
ranges from 0.10–0.50 for intensively managed captive GD) such that
populations (Lacy 1995).
The effective size of a population influences the rate at F ¼ 1 GD
which gene diversity is lost. A population with a small effec-
tive population size relative to actual census population size Therefore, reducing inbreeding in a captive population
(Ne/N ratio) loses gene diversity more quickly than an equal- should increase gene diversity. Given this relationship,
sized population with a greater Ne/N ratio because individuals breeders have often approached the creation of breeding
groups with a maximum-avoidance-of-inbreeding (MAI) Remember the inverse relationship between mean kinship
strategy, which avoids matings between relatives. To imple- and gene diversity?
ment the MAI strategy, managers examine the kinship
between potential mates, which is equivalent to the 1 MK ¼ GD:
inbreeding coefficient of potential offspring of the pair.
Pairs with little or no kinship to each another (i.e., pairings If population mean kinship is minimized, gene diversity is
that would produce offspring with a low or zero inbreeding maximized. Minimizing population mean kinship (in other
coefficient) are then selected to breed. For example, if you, as words, lowering the average mean kinship of the population)
a population manager, had pedigree data from the okapi can be accomplished by breeding individuals with low mean
studbook for the population under your care, like that kinship values because these animals are the least related to
displayed in Table 5.7, you could create pairs with little or others in the population. Conversely, preventing the matings
no individual kinship to produce offspring with low of individuals with high mean kinship values has the same
inbreeding coefficients. But is this the best strategy available? effect. Recall that an individual animal’s mean kinship value
(mk) reflects its average relatedness to all other individuals in
5.8.4.3 Mean Kinship Breeding Strategies the captive population. An animal with a high mean kinship
A strategy to avoid inbreeding reduces deleterious effects of value is related to many individuals and thus carries many
inbreeding depression in a population, but geneticists have common alleles. But an animal with a low mean kinship
discovered that this strategy is still not the best approach for value has few relatives in the population and carries fewer
maintaining genetic diversity (Ballou and Lacy 1995). A common alleles. When choosing individuals for a breeding
more effective strategy is to select matings between captive pair, it is also important to match the mean kinships of the
animals that minimize population mean kinship (MK), rather individuals. Consider what happens when a breeding pair is
than minimizing individual kinship. Ballou and Lacy (1995) mismatched in their kinship. When a pairing is composed of
verified the efficacy of this strategy by comparing the individuals with disparate mean kinships (low and high), the
outcomes of a variety of mating designs through computer offspring will carry rare alleles from the low-MK parent and
simulation. They found that a strategy of minimizing popula- common alleles from the high-MK parent. In the future, when
tion mean kinship was best for maximizing the retention of that offspring matures and is itself chosen for breeding, its
genetic variation. Today the mean kinship (MK) strategy is offspring will carry both rare and common alleles. As a result,
the strategy of choice in managed captive populations. In a the rare and common alleles are perpetuated in equal fre-
MK strategy, individuals are paired based not only on their quency and representation is never equalized. On the other
relationships to one another, but also to the population as a hand, by breeding animals with low mean kinship, managers
whole. can increase the proportion of rare alleles in the captive
Most indices of diversity, such as the Shannon Index, population, and thereby raise the overall level of its gene
increase in value not only in response to the number of diversity.
species present but also in response to an increasing fre- Thus, an animal’s mean kinship value can serve as an
quency of rare species. Similarly, success of mean kinship index of its genetic value within the breeding program and
breeding strategy rests on the premise that gene diversity is a can be used to prioritize breedings for the purpose of
measure not only of the number of different alleles in the maintaining gene diversity. Assuming that the founding
captive population (which is a function of the gene diversity animals of a captive population are a representative sample
of the founders and cannot thereafter be altered), but also a of the wild population, an appropriate goal of managers is to
measure of the frequency of those different alleles preserve the genetic diversity present in the founders.
(Table 5.8), a variable which managers can manipulate Managers minimize changes from the starting gene pool by
through managed mating pair selections. equalizing founder contributions to future generations.
Individuals with low mean kinships represent founder
lineages that are under-represented in the general population.
Table 5.8 Two populations showing different frequencies of the same If these low mean kinship animals are allowed to reproduce,
four alleles (A, B, C, D)
their genetic contribution to the population will be increased
Population 1 25A 25B 25C 25D in the next generation. Likewise, by preventing over-
Population 2 97A 1B 1C 1D represented animals from reproducing, the contribution of
Based on data from S. Long, Association of Zoos and Aquariums. their common alleles can be decreased. By using mean kin-
Format design by M. J. Bigelow
In population 2, alleles B, C, and D are rare relative to allele A. Increased ship to equalize the contribution of the various founder
frequency of these alleles in population 1 gives population 1 a higher lineages in the population, managers can maintain and possi-
level of gene diversity bly even increase gene diversity in the next generation.
5.9 Applying Genetic Information in Conservation 203
5.9 Applying Genetic Information protection. Some endangered species might not be
in Conservation recognized (because they are morphologically similar to
common species) and allowed to become extinct. In a captive
5.9.1 General Considerations population, separate species mistakenly identified as the same
species might hybridize, resulting in lost genetic distinc-
The applications of genetic understanding and techniques in tiveness and reduced reproductive fitness. Until recently,
conservation that we have covered so far are varied, but can under the terms of the US ESA, protection could only be
be broadly grouped into six categories: (1) clarification of given to a distinct species, subspecies, or population seg-
relatedness, taxonomy, and phylogeny among populations; ment. This condition has been interpreted by US courts and
(2) determination of population management units based on agencies to mean that the protected group must have unique
genetic criteria; (3) estimation of gene flow and dispersal mitochondrial DNA sequences not shared with other groups,
among populations; (4) determination of the time since past and that there should be differences in allelic frequencies in
genetic bottlenecks in a population or the time since signifi- nuclear DNA loci, or evidence of genetically determined
cant differentiation between populations; (5) understanding morphological, behavioral, or life history difference
patterns of reproductive ecology; and (6) locating original (Frankham et al. 2002: 15). But, without proper genetic
sources of wildlife products, an important issue in the assessment, these questions cannot be answered. Thus,
enforcement of laws protecting wildlife from commercial genetic information can resolve taxonomic uncertainties and
exploitation. We have already seen examples of these define appropriate management units for conservation
applications in various ways, but now let’s make the (Frankham et al. 2002:365), especially through the aforemen-
applications and examples more clearly specific to these tioned technique of DNA barcoding, where a single gene,
conceptual categories to better grasp the full range of things the cytochrome oxidase I site gene, has been shown to con-
that genetic analysis can do in and for conservation efforts. sistently display species-specific sequences that permit pre-
cise identification and designation of different species of
animals.
5.9.2 Genetic Analysis Can Clarify Relatedness, An example of this application can be seen in recent
Taxonomy, and Phylogeny molecular genetic studies of sea turtles which compared the
Kemp’s ridley turtle (Lepidochelys kempii) and the similar
The clarification of relatedness and taxonomy has olive ridley turtle (L. olivacea). Molecular analysis
implications for one of the most basic questions in conserva- demonstrated that the Kemp’s ridley turtle deserved recogni-
tion biology: how many species are there? Thus, taxonomic tion as a separate species (Avise 1998). Similar applications
assessment is one of the most important applications of of molecular genetic techniques have shown genetic
conservation genetics. Until the 1970s, taxonomists classified differences in sympatric populations of killer whales
species, subspecies, and other taxonomic units using mor- (Orcinus orca) (Hoelzel and Dover 1991; Hoelzel et al.
phological characteristics of living or preserved specimens, 1998). The case of killer whales is particularly interesting,
often from very small samples. Systematic and taxonomic as genetic analysis demonstrated that observed differences in
uncertainties were large, but of limited interest outside pro- behavior in sympatric populations, so called “resource
fessional circles. As we have seen earlier, in the United polymorphisms,” could be genetically based (Hoelzel 1998).
States, all this changed with the passage of the Endangered
Species Act (ESA) of 1973. The Act extended legal protec-
tion to species and subspecies, making taxonomic determina- 5.9.3 Genetic Analysis Can Define
tion a matter of life and death, as well as the basis of legal Management Units of Fragmented
protection and the criterion for legal challenge. Geneticist Populations
Stephen J. O’Brien, reflecting on the “innocence” of taxo-
nomic science in the years before the US ESA, wrote, One of the fundamental questions of conservation biology is:
“When taxonomic distinctions became the basis for legal on what basis do we identify and delimit meaningful man-
protection afforded by the Endangered Species Act of 1973, agement units of populations, especially populations of con-
this innocence was lost forever. Disagreements over taxo- servation priority, in such a way that our management plans
nomic status fueled legal assaults on the Act, and misclassi- are sensitive and appropriate to real differences in population
fication led to inappropriate conservation measures resulting subunits? This is why the measurement of FST, previously
in losses of some species” (O’Brien et al. 1996). The rise of introduced in our study of landscape genetics (Sect. 5.6), is
taxonomy as a legal basis for conservation meant that taxo- such an important metric in many aspects of conservation.
nomic status had to be established with clarity, or endangered The genetic differences in subpopulations that can be
species might be denied protection under law. Alternatively, measured by FST can furnish an objective criterion for defin-
if the determination of taxonomic status was in error, effort ing population management units. We can appreciate its
might be wasted on abundant species that did not need value in the next two examples.
The nation of India harbors over half of the world’s highly structured, with large genetic differences between
population of the endangered Asian elephant (Elephas population segments. Differences were best explained by
maximus), but populations are fragmented and widely treating populations from the western Aleutians, central
separated. By analyzing mitochondrial DNA control region Aleutians, mainland Alaska and British Columbia, northern
sequences as well as six nuclear DNA microsatellite markers, California, and central California as five distinct genetic
conservation scientist T. N. C. Vidya and her colleagues management units. The authors asserted that “These
examined genetic structures of elephants from throughout populations are probably demographically independent and
the country, identifying four demographically autonomous non-exchangeable [and]. . .peripheral populations may be
population units, a northeastern India unit, a central India especially vulnerable to extinction. . .[given their] generally
unit, and two genetically distinct units in southern India, small size, and often marginal habitat” (Friesen et al.
Nilgiris and Anamalai-Periyar, that should be managed sepa- 2005:612).
rately based on their genetic uniqueness (Vidya et al. 2005).
Genetically identified units were different from government-
proposed management units, which had been influenced by 5.9.4 Genetic Analysis Can Determine Rates
political considerations. of Gene Flow Among Populations
Genetic techniques have been used in a similar way in
efforts to define appropriate population units for management Managing gene flow among populations, again most effec-
in the marbled murrelet (Brachyramphus marmoratus) tively assessed by measurement of FST, can help conservation
(Fig. 5.26), a seabird of the US and Canada that forages in biologists resolve questions of which animals to translocate
offshore ocean waters, but nests in large trees in coastal old to new sites, how often to do so, and when to begin and end
growth forests from central California to the Aleutian Islands. such translocations (Frankham et al. 2002:407). For example,
Marbled murrelets face a variety of threats in both foraging in Rocky Mountain bighorn sheep, Lukart and Allendorf
and nesting habitats, being vulnerable to deforestation and (1996) used mtDNA to infer rates of gene flow and genetic
forest fragmentation, oil spills, and gill netting. Murrelet differentiation in populations in the western United States,
populations have declined throughout their range and are and found a wide distribution of mtDNA groups (haplotypes)
listed as threatened species by both Canada and the over a large geographic region, suggesting that, in the past,
US. Victoria Friesen and her colleagues examined the gene flow among populations was high. However, many
sequence variations in 547 base pair fragments from control populations of bighorn sheep are now highly differentiated,
regions of murrelet mtDNA sampled from throughout their and some have become genetically fixed for a single haplo-
range. They found that genetics of murrelet populations were type, suggesting more recent isolation and fragmentation of
individual populations, a trend consistent with the results of
Epps et al.’s (2004) study of the effect of climate change on
these populations and the increasing fragmentation that
would result as more and more populations at lower, warmer,
and drier elevations disappear under current conditions of
global warming (Chap. 4).
Gene flow is also an important consideration in plant
conservation because it is often low in small populations of
rare and endemic plant species. Increasingly conservation
biologists have been able to assess gene flow indirectly by
examining distribution of genetic variation among
populations (Hamrick and Nason 2000). For example,
Dolan et al. (1999) determined such variation in three species
of rare perennial shrubs in Florida (USA) scrub vegetation.
The level of interpopulation gene flow was estimated by
calculating differences among populations. Although levels
of gene flow were different in each species, one species,
Fig. 5.26 The marbled murrelet (Brachyramphus marmoratus) of the Hypericum cumulicola, was experiencing almost no gene
Pacific coast of Canada and the United States, a threatened bird species flow among populations. This species had the lowest propor-
which forages in offshore waters but nests in large trees in old growth tion of species- and population-level polymorphic loci, the
forests. Genetic analysis of mitochondrial DNA in murrelets has enabled
conservation biologists to identify separate and genetically distinct fewest number of alleles per polymorphic locus, and the
population segments that can serve as the basis of population manage- lowest level of heterozygosity (Dolan et al. 1999). More
ment units and aid in the development of unit-specific recovery plans. recent studies (Oakley 2015) of the same species confirmed
(Photo courtesy of US Fish and Wildlife Service, public domain) that there has been limited genetic variation for ecologically
5.9 Applying Genetic Information in Conservation 205
important traits in these populations, which could limit adap- (Antarctic) stock of minke whales. Rather, it originated from
tive responses to future environmental change, possibly the so-called “J” stock, a depleted and protected population
increasing extinction risk. living in the northern Pacific, including the Sea of Japan.
In a population with these characteristics and little or no Taking the analysis further, Baker et al. (2000) examined
gene flow, conservationists must protect many individual genetic identifiers from minke whales known to have been
populations, not just a few in designated reserves, if they taken for scientific research and compared them to minke
intend to preserve genetic diversity in the species. And whale products sold in commercial markets. The market
because genetic diversity in this perennial shrub species is sample was significantly different in genetic characteristics
primarily found among populations rather than within from the scientific catch, indicating that most of these
populations, even small populations are worth preserving products did not come from scientific hunting, but from
(Dolan et al. 1999). undocumented, illegal exploitation of the protected J stock.
Using their knowledge of the market proportion of minke
whale products and the known scientific catch (from a differ-
5.9.5 Genetic Analysis Can Expose Exploitation ent stock of minke whales), Baker and his colleagues were
of Protected Species able to estimate that approximately 100–150 minke whales
were being taken each year from the J stock to supply
Following decades of over-exploitation of whaling stocks of observed proportions of it in Japanese and Korean markets.
all species worldwide, the International Whaling Commis- Taking their analysis to its conclusion in an insightful final
sion (IWC) voted, in 1982, to impose a moratorium on step, they used their data to model expected future trends in
commercial whaling. Taking effect in 1986, most whale the J stock of minke whales based on this level of annual take.
species were completely protected from commercial exploi- Their model (Fig. 5.27) indicated that, at these take levels, the
tation. Two exceptions were the northern Pacific and Antarc- “protected” J stock of minke whales would continue to
tic minke whales (Balaenoptera acutorostrata and B. decline (Baker et al. 2000).
bonaerensis, respectively), relatively small (25–30 ft)
baleen whales whose populations were somewhat less
depleted than larger species. Japan is one member-nation of
the IWC that was permitted a small annual take of minke
whales for “scientific” purposes. The nation of South Korea
also reported “incidental” take of minke whales in their
commercial fishing operations. Both of these sources of
minke whale “take” are legal under international law, and
the whale products can be sold in commercial markets in both
countries. To test whether the “whale” products in these
countries were derived from these sources, C. S. Baker and
his colleagues purchased products labeled as “whale” meat
from retail outlets from 1993 through 1999. Using a
phylogenic analysis of base sequences from the mtDNA
control region that could be compared to existing data on
various cetacean species, they determined that the whale
products included tissues and organs from eight species or
subspecies of baleen whales (including the two species of
Fig. 5.27 Estimated history and predicted future decline of the J-stock
minke whales), two species of sperm whales, two species of (northern Pacific/Sea of Japan) of minke whales (Balaenoptera
beaked whales, porpoises, killer whales, numerous species of acurostera) under three assumed levels of incidental take (50, 100, and
dolphins, as well as sheep and horses. Although genetic 150 whales per year) from 1998 onward. Incidental take levels represent
estimated take needed to account for past frequency of minke whale
analysis revealed that the majority of the “whale” products
products in Japanese and Korean markets (100–150 whales per year) and
(68%) in these markets were indeed from minke whales, the a conservative, lower estimate (50 whales per year) in case of over-
other six baleen species and the sperm whales represented in estimation or more effective enforcement of protective statutes. Note
the markets were protected by international law and had not that, regardless of level of incidental take, the J stock continues a steady
decline through 2050. (Reprinted by permission from the Royal Society
been hunted legally for years. Further analysis revealed that
(Great Britian), from Baker, C.S., 2000. Predicted decline of protected
the proportion of minke whale products present could not whales based on molecular genetic monitoring of Japanese and Korean
have been generated from scientific and incidental take alone, markets. Proceedings of the Royal Society of London B: Biological
and that much of this product did not come from the southern Sciences 267, 1191–1199. # 2000 The Royal Society)
Three important questions were answered in this investi- Discouraging as these findings are, their publication in this
gation. First, were whale products sold in legal commercial research effort and related studies exposed illegal activity and
markets derived from legal scientific take? No. Many whale contributed to increased international efforts to stop illegal
products represented unreported exploitation of endangered exploitation of protected stocks and species, as well as con-
whale species, as well as products not from whales at all. sumer fraud. Such exposure provided the basis for more
Second, were the products from scientific take (minke vigorous enforcement by wildlife enforcement agencies, as
whales) from abundant (Antarctic) stocks? No. Six times well as incentive for heightened awareness among consumers
more of the minke whale products were from the protected who did not want to be party to illegal exploitation of
J stock than from the Antarctic stock. Finally, what propor- endangered animals. More recently, researchers have
tion of undocumented whale products in these markets came continued highlighting ongoing efforts to establish a DNA
from “protected” species and populations? Sadly, approxi- register, or a tissue bank system for biological material, for all
mately 10% of the products came from protected species and minke whale bycatch. A comparison of market data to regis-
31% from protected stocks of minke whales (Baker et al. ter samples could help determine whether the whales are
2000). legally bycaught or illegally caught (Song 2016).
Synthesis primary concerns in most population management efforts

Advances in genetic technology and assessment have for either common or rare species. The best available
given conservation genetics an increasing role in the man- models for population growth and viability analysis,
agement of captive populations, conservation forensics, such as VORTEX, RAMAS, ZooRisk, and GAPPS,
taxonomic clarification, and conservation law. Increasing increasingly incorporate capacities for genetic manage-
precision and quantification of the genetic characteristics ment of populations, and modelers now build custom
of individuals and populations have given added force, models for specific populations that incorporate a full
with occasional dilemmas and confusion, to landmark range of genetic effects. In the field, however, manage-
legislation like the Convention on the International ment of populations usually still underestimates and
Trade in Endangered Species (CITES), the Convention under-incorporates genetic considerations. Creating a
on Biological Diversity, and the US Endangered Species larger role for genetic considerations in management of
Act and its counterparts in countries throughout the world, wild populations will require conservation biologists to:
as well as many other conservation laws that require
taxonomic clarity to be applicable to the protection of 1. Intentionally build genetic considerations into long-
species. And in cases where taxonomic distinctness is an term management strategies;
important criterion for setting conservation priorities, 2. Extrapolate appropriate strategies for most taxa from
genetic analyses decisively answer questions of taxo- relatively few studies of model species, at least for the
nomic status (Schemske et al. 1994). Despite these foreseeable future;
developments, genetic considerations still often play rela- 3. Identify more precisely under what circumstances
tively little part in actual management of in situ genetic considerations could limit the size or distribu-
populations. But they should play a far greater role, tion of managed species, and
because we now know that loss of individuals in a popu- 4. Use genetic data more often, more systematically, and
lation, even in common species, results in loss of genetic more intentionally to better understand population and
diversity in that population. In logged tropical forests, for extinction processes.
example, geneticists Wickneswari Ratnam and Timothy
Boyle determined that logging, which reduced average An understanding of genetic conservation begins to reveal
basal area of trees by 56%, also reduced expected hetero- to us exactly what genetic diversity is, why inbreeding can
zygosity and genetic diversity (Shannon Index) by lead to a population’s decline, and why processes of gene
5–23.4% in five species, each with a different life history flow matter to conservation efforts, not only for
strategy, and loss of species-specific alleles ranged from populations per se, but with regard to questions of habitat
7.7 to 25.0% (Ratnam and Boyle 2000). fragmentation, dispersive ability, and population struc-
Ratnam and Boyle’s study is but one of many examples ture. These considerations are becoming more common.
that precisely quantify how much genetic diversity may be When they are normative, “ordinary,” and part of standard
lost even in “ordinary” use and management of biological of “best practices” in all biological studies and manage-
resources. Yet genetic considerations are still often not ment, all efforts in biodiversity conservation will benefit.
(continued)
Appendix 5.1: A Summary of Mathematical Equations Used in Common Estimations of Genetic

Diversity and Effective Population Size
Symbol Definition Equation Notes

P Polymorphism P ¼ number of polymorphic loci
total number of loci
Expressed as a probability of encountering a polymorphic locus within all
loci in the population.
P
H Average heterozygosity Hi Where H is the average heterozygosity at locus i and N is the total number of
H¼ N loci used.

1 t
Ht Change in H t ¼ H 0 1 2N Where H0 represents the original level of heterozygosity and H1 represents
heterozygosity by the new level of heterozygosity after any number of t generations.
generation

ΔH Change in ΔH ¼ H m 2N
1 Where m is the addition of heterozygosity through mutation.
heterozygosity by
mutation
H Heterozygosity H ¼ 2Nm Occurs when ΔH ¼ 0.
equilibrium
P
A Allelic diversity ðA1 þA2 ...þAn Þ Where A1 is the number of alleles at locus 1, A2 the number of alleles at locus
A¼ N 2, and so on through all N loci.
Δq Genetic drift Δq ¼ qð2N
1qÞ Where q is a randomly selected allele in one generation, and Ne is the
e
variance effective population size.
Ne Effective population size N e ¼ 2þσ
4N
2 Where N is the population’s size and σ 2 is the variance in progeny number.
4N N
Effective population size N e ¼ N m þN
m f
f
Where Nm is the number of males and Nf is the number of females.
with unequal sex ratios
Effective population size N e ¼ P tð 1 Þ Where Nt is the effect population size at generation I, and Nei is the effect
N i
by generation e population size in generation i.
E(n) Expected number of E(n) ¼ m ∑ (1 pi)2Ne Where m is the number of alleles prior to the bottleneck, p is the frequency
alleles after a bottleneck of the fth allele, and N is the effective number of individuals at the
bottleneck.
FST Fixation index F ST ¼ FTFF
T
S Where FT is the expected heterozygosities of the total population and FS is
the expected heterozygosities of the subpopulations
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The Conservation of Populations: Theory, Analysis,
Application 6
I have seen something else under the sun: the race is not to the swift, or the battle to the strong, nor does food
come to the wise, . . . but time and chance overtake them all.
Ecclesiastes 9:11
Keywords 6.1 Defining Populations and Population

Population demography · Stochasticity · Density depen- Processes
dence · Metapopulations · Occupancy estimation · Mini-
mum viable populations · Population recovery · 6.1.1 What Is a Population?
Population viability analysis · Conservation-reliant spe-
cies · Conservation management agreements The traditional definition of population is “all coexisting
individuals of the same species living in the same area at
the same time.” Populations are composed of individuals and
change through time because these individuals die, give birth,
Overview immigrate (into the population) or emigrate (to leave and join
In this chapter, you will learn: a different population). We attempt to understand populations
by our measurement and prediction of the probabilities of
1. What populations are and what determines their size these and other population events (the study of population
and persistence. demography) and the factors that influence these
2. What metapopulations are. probabilities. Some influences come from characteristics
3. How populations can be detected and measured. and behaviors of the population itself (intrinsic factors) and
4. What is a “minimum viable population” (MVP). some from environmental or ecological events and processes
5. What is population viability analysis (PVA). external to the population (extrinsic factors). Populations can
6. How PVA results can be used in conservation be differentiated by genetics, demography (distinguishing
management. patterns of rates of birth and death) and spatial distribution
7. What is a conservation-reliant species. (the area over which they occur). Differences in spatial dis-
8. What are practical steps for determining how to tribution are the most important, because, if the populations
conserve populations. are not separated spatially, differences in genetics and
demography will not exist.
The concept and definition of “population” can also be
made more complex in some situations. This is often the case
where populations are threatened, harvested, or are otherwise
controversial in regard to public perception. In such cases,

212 6 The Conservation of Populations: Theory, Analysis, Application
populations may be defined politically rather than biologi-

cally because politics, not science, is often the determining Information Box 6.1 (continued)
factor in management decisions and jurisdiction. For exam- explanation of what you study, you must start to study
ple, in the United States, the northern Great Lakes states of the inconvenient. In plants, for example, dispersing
Wisconsin and Michigan both independently manage their individuals (transients) contribute nearly 50% to
wolf (Canis lupus) “population” even though the two states patterns of population change across multiple
share a common boundary (which is meaningless to wolves). lifeforms, and the effects of dispersers on population
Artificial as political definitions may be, they must be con- growth increases with increases in the number of plant
sidered just as carefully as spatial, demographic, and genetic life stages (McDonald et al. 2016). Population connec-
considerations in real conservation management plans. tivity, vital for gene flow in general and
metapopulation dynamics in particular, is strongly
affected by a species’ dispersal ability (Kool et al.
6.1.2 Population Demography 2013). As we shall see later in the case history of the
growling grass frog (Sect. 6.2.3), knowing a species’
Extinction, the quality of “ceasing to be,” marks the termina- average dispersal distance enables managers to deter-
tion of a population’s existence. Extinction is an event that mine which currently unoccupied habitat subunits can
conservation biologists seek to avoid, but few populations be reached by dispersers and which cannot. In the past,
suddenly disappear when they are large and growing. Extinc- dispersal was measured indirectly by measurements of
tion is preceded by decline. genetic similarity, or directly by visual observation or
Population growth is defined by birth, death, immigration, mark and recapture studies. Modern technologies,
and emigration. In a group of individuals born at the same including telemetry (using both surface and satellite
time (a “cohort”), the sum of the probabilities of survival of monitoring systems), global positioning system (GPS)
each individual to a particular age (survivorship) influences sensors implanted on individuals, and radar which can
the trajectory of population change through time. Individuals track large groups or airborne individuals (such as
are added by births, a process known as recruitment. Recruit- insects or birds) have made direct measurement of
ment is the net additions/subtractions of reproductively via- dispersal more accessible and less mysterious (Kool
ble individuals, added to the population after adjustment for et al. 2013). Advances in occupancy modeling and
early or neonatal mortality. As such, recruitment is strongly estimation of detection probabilities (Sect. 6.3) have
influenced by fecundity, the number of young or eggs pro- potential to make indirect measurements of dispersal
duced per female (animals) or seeds per individual (plants) more precise. Unfortunately, recent reviews of the lit-
per unit time. Other increases accrue to the population erature in conservation biology still find that dispersal
through immigration, while additional losses are incurred is most commonly measured in indirect ways, includ-
through emigration. Both are functions of dispersal, the ing expert opinion, modeling, and (poorly collected)
permanent movement of an organism from its area of birth occupancy data, all leading to a multiplication of low
to a new area. Dispersal must be measured in terms of rate quality information about dispersal. More than half of
(proportion of individuals that leave the natal area), distance 655 publications recently reviewed in conservation
(how far an organism travels from the natal area before it literature described measurement of dispersal as an
resumes a settled existence) and direction (which way is the area of “knowledge gap” (Driscoll et al. 2014). Yet
dispersing individual going, especially in relation to its natal without accurate, high quality information about dis-
area, other populations, suitable habitat, or human habitation persal, it will be impossible to fully understand pro-
or disturbance). All three provide essential information in cesses affecting target populations or to achieve
population management and restoration. (Information Box successful outcomes in population restoration. Dis-
6.1). persal is a subject where there remains a vital need
for continued investigation and improved quality of
research. If you enter the field of conservation biology,
Information Box 6.1: Dispersal – A Neglected Ele-
perhaps you can make a contribution.
ment in Population Conservation
What is hard to measure is convenient to ignore. This
has long been the case of the dispersal in the study of
plant and animal populations. But when what you are If we had perfect knowledge of the exact state of each
ignoring is important to the understanding and individual in a population, we could perfectly predict the
behavior of the population through time. But we don’t.
(continued) Lacking perfect knowledge, we construct models based on
6.1 Defining Populations and Population Processes 213
our assumptions about a population. We begin with very

simple models and then add (and later, sometimes subtract)
complexity as we learn, from observing real populations,
what is actually needed to make accurate predictions of
population behavior. Models are ways of understanding the
complexity of populations, but the models and their
assumptions are not laws (like the “laws” of physics or
chemistry) that invariably and inviolably describe how
populations always behave. Models must be informed by
study of real populations, and critically evaluated according
to how well they describe real population behavior.
Simple models of population growth integrate
complexities of these multiple factors with relatively few
Fig. 6.1 The exponential growth curve, a graphical depiction of a
mathematical concepts. The simplest model of population population increasing at an ever increasing rate over time. (Illustration
dynamics can be understood as the sum of birth (B) and by M. J. Bigelow)
ingress (I) minus mortality or death (D) and egress (E).
Sometimes referred to as the “fundamental” equation of has an ecology. An accurate model must reflect that. The
population dynamics, it can be expressed mathematically as simplest model of population growth affected by the environ-
ment is logistic growth, which includes an environmental
N ðtþ1Þ ¼ N ðtÞ þ ðBt þ I t Þ ðDt þ E t Þ limit (K, carrying capacity) on population size that slows
population growth as N approaches this limit. Logistic
where the size of the population at the next time increment growth is defined as
(Nt + 1) is a function of its present size (Nt) plus additions of

birth and ingress (immigration) and subtractions of death and dN KN
¼ rN :
egress (emigration). dt K
If a population is growing exponentially (increasing at an
increasing rate over time), then its growth is defined by the When N is small relative to K, KN K approaches unity, and
population’s size, N, and rate of increase, r, which is the the population grows at a nearly exponential rate. But as
difference between its rate of birth and rate of death. In a N approaches K, the value of the expression approaches
closed population (no immigration or emigration), the num- zero, and so does the rate of growth, dN dt . Thus, a visual
ber of births (B) is a function of the per capita birth rate projection of the model depicts a population growing through
(b) times the population size (B(t) ¼ bN(t)). Similarly, the time at a nearly exponential rate during early stages of
number of deaths (D) is a function of the per capita death rate growth, but gradually slowing until it reaches a stable equi-
(D(t) ¼ d(N(t)). We drop the subscripted t on b and d because librium (Fig. 6.2).
we assume they are constant through time. With this under- Notice, in this equation, that changes in the size of the
standing, one of the most common formulations of popula- population (N ) directly affect population growth, an expres-
tion growth rate is then r ¼ b-d, where r is the per capita or sion of density dependence. Animal population demographer
instantaneous rate of growth. In exponential growth, change Marlené Gamelon and her colleagues express this concept of
in numbers (dN) in the population over change in time (dt) is density dependence precisely. “Density dependence operates
expressed as through a negative feedback between the growth rate and the
population size at one or more steps. . .In many cases, density
dN dependence results in regulation of the population
¼ rN:
dt fluctuations around a mean population size, the carrying
capacity. . .” (Gamelon et al. 2016). Density dependence is
Viewed graphically, exponential growth is a J-shaped curve in fact the reason that logistic growth occurs, because it is the
showing a population growing over time at an ever- governor of the interplay between different and competing
increasing rate (Fig. 6.1). sources of mortality. At low population densities, different
Exponential growth can be a helpful starting point for kinds of competing forms of mortality (for example, disease,
understanding the foundational process of population predation, starvation) may not be not strongly interactive.
increase – the size of the population times its rate of growth. Therefore, different kinds of mortality are additive in their
But exponential growth is not a realistic, long-term phenom- effects. As the population’s density increases, density depen-
enon in any population. As biologist Mark Boyce remarked dent mechanisms increasingly couple competing causes of
bluntly, “it has no ecology” (Boyce 1992). A real population
stochasticity, demographic stochasticity, environmental

stochasticity, and natural catastrophes (Shaffer 1981).
Their effects are not predetermined, but subject to uncertainty
(stochastisities) whose probabilities are unique to particular
populations and their environments. If a population is large
relative to the magnitude of uncertainty, the outcomes
associated with these sources of uncertainty (stochastic vari-
ation) are predictable using an average of projected
outcomes. But if the population is small relative to the mag-
nitude of uncertainty, its success or failure might be unpre-
dictable because of chance events affecting only a few
individuals and the disproportionate effects these events
might have on the population.
The heath hen (Tympanuchus cupido cupido), a bird simi-
Fig. 6.2 The logistic growth curve, a graphical depiction of a
population’s growth as it approaches an environmental limit or carrying lar in appearance and behavior to the North American prairie
capacity. (Illustration by K. DeVoss) chicken (Tympanuchus spp.), was once common throughout
the northeastern US. By 1876, overhunting and habitat
destruction had restricted its range to the island of Martha’s
mortality into compensatory relationships – relationships in Vineyard in Massachusetts (Shaffer 1981). By 1900 there
which mechanisms like the behavior of individual members were fewer than 100 birds, and a refuge was established for
of the population (intrinsic factors) limit population growth them on the island in 1907. By 1916, the population had
before the population begins to be limited by its environment increased to around 800 individuals and seemed to be headed
(extrinsic factors). for recovery. Then a series of unfortunate environmental and
The logistic model assumes that all individuals in a popu- demographic events befell the survivors. In the year of their
lation are demographic equivalents of one another in repro- peak population, a fire (a stochastic event) devastated the
duction and survivorship. This is rarely so, especially in island and destroyed most of the remaining habitat and
populations with long-lived individuals. In these cases, pop- nests. A high winter concentration of goshawks (Accipiter
ulation growth is often defined and determined by age struc- gentilis) (another stochastic event), a predator of heath hens,
ture and sex ratio. Age structure and sex ratio are important deepened the decline. After a minimal recovery in 1920,
variables in population dynamics. But to understand their disease (a third stochastic event) swept through the popula-
roles more precisely, we must first distinguish between the tion, eliminating all but 100 birds. Losses continued from this
two categories of factors that influence population size and point on, with increasing numbers of birds experiencing
trajectory – density–dependent factors and density–indepen- sterility. The proportion of males increased until, in the
dent factors. final years, there were no females at all! The population
was extinct by 1932.
Factors of environmental stochasticity, demographic
6.1.3 Stochastic Perturbations – stochasticity, genetic stochasticity, and natural catastrophe –
Density-Independent Factors the “Four Horsemen of the Extinction Apocalypse” – all
of Population Growth contributed to the heath hen’s demise. These factors acted
and interacted on the population independent of its size and
6.1.3.1 Stochastic Factors density. But when the size and density of this population
Many factors that influence population size are described as were reduced, such factors overwhelmed remaining
stochastic factors whose values vary according to a individuals and led to the heath hen’s extinction. Stochastic
randomly-generated probability distribution. A population factors that act in density independent ways are complex and
whose growth is solely a function of chance events, or sto- interactive, but we will examine each one individually.
chastic variation, can be described as being density indepen-
dent. That is, factors and forces which act on increasing or 6.1.3.2 Genetic, Environmental Stochasticity
decreasing the size of the population act independently of its and Demographic Stochasticity
density (the number of individuals per unit area). What are Small populations suffer increased inbreeding, genetic drift,
these “stochastic factors”? and accumulation of unfavorable mutations (Chap. 5). Many
In his classic paper, “Minimum Population Sizes for Spe- small populations in the wild, such as the Florida panther
cies Conservation,” biologist Mark Shaffer identified four (Puma concolor coryii) (O’Brien et al. 1996), as well as
“sources of uncertainty” affecting population size: genetic captive-bred populations of endangered species, show
Fig. 6.3 Effects of

environmental and demographic
stochasticity on persistence time
of a population. Note that while
probability of extinction from
such forces is low in large
populations, these factors create a
high probability of extinction at
low populations. More
information in text (After
Simberloff 1998)
detrimental influences associated with these effects, which Demographic stochasticity refers to random fluctuations
can lead to further population declines and eventual in birth and death rates, emigration and immigration, or sex
extinction. ratio and age structure of a population. Biologist Robert Lacy
Environmental stochasticity refers to fluctuations in the noted that “with the exception of aging, almost all events in
probability of birth and death due to temporal variation of the life of an organism are stochastic” (Lacy 1993). In large
habitat parameters; populations of competing, parasitic, or populations, variation among individuals rarely matters; in
predatory species; and incidence of disease. The importance small populations, it matters a lot. Loss of a pregnant female,
of environmental stochasticity can be best understood rela- a skewed sex ratio, or accidental deaths of a few breeding
tive to the average rate of increase of the population. Let rav adults can significantly affect small populations. Like envi-
represent that average rate and Ve represent the variance in ronmental stochasticity, effects of demographic stochasticity
population growth attributable to environmental variation. If diminish with increasing population size, and population
rav is greater than Ve, then expected persistence time of a persistence is all but assured if r (the population’s intrinsic
population increases directly with increasing population size rate of increase) is positive (Fig. 6.2). In small populations,
at an ever-increasing rate (Fig. 6.3). In this scenario, environ- effects of demographic stochasticity make extinction a real
mental stochasticity is unlikely to cause extinction as long as risk.
the population is not very small. On the other hand, if Ve is
greater than rav, the shape of the population persistence curve 6.1.3.3 Natural Catastrophes
is different. Persistence time still increases as the size of the Some (for example, Simberloff 1998) have argued that natu-
population increases, but it reaches an upper asymptote, ral catastrophes are just special cases of environmental
beyond which further increases in population size do not stochasticity because they are often extreme forms of normal
significantly increase expected time of population persis- environmental variation (e.g., prolonged drought or flash
tence. This second case describes a population with large, floods resulting from intense, heavy rain). But catastrophes
environmentally induced population fluctuations and a rela- not only lie outside the normal probability distribution of
tively small rate of increase. In such a case, even a large random events associated with environmental variation they
population would be vulnerable to extinction. The best rem- may be qualitatively as well as quantitatively different in their
edy would not necessarily be to generate the largest popula- effects. In the words of biologist Robert Lacy, “a forest fire is
tion, but to ensure that the total population did not all not just a very hot day” (Lacy 1993). Catastrophes pose
experience the same environmental variations at once, threats for small populations because they can eliminate
which is also the best remedy for reducing risk of extinction every individual. The most viable protection against risks
from natural catastrophes. This is a problem we will examine associated with catastrophes is spatial segregation of different
shortly. population subunits. You can see an example of how to
reduce such risks in Information Box 6.2.
Information Box 6.2: Avoiding Extinction from Information Box 6.2 (continued)
Natural Catastrophe: Reducing Risk Through Spa- times as high for the single large population as for the
tial Segregation separated populations.
What are the relative risks of keeping a small population This kind of analysis has been used in the past on
intact on a single site versus subdividing the population actual endangered species, such as the whooping crane
into separate units on separate sites, the better to avoid (Grus americana) to determine whether it was better to
the tragedy of all individuals being exterminated by a manage the species as a single large population or as
natural catastrophe in one place? Using a method known two smaller populations (Information Box Fig. 6.2)
as Decision Analysis (DA), we can approach this prob- (Maguire 1986). In this case, the greatest environmen-
lem analytically and display our choices and their tal risk was considered to be that of a severe storm
probabilities as a decision tree (Information Box striking the population, particularly on its historic win-
Fig. 6.1), evaluating all possible outcomes in this exam- tering grounds on the Texas Gulf Coast (USA). The
ple. If the population remains as a single unit and there is single decision criterion is to minimize extinction risk.
no fire, the extinction probability is 0.05. If a fire occurs, Ideally, DA would be coupled in making management
their extinction is certain (pE ¼ 1.0). Their expected decisions to a Population Viability Analysis (PVA), a
probability of extinction is then the sum of the tool we will work with in detail later in this chapter
probabilities of these events, or (Sect. 6.5). Without the added detail and sophistication
(0.05 0.9) + (0.1 1) ¼ 0.045 + 0.1 ¼ 0.145. If the of a PVA, our DA is obviously one dimensional. In the
population is managed as two units, there are four whooping crane decision tree, the only serious threat
possibilities: (1) a fire occurs in both units; (2) a fire considered is environmental catastrophe (a violent
does not occur in either unit; (3) and (4) a fire occurs in storm), but other threats could also cause extinction in
one unit but not in the other. The probability of a fire in this population. Our DA also assumes that natural
both units is 0.10 0.10 ¼ 0.01; and the probability that catastrophes are independent events in the two areas
a fire does not occur in either unit is 0.9 0.9 ¼ 0.81. (they may not be if the two areas are not sufficiently far
The probability of a fire in at least one unit but not in apart). Informed by a carefully constructed PVA, our
both is 0.1 + 0.1 ¼ 0.2. Therefore, the extinction proba- DA would be more complex, but also more realistic,
bility of the two subunits is (0.01 1) + 2 and our management decision would lead to better
(0.040.15)+(0.810.0225)¼0.01+0.012+0.018225¼ management.
0.040225. In this case, extinction risk is more than three
Information Box Fig. 6.1 A decision tree of risk analysis for the viability analysis, risk analysis can provide valuable insight into the
probabilities of extinction for a population managed under two different relative risks of different management strategies for a population with
management strategies. When combined with results of a population given characteristics. (Illustration by K. DeVoss)
Another is that each new individual has the same effect on

Information Box 6.2 (continued) slowing or “braking” the growth of the population as the last
individual, such that all individuals contribute equally to this
negative feedback between population growth and popula-
tion density. The equation also assumes the effect is instanta-
neous, without any “lag” in time between the addition of the
new individual and the effect of the addition on population
growth. And it assumes that all individuals (members of N )
are equally adept at getting resources from the environment,
K, and that K itself remains constant and is unaffected by
members of the population or by other factors. These
assumptions are almost always false in real populations and
would not help a conservation biologist predict real popula-
tion change. What can be done to understand and model
density-dependence in a more realistic way?
6.1.4.2 The “Instantaneous” Assumption –

Detecting Time Lag in Density Dependence
Over 60 years ago, in their monumental work, the book The
Information Box Fig. 6.2 A decision tree representing probabilities of Distribution and Abundance of Animals, published in 1954,
extinction of different sized whooping crane (Grus americana) population ecologists H. G. Andrewartha and L. C. Birch
populations under different management scenarios relative to the risk
of potential elimination by severe storms. (Reprinted by permission asserted that most animal populations, especially populations
from Springer Nature, from Drechsler, M., Burgman, M.A., 2004. of invertebrates, fluctuated randomly in response to environ-
Combining Population Viability Analysis with Decision Analysis. Bio- mental variation – that is, changes in their populations were
diversity and Conservation 13, 115–139. # 2004 Kluwer Academic mostly density independent (Andrewartha and Birch 1954).
Publishers)
This was sometimes described as the “fencepost model”
because it assumed that population size was limited by the
total amount of some essential resource and a sharp, defin-
able, and often sudden “cutoff” occurred above this level (the
These and other forms of stochastic factors are not directly
“fencepost”), with no effect below this level (e.g.,
influenced by members of the population. They can be referred
Charlesworth 1972).
to as mechanisms of population control because they may limit
An alternative view of population variation, density
the size of the population. But some factors that influence
dependent population regulation, was being articulated at
population size and density are intrinsically affected by the
about the same time, first by the Dutch ornithologist Huyb
population itself, especially by the density of the population.
Kluyver through his long-term studies and experimental
Many, some would say most, natural populations are not only
manipulations of populations of a common European forest
controlled by density-independent factors, but also regulated
bird, the great tit (Parus major) (Kluyver 1951), then more
(kept within a predictable range of variation) by density depen-
comprehensively expressed in the classic work of British
dent factors. What are examples of density dependent factors
ornithologist David Lack in his book, The Natural Regula-
and how do they affect populations?
tion of Animal Numbers (Lack 1954). Density dependent
regulation in populations of the great tit (Fig. 6.4) was
6.1.4 Density Dependent Population inferred by Kluyver, Lack, and others from observations of
Regulation negative correlation between population growth and popula-
tion size. Using data compiled through annual monitoring of
6.1.4.1 Density Dependence and Logistic Growth a population of great tits using a small forest called Marley
We have seen how density independent factors can exercise a Wood near Oxford University in England, Lack
measure of control over a population’s growth, often keeping demonstrated that, over time, great tits achieved relative
it below some particular size. But density dependent factors constancy in numbers through density dependent regulation –
affect a population more precisely, and so are often referred manifested in, among other ways, inverse relationships
to as factors of regulation that not only limit a population’s between number of chicks raised in relation to the number
overall size or density, but keep its average size and density, of breeding pairs (Fig. 6.5) and number of breeding pairs in a
over time, near a long term average. How does this happen? given year compared to change in population size relative to
As an expression of density dependence, the logistic the previous year (Lack 1964).
growth equation makes several assumptions. One is that While the studies of Kluyver, Lack and others had already
individuals are added to the population at a constant rate. begun to suggest that additions of new individuals might not
assumption in his classic examination of populations of forest

insects. He noted that most past analyses (including those of
Andrewartha and Birch) of population growth in insects
“look only for direct (not-lagged) density dependence.
Thus, there is a real danger that populations characterized
by delays in regulation will be relegated to a density-inde-
pendent limbo by an analysis not-equipped to recognize such
behavior” (Turchin 1990:660). Turchin attacked the problem
by changing the rules – creating approaches, models and
equations that related populations in 1 year to populations
in preceding years (technically, an approach called “autocor-
relation,” examining how strongly population size in a given
year was correlated to population size in previous years).
Searching for signs of delayed density dependence, Turchin
Fig. 6.4 The great tit (Parus major), a common Eurasian songbird found that eight of 14 insect populations he examined did
whose populations were the subject of foundational research leading to exhibit delayed density dependent regulation. “Delayed den-
an understanding of density dependent population regulation in animal sity dependence,” noted Tuchin, “can arise in natural
populations. (Photo courtesy of F. Franklin, reprinted under CC3.0 populations as a result of interactions with other members
BY-SA)
of the community, such as natural enemies, or because high
population densities may adversely affect the fecundity of the
next generation. . .an analysis that does not consider time lags
will not. . .detect density-dependent regulation in population
oscillations driven by delayed density-dependent
mechanisms” (Turchin 1990:662–663).
6.1.4.3 The Assumption of Constant Carrying

Capacity (K)
Traditional logistic growth assumes that carrying capacity
(the maximum size a population can attain in a given envi-
ronment) is constant, when in reality it changes according to
environmental variation and interaction with the effects of the
population on available resources. It is more accurate to
define carrying capacity (K ) not as a constant value a popu-
lation cannot exceed, but as a population size reflecting
equilibrium between a population and its resources created
by their dynamic interaction. Thus, the value of K varies
environmentally. For example, herbivore populations can
lower the value of K by damaging their plant food resources
in ways that reduce the plants’ long term productivity. Con-
versely, changes in plant abundance, distribution, and
nutritional quality can affect demographic processes in the
population that influence the strength of density dependence
(Wang et al. 2006; Riotte-Lambert et al. 2017).
Interactions between density dependence and environ-
mental variation can be even more complex. For example,
Fig. 6.5 Production of young in relation to density of breeding pairs in a Wang et al. (2006) modeled long-term population trends in
population of great tit (Parus major) from 1947–1960 in Marley Wood near
Oxford, England, UK, one of the earliest demonstrations of density depen-
bison (Bison bison) in Yellowstone National Park (USA) and
dent population regulation. (Reprinted by permission from Wiley, from four western US populations of elk (Cervus elaphus) using
Lack 1964. A Long-Term Study of the Great Tit (Parus major). Journal of long-term time series analyses of population size. Their
Animal Ecology 33, 159–173. # 1964 British Ecologial Society) models revealed that elements of both temporal and spatial
variation affected strength of density dependence. The
have instantaneous effect on population growth, as would be greater the variation in one aspect of temporal variation,
assumed by the logistic equation, entomologist Peter Turchin winter temperatures, the greater the strength of density
later made one of the most direct refutations of this dependence in reducing population growth. The greater the
Fig. 6.6 Effects of spatial 0.6

heterogeneity in (a) vegetation a
(Normalized Vegetation Index 0.5 r2 = 0.51
(NDVI)) and (b) temporal wi = 0.92
variability (based on coefficient of 0.4
variation (cv)) in winter
Model-averaged strength of density dependance

temperatures on strength of 0.3
density dependence on one
population of bison (Bison bison) 0.2
in Yellowstone National Park,
USA and four populations of elk 0.1
(Cervus elaphus) in two national
parks, the National Elk Refuge, 0.0
and the Gravelly Mountains in the
western United States. r2
represents amount of variation in 0.30 0.32 0.34 0.36
strength of density dependence cv of NDVI
explained by each variable.
(Reprinted by permission from 0.6
Wiley, from Wang, G., Hobbs, N. b r2 = 0.70
T., Boone, R.B., Illius, A.W., 0.5
wi = 0.79
Gordon, I.J., Gross, J.E., Hamlin,
K.L., 2006. Spatial and temporal 0.4
variability modify density
dependence in populations of 0.3
large herbivores. Ecology
87, 95–102. # 2006 Ecological 0.2
Society of America)
0.1
0.0
0.0 0.1 0.2 0.3 0.4

cv of winter temperature
variability of vegetation in the environment (an example of animals to remember how resources are distributed in their
spatial variability), the weaker the effects of density depen- local environment, is a common, in fact, normal feature in the
dence on population regulation (Fig. 6.6). Regarding effects behavior of most animals, and, as more recent research is
of winter temperatures (temporal variability), Wang et al. beginning to reveal, a powerful interactant with and on den-
explained that “. . .increasing temporal variability in sity dependent processes in animal populations.
resources that determine carrying capacity also amplifies
density–dependent feedbacks to population growth, causing 6.1.4.4 The Assumption of Equal Contribution: The
reduced population density and equilibrium densities relative Concept of the Critical Age Class
to the deterministic case. . .It is plausible, in turn, that these As previously noted, traditional expressions of density
reductions in carrying capacity amplify the intensity of feed- dependence, like the logistic growth equation, assume each
back from population density to individual survival and on individual makes an equal contribution to density-dependent
reproduction” (Wang et al. 2006:99–100). Regarding effects effects, regardless of age, sex, or reproductive status. In real
of spatial heterogeneity (spatial variability) expressed in var- populations, this is rarely the case. Over 40 years ago, popu-
iation in vegetation, Wang et al. noted that “. . .spatial hetero- lation geneticist Brian Charlesworth demonstrated that den-
geneity appeared to weaken the effect of density dependence. sity dependence in populations could be, in fact, an
We propose that this effect is mediated by the ability of age-dependent process. Charlesworth explained “. . .this den-
herbivores to exploit spatial heterogeneity by selective sity [dependent] regulation must occur in response to the
feeding. . .[which] allows herbivores to cope with food number of individuals in a specific group of ages, what I
resources containing concentrations of nutrients that vary shall call the ‘critical age group.’ This critical age group
over time” (Wang et al. 2006:100). might, for example, be composed of all individuals of repro-
What Wang et al. are demonstrating in this study is an ductive age, or of newborn individuals, depending on the
example of diet selectivity, which is manifested in animals, in biology of the population” (Charlesworth 1973:306–307).
part, as habitat selectivity, something we will examine in the The value of Charlesworth’s work in this and other aspects
next chapter. Such selectivity, which involves and requires of population genetics and evolutionary biology was widely
recognized, but the implications of his insights about density segregation through management are often difficult and costly.
dependent processes of population regulation being driven by But, in natural environments, many populations are spatially
one or more “critical age groups” has been one that conser- segregated and widely dispersed whether managed or not.
vation biologists and population managers have not always When such segregation is present, gene flow via exchange of
appreciated or applied. More recent studies have begun to individuals must occur to retain connectedness between
examine the effects of age-structure on density dependence subunits. In such cases we must modify our understanding of
more closely. For example, Gamelon et al. (2016) studied “population” to take these conditions into account, and so
how density dependence affected population growth rate in begin to understand the concepts, functions, and processes
age-specific population sizes in the great tit, the same bird associated with metapopulations.
species studied by Huyb Kluyver and David Lack. Working In their previously mentioned work, The Distribution and
with a 38 year data set, Gamelon et al. demonstrated that the Abundance of Animals, ecologists H. G. Andrewartha and
strength of density dependent processes was not independent L. C. Birch asserted that “a natural population occupying any
of sex or age, but concentrated in one sex-age class, females considerable area will be made up of a number of. . .local
in “age class 1” (females in their first year of breeding). populations” (Andrewartha and Birch 1954). Accompanying
Gamelon et al. determined that “. . . the number of females their description, Andrewartha and Birch provided a schematic
in age class 1. . .strongly affected survival and recruitment of illustration of a series of spatially subdivided populations of a
all age classes and also the population growth rate. . .These species with different densities in each subunit. This seemingly
findings question the assumption that all individuals can be innocuous statement and simple illustration generated little
treated as having an equal contribution to density regulation attention, but together represent one of the first expressions
and that the effect of the number of individuals is age inde- of the concept of a metapopulation – a population existing as
pendent” (Gamelon et al. 2016:2486). spatially disjunct subunits at different densities in habitat
Determining overall population growth is what integrates, patches of varying carrying capacities. Andrewartha and
and therefore simplifies, the enormous complexities of the age- Birch further noted that individual subunits suffered periodic
and sex-specific mortalities (and the factors that influence extinction, followed by recolonization by individuals dispers-
them) into a single measure of fitness, the population’s rate of ing from neighboring subunits. These conditions are also part
growth, r (as it has been previously represented in the logistic of our understanding of metapopulations today.
growth equation) or λ (lambda (λ ¼ er), as it will be subse- The view of populations as subdivided by interacting units
quently represented in our study of population viability analy- was made more explicit by biologist C. B. Huffaker and his
sis). Much of the work of conservation biology is focused on co-workers in an elegant series of experiments involving
understanding factors causing mortality in populations, espe- mites and oranges in the late 1950s and early 1960s (Huffaker
cially novel, unexpected, or human-induced mortalities like 1958; Huffaker et al. 1963). Huffaker’s work, designed to
hunting and harvest, interactions with endangered species, evaluate dynamics of predator-prey relationships, used the
climate change, or emergent diseases. That is why understand- six-spotted mite (Eotetranychus sexmaculatus) as the prey
ing density dependence matters. For example, in species species, and another species of mite (Typhlodromus
harvested commercially, trapped, or hunted for sport, the occidentalis) as the predator. Both mites can sustain large
assumptions and form of density dependent regulation populations on the skin of an orange, so Huffaker created
operating in a population will affect the estimate of the sustain- “habitats” of oranges and “non-habitats” of rubber balls
able rate of harvest. For threatened or endangered species, the placed in various combinations on a tray. Mites could not
assumptions and form of density dependence can affect or even leave the tray but could move among the balls and oranges
determine persistence time for that population. (Fig. 6.7). When the prey species was forced to feed in
habitats (oranges) concentrated in large areas and grouped
at adjacent positions, predators exterminated prey within
6.2 Populations and Metapopulations: 2 weeks, whereupon all predatory mites starved. When
Complexities of Population Subdivision oranges were dispersed, prey survived longer, and predator
and Fragmentation and prey populations followed regular cycles of increase and
decrease (Fig. 6.8) (Huffaker et al. 1963).
6.2.1 Origins of Metapopulation Theory In these experiments, Huffaker and his colleagues
demonstrated the importance of environmental heterogeneity
We noted in our examination of the role of natural catastrophes in maintaining the stable predator-prey interactions. They
on populations that one of the most viable means of protection also demonstrated that populations can persist as
for small populations against such catastrophes, as with less “subpopulations” that occupy fragmented habitats on a tem-
severe forms of environmental stochasticity, is spatial segre- porary basis, with individuals moving regularly from one
gation of population subunits. Attempts to achieve such habitat subunit to another. Individual subpopulations suffer
6.2 Populations and Metapopulations: Complexities of Population Subdivision and. . . 221

representation of Huffaker’s
experiment (Huffaker 1958;
Huffaker et al. 1963) on
persistence of a predator-prey
system of two species of mite.
Dark circles represent oranges
mites could colonize. White
circles represent rubber balls they
could not colonize. (Illustration by
K. DeVoss)
Fig. 6.8 Oscillations in densities of the predatory mite, Typhlodromus prey populations follow a series of regular synchronized fluctuations.
occidentalis, and its prey, the six-spotted mite, Eotetranychus (Reprinted by permission from the University of Californa,
sexmaculatus, in Huffaker’s experimental system of oranges (habitat from Huffaker, C.B., 1958. Experimental studies on predation: disper-
for Eotetranychus sexmaculatus which feeds on oranges) and rubber sion factors and predator-prey oscillations. Hilgardia
balls (non-habitat) over a period of 60 weeks. Note that predator and 27, 343–383. # 1958 Regents of the University of California)
extinction, and only a portion of all available habitats are survive for a while, send out migrants, and eventually disap-
occupied at any one time, yet the population persists. pear” (Levins 1970:82) (Fig. 6.9).
Levins’ was a mathematical ecologist whose theory of
metapopulations arose from his examinations of habitat het-
6.2.2 The Definition and Development erogeneity and its relationship to problems associated with
of Metapopulation Concepts the control of insects that damaged crops. In a paper
presented at the symposium “Genetics in Biological Control”
Huffaker created fragmented habitats by random placement at the 1968 Meeting of the Entomological Society of Amer-
of oranges and rubber balls. The process of fragmenting ica, Levins stated that his purpose was “to show that the
habitats similarly divides formerly contiguous populations pattern of environmental variation in space and time can be
into spatially discrete population subunits. The concept of utilized in the control of pests and to indicate the information
metapopulations was developed to describe such conditions which is needed for the selection of the most promising
as an alternative to the traditional view of populations as predator” (Levins 1969). Levins’ objective was to determine
demographically homogeneous units having no group- the optimum properties of predator populations that could
specific structure. The metapopulation alternative emerged control pests, and then produce such a population through
as an explicit model in the late 1960s when Richard Levins genotypic selection. His long-term goal was to direct
(1970) offered the first definition of a metapopulation, a word entomologists away from thinking about “average”
he coined to describe “any real population [that] is a popula- conditions and focus instead on using specialized predators
tion of local populations which are established by colonists, that would not be uniformly effective in all environments
conservationists perceived that population persistence also

would depend on the ability of individuals to disperse suc-
cessfully among habitats. Metapopulation theory suggested
that vacant habitats might be recolonized on a regular basis
and unoccupied habitat could be as important as occupied
habitat in long-term population persistence. It also indicated
that a fragmented group of population subunits could,
because of its spatial structure, actually enhance other aspects
of population structure as well as population persistence,
giving conservationists a rationale for valuing fragmented
populations and fragmented habitats (Simberloff 1997). As
a result of these implications, spatial structure became a key
concept of metapopulation theory and modeling (Hanski and
Simberloff 1997).
Further efforts in modeling and field studies led to creation
Fig. 6.9 The Levins’ Model of Metapopulations. Levins’ of new, alternative population concepts, or re-use, with mod-
metapopulation model portrays extinction and migration patterns of
ification, of old ones. So-called “patchy-populations”
individuals living in discrete subunits. Without recolonization of the
habitat, each local population is in danger of becoming extinct. Arrows (Harrison 1991) were conceived as panmictic interbreeding
represent population recolonization. White patches represent populations occupying discrete habitat units, but in which
populations that have become extinct due to the lack of immigrants. exchange of individuals between units was so frequent that
Gray patches are occupied habitats. (Illustration by M. J. Bigelow,
demographic events were synchronous and population
modified by K. DeVoss, based on concepts from Harrison 1991)
subunits had no unique genetic structure. In this arrangement,
changes in habitat use, not changes in rates of colonization
(Levins 1969). Levins offered his theory to exterminate and extinction, are primary drivers of population size and
populations. Ironically, his view of metapopulations would distribution. In contrast, “source-sink” populations (Pulliam
come to be employed in conservation biology as a way to 1988) were understood as those in which a large and more
preserve them. stable “source” population, with positive growth and a sur-
The original Levins concept was a spatially implicit model plus of recruitment, supplies new individuals to smaller, more
with discrete habitat patches and local populations, all extinction-prone “sink” populations with little or no recruit-
assumed to be equally connected to one another (Hanski ment. Yet another variant, “mainland-island” populations,
and Simberloff 1997). Spatially implicit models, in their were originally proposed in the older theory of island bioge-
elegance and simplicity, facilitated mathematical and concep- ography (MacArthur and Wilson 1967), and constituted a
tual analyses of how metapopulations might work. Unfortu- special case of source-sink populations in which a single,
nately, spatially implicit models were unrealistic, and their large, and persistent “mainland” population provided
dependence on other assumptions about populations limited immigrants to surrounding “island” populations, limiting
questions that could be asked. their rates of extinction. The Florida scrub jay (Aphelocoma
Subsequent efforts in modeling metapopulations relied on coerulescens), for example, once considered a classic exam-
spatially explicit models, which assumed differing degrees of ple of metapopulation structure, has, upon more detailed
connectedness between population subunits and featured study, revealed a variety of population arrangements (Stith
“localized interactions.” Localized interactions are those in et al. 1996). Some have corresponded to a classical (Levins)
which population subunits interact primarily or exclusively metapopulation model, others to the “patchy-population”
with neighboring subunits, not all subunits. A further refine- model (Harrison 1991), and still others to the mainland-island
ment in metapopulation modeling has been the development model or variations of it (Stith et al. 1996).
of “spatially realistic models” (Hanski and Simberloff 1997) These and other variants of metapopulation theory and
that include considerations of the geometry of particular models relaxed Levins’ original assumptions about what
patches (especially on issues of size, shape, and arrangement constituted a metapopulation. But relaxed assumptions can
of patches). Metapopulation theory’s view of populations as lead to incorrect judgments, which in turn lead to misguided
spatially discrete subunits in fragmented, yet still connected, management recommendations. In a more recent examina-
habitats offered a picture of what biologists perceived to be tion of metapopulation concepts and their use, Emmanuel
the case in nature. Biologists realized that any plan for Fronhoffer of the University of Wartburg (Germany) and
maintaining extant populations would have to incorporate his colleagues defined four conditions required to meet the
preservation of many habitat fragments rather than relying criteria of a classical metapopulation. First, discrete habitat
exclusively on large, contiguous habitat blocks. Further, patches where individuals occur must be able to support
6.2 Populations and Metapopulations: Complexities of Population Subdivision and. . . 223
breeding populations. Second, any of these breeding

assemblages (subpopulations) must be prone to extinction.
Third, recolonization of any habitat patch containing, or
formerly containing, a population subunit must be possible.
Fourth, subpopulation dynamics must be asynchronous
(demographic events like population increases and decreases
must occur at different times in different units) (Fronhoffer
et al. 2012).
These conditions are rarely met in real populations
because only specific environmental conditions and life-
history attributes promote emergence of classical
metapopulation structures. Many instances in which target
species are described as “metapopulations” are really cases of
spatially structured populations (different population
subunits having different characteristics), but not cases
where the population actually meets the four criteria. This
can lead to serious errors in management prescriptions. For
example, emphasizing connectivity between habitat patches
Fig. 6.10 The growling grass frog (Litoria raniformis), a species native
would enhance persistence of a true metapopulation (because to southeastern Australia endangered by encroachment from develop-
it would increase colonization rates of individual patches), ment associated with the city of Melbourne. (Photo courtesy of
but would be wasted on a mainland-island or source-sink Museums Victoria and David Paul)
population where the primary concern should be protecting
the mainland or source group. In conservation, like medicine, because extinct habitats may be recolonized by individuals
the right prescription requires the right diagnosis. Carefully from neighboring populations, (5) the probability of habitat
applied, conservationists can use metapopulation theory to colonization will be strongly influenced by the proximity of
accurately understand characteristics of endangered neighboring subunit populations (Heard et al. 2012).
populations, and from such understanding, make intelligent Surveying 167 wetlands 1380 times over a 7-year period
recommendations that can prevent their extinction. We can from 2001 to 2007, as well as conducting a mark and recap-
learn from one study that made such application. ture effort that provided estimates of rates and distances of
dispersal, Heard et al. confirmed each of the five predictions
for this population. Occupancy of surveyed wetlands by the
6.2.3 Does Metapopulation Theory Predict growling grass frog was not random, but spatially clustered,
Behavior of Real Populations? The Case confirming Prediction 1 (Fig. 6.11). Most frogs showed dis-
of the Growling Grass Frog persal distances of less than 200 m, supporting the prediction
of strong site fidelity in individual populations (Prediction 2)
One of the most rigorous examinations of the match between (Fig. 6.12) Repeated surveys confirmed frequent site-specific
theory and observation in metapopulation study was colonization and extinction (Prediction 3) (Fig. 6.13). The
undertaken by Geoffrey Heard and his colleagues in their influence of proximity on extinction was weak (i.e.,
investigation of an endangered amphibian species, the growl- extinctions were not strongly clustered, Prediction 4)
ing grass frog (Litoria raniformis) (Fig. 6.10), native to (Fig. 6.13), but colonization sites were strongly clustered,
southeastern Australia and currently threatened by the expan- with all colonized wetlands lying within 1300 m of an
sion of the city of Melbourne. Heard and his colleagues occupied wetland (average distance 510 m, Prediction 5)
framed five predictions derived from classical (Fig. 6.13).
metapopulation theory and tested them against observed For the growling grass frog, predictions of classical
characteristics of the frog population around Melbourne. metapopulation theory proved true, and offered a sound
The predictions were: (1) probability of habitat (wetland) basis for conservation planning. Such implications in this
occupancy is positively influenced by proximity of neighbor- case would include management recommendations for pre-
ing populations; (2) habitat fidelity will be high, such that the serving presently unoccupied wetlands adjacent to existing
majority of individuals will occupy only one wetland during population subunits; increasing efforts in wetland enhance-
their lifetime; (3) habitat occupancy will be highly dynamic ment and creation at specific locations to increase
over time because populations are independent and metapopulation size, diversity, and connectivity; and
extinction-prone, and, therefore (4) the probability of extinc- identifying and giving priority of preservation and enhance-
tion in a given subunit population will be only weakly ment to wetlands containing the most important population
influenced by proximity of neighboring subunits, but, subunits (Heard et al. 2012).
Fig. 6.11 Observed wetland

occupancy by the growling grass
frog (Litoria raniformis) in
wetlands adjoining the
metropolitan area of Melbourne,
Australia (blue area), 2001.
Circles denote wetlands (pools
adjacent to streams and lentic
(lake adjoining) wetlands) where
the frog was detected (filled
circles) or not detected (open
circles). Rectangles enclose areas
of mark-recapture studies used to
estimate dispersal and site fidelity.
Elsevier, from Heard, G.W.,
Scroggie, M.P., Malone, B.S.,
2012. Classical metapopulation
theory as a useful paradigm for the
conservation of an endangered
amphibian. Biological
Conservation 148, 156–166.
was not always the case. Early iconic figures in animal

6.3 Detecting Populations for Conservation ecology, like Charles Elton, and in wildlife management,
Management Aldo Leopold, wrote in detail about “census” methods that
should be used to estimate population distribution and den-
6.3.1 The Problem of Detection sity (Elton 1927; Leopold 1933). Such methods were in fact
not censuses, but no methodology was available to correct for
Accurate knowledge of the number of organisms in an area of undetected individuals. An absence of a solution does not
interest is critical to effective conservation, but current remove a problem, and three dilemmas arise from calling
managers of wildlife populations recognize that no survey such sampling efforts “censuses.” First, a great deal of effort
methodology perfectly detects every individual in the sur- is expended to produce results with low reliability. Second,
veyed area, much less the entire population. Such perception management decisions “guided” by such unreliable data are
6.3 Detecting Populations for Conservation Management 225
Fig. 6.12 (a) Dispersal distance a

and proportion of observed 1.0 1.0
movements of the growling grass
Proportion of observed movements

frog (Litoria raniformis) in
wetlands adjoining the 0.8 0.8
Probability of dispersal
metropolitan area of Melbourne,
Australia. Dashed line shows 0.6 0.6
proportion of movements (y axis)
at least as far as corresponding
distance (x axis). Solid line is 0.4 0.4
resulting negative power function
based on observed distances that
an individual frog would disperse 0.2 0.2
as far as each distance during its
lifetime. (b) Proportion of frogs
captured during study that would 0.0 0.0
disperse at least as far as next 10 50 90 130 170 210 250 290 330 370 410 450
nearest wetland. Solid black line
represents mean estimate, dashed Distance (m)
lines 95% CI. Estimates based on b
10,000 computer simulations per 2000
frog. (Reprinted by permission
from Elsevier, from Heard, G.W.,
Scroggie, M.P., Malone, B.S.,
1500
2012. Classical metapopulation
theory as a useful paradigm for the
conservation of an endangered
Frequenct
amphibian. Biological 1000

500
0.0 0.1 0.2 0.3 0.4

Proportion dispersing to nearest wetland
likely to be wrong. Third, when the same “census” methods from numbers of individuals flushed. Biologist J. T. Emlen,
are used year after year with no estimate of the probability of conducting surveys of songbirds in Wisconsin (USA),
detecting individuals, even long-term collections of data have attempted to solve the same kind of problem by developing
low reliability, accumulate high levels of error, and produce species-specific “detection threshold distances” that
incorrect inferences. estimated the maximum distance at which an he could hear
Ecologists engaged in “distance sampling,” which has the song of a given species, varying from 80 m for a black-
become a subdiscipline of its own in modern ecology, were capped chickadee (Parus atricapillus) to 300 m for an oven-
among the first to perceive and attempt to correct for bird (Sierus aurocapillus) (Table 6.1) (Emlen 1984).
problems associated with imperfect detection. They Similar kinds of efforts were used in mammal surveys,
recognized, for example, that probability of detection especially in large, wide-ranging species, where investigators
declines with distance from the observer. One attempt to developed methods of detecting the presence of individuals
address the distance-detection problem was the calculation of different population status (male or female, resident or
of “flushing distances” used in estimating densities of game non-resident) using indirect methods such as track surveys
birds like pheasants, grouse and quail, which were often on roads with different kinds of substrates, as Fred Van Dyke
counted when “flushed” into flight from a position on the and his colleagues did for mountain lions (Puma concolor) in
ground along or near a transect line walked by an observer the western United States (Van Dyke et al. 1986). A widely
(e.g., Guthery 1988). Managers determined that nearly 100% used methodology that permitted a quantifiable estimate of
of individuals present would be flushed within short distances undetected individuals was that of “mark and recapture,”
of the line but decreasing proportions at increasing distances. which represents an attempt to determine the size of a popu-
Where such distance-detection relationships could be lation only imperfectly detected by capture effort, as was
quantified, managers attempted to derive estimates of density used in the previously described study of the growling grass
Fig. 6.13 Observed turnover in

wetland occupancy by the
growling grass frog (Litoria
raniformis) in wetlands adjoining
the metropolitan area of
Melbourne, Australia (blue area),
2001–2007, showing wetlands
where frogs were detected every
year (filled circles), wetlands
where frogs were never detected
(open circles), wetlands at which
extinction was observed
(x enclosed by circle), wetlands at
which recolonization was
observed (partially filled circle)
and wetlands at which both
extinction and colonization were
observed (open circle surrounding
smaller filled circle). (Reprinted
by permission from Elsevier, from
Heard, G.W., Scroggie, M.P.,
Malone, B.S., 2012. Classical
metapopulation theory as a useful
paradigm for the conservation of
an endangered amphibian.
Biological Conservation
148, 156–166. # 2012 Elsevier
Ltd)
frog (Heard et al. 2012). To understand the logic of a mark (proportion of marked individuals to total individuals at first
and recapture study, let us frame relationships between cap- captures) should be equivalent to R/T (the ratio of recaptured
tured and uncaptured individuals this way. Let N be the individuals to all individuals captured in the second effort) or
number of individuals in a population; M the number of
individuals “marked” in some way and then released follow- M=N ¼ R=T:
ing an initial effort; R the number of marked individuals
caught in a subsequent capture effort (“marked recaptures”) Then, by cross multiplication
and T the total number of individuals captured in that
subsequent effort (second sample). The ratio of M/N MT ¼ RN:
Table 6.1 An early attempt to incorporate differential detection probability into animal surveys by ornithologist J.T. Emlen
Total number of
Strip size Density (territory equivalents per km2)d detectionse
Widtha Areab Number of territories June (weeks) July (weeks) Territory sizef
Species (m) (ha) locatedc 1st 2nd 3rd 4th 1st 2nd 3rd 4th within between (ha)
Mourning Dove (Zenaida macroura) 200 90 14 9.5 10.7 11.6 13.6 13.6 13.3 13.3 12.1 162 3 1.8
Red-bellied Woodpecker (Melanerpes 120 54 10 15.0 15.0 15.0 15.0 16.9 16.9 16.9 16.9 103 6 1.6
carolinus)
Downy Woodpecker (Picoides 120 54 (?)g – – – – – – – – – – –
pubescens)
Northern Flicker (Colaptes auratus) 240 108 6 1.9 3.3 4.3 4.3 4.3 4.3 4.3 4.3 54 5 1.6
Eastern Wood-Pewee (Contopus 140 63 5 4.9 4.9 5.9 5.9 5.9 5.9 5.9 5.9 66 2 1.8
virens)
Great Crested Flycatcher (Myiarchus 200 90 16 13.9 13.9 13.9 13.0 11.0 10.1 10.1 8.1 157 12 2.0
crinitus)
Eastern Kingbird (Tyrannus tyrannus) 120 54 1 1.0 1.0 1.0 1.0 1.0 0 0 0 15 0 2.5
Blue Jay (Cyanocitta cristata) 160 72 35?g – – – – – – – – 318 102 0.9
Black-capped Chickadee (Parus 80 36 15 10.8 12.5 12.5 12.5 12.5 12.5 12.5 12.5 146 0 3.5
6.3 Detecting Populations for Conservation Management
atricapillus)
Tufted Titmouse (Parus bicolor) 300 135 3 1.7 1.7 1.7 1.7 1.7 1.7 1.7 1.7 19 2 5.0?
White-breasted Nuthatch (Sitta 120 54 12 13.5 13.5 13.5 13.5 13.5 13.5 13.5 13.5 124 0 3.0
carolinensis)
House Wren (Troglodytes aedon) 140 63 33 26.3 26.0 28.4 25.0 23.5 23.5 25.4 20.5 455 13 0.8
Blue-gray Gnatcatcher (Polioptila (?)h (?) 1 – – – – – – – – – – –
caerulea)
Wood Thrush (Hylocichla mustelina) 300 135 17 9.9 9.9 10.4 9.8 9.8 10.5 10.5 9.8 325 11 1.3
American Robin (Turdus migratorius) 140 63 14 16.2 16.2 16.2 16.2 16.5 16.3 18.3 18.3 252 47 2.2
Gray Catbird (Dumetella carolinensis) 140 63 36 42.7 42.7 39.5 38.3 32.9 32.1 31.6 28.4 484 7 1.0
Brown Thrasher (Toxostoma rufum) 200 90 9 7.6 7.6 8.1 8.1 8.1 8.1 7.6 7.6 32 0 1.4
White-eyed Vireo (Vireo griseus) (?)h (?) 1 – – – – – – – – – – –
Red-eyed Vireo (Vireo olivaceus) 200 90 0 0 0 0 0 0 0 0 0 0 17 –
Blue-winged Warbler (Vermivora (?)h (?) 1 1.0 1.0 0 0 0 0 0 0 12 0 –
pinus)
Ovenbird (Seiurus aurocapillus) 300 135 3 1.9 1.9 1.9 1.3 1.3 1.3 0.5 0.4 37 15 3.0
Common Yellowthroat (Geothlypis 180 81 9 5.7 5.7 5.7 5.7 5.7 3.6 2.3 2.3 44 2 0.7
trichas)
Scarlet Tanager (Piranga olivacea) 200 90 3 2.3 2.3 1.5 1.5 1.5 1.5 0.7 0.7 17 3 2.5
Northern Cardinal (Cardinalis 280 126 27 16.1 16.1 16.1 16.1 16.1 16.1 16.1 16.1 460 10 3.0
cardinalis)
Rose-breasted Grosbeak (Pheucticus 200 90 8 6.4 6.4 8.4 8.4 8.4 6.2 4.0 0 55 21 0.8
ludovicianus)
Indigo Bunting (Passerina cyanea) 200 90 9 6.7 6.7 6.7 5.8 5.2 5.2 3.6 3.0 88 5 1.3
200 90 2 2.2 2.2 2.2 2.2 2.2 2.2 2.2 2.2 31 4 1.5
227
(continued)
Table 6.1 (continued)
228
Total number of
Strip size Density (territory equivalents per km2)d detectionse
Widtha Areab Number of territories June (weeks) July (weeks) Territory sizef
Species (m) (ha) locatedc 1st 2nd 3rd 4th 1st 2nd 3rd 4th within between (ha)
Rufous-sided Towee (Piplio
erythrophthalmus)
Field Sparrow (Spizella pusilla) 160 72 4 3.8 3.8 3.8 3.8 4.4 4.4 4.4 4.4 43 0 0.8
Song Sparrow (Melospiza melodia) 140 63 6 2.2 2.2 2.2 3.5 3.5 3.5 3.5 3.5 21 0 1.3
Brown-headed Cowbird (Molothrus (?) (?) (?)g – – – – – – – – – – –
ater)
Northern Oriole (Icterus galbula) 200 90 2 2.2 2.2 2.2 2.2 2.2 0 0 0 10 4 1.8
American Goldfinch (Carduelis tristis) 200 90 1 0 1.1 1.1 1.1 1.1 1.1 1.1 1.1 28 29 2.3
The column designated “width” refers to the maximum distance at which Emlen could hear the song of a particular species.
Reprinted by permission from American Ornithology Society, from Emlen, J.T., 1984. An Observer-Specific, Full-Season, Strip-Map Method for Censusing Songbird Communities. The Auk
101, 730–740
a
Observer’s detection-threshold distance times 2
b
Strip width strip length (4,500 m)
c
Number of territories (represented wholly or fractionally) located within the strip
d
Sum of territories (total of fractions and wholes) per km2 for each of 8 weeks – June and July
e
Number of detections recorded (a) within and (b) between the detection-point clusters (territories) during the 45 traverses
f
Mean area of detection-point clusters (territories)
g
Clustering of detection points too loose to indicate clearly the location of localized (territorial) resident birds. Indications of clustering, presumably near nests, prompted a guess of ca. 35 home areas
for the Blue Jay
6
h
Data were insufficient to provide a useful estimate of the detection-strip widths
The Conservation of Populations: Theory, Analysis, Application
If we now divide both sides by R, we have surveys of the same site and development of a specific prob-
ability theory of detection that could employ data based on
MT=R ¼ N, repeated visits to the same site. Along with scientists
associated with ARMI (e.g., MacKenzie et al. 2002), other,
and can now solve for the value of N. similar methodologies were independently created at about
A mark and recapture efforts is itself an attempt to esti- the same time by investigators attempting to solve the same
mate detection probability. In this case, R/T is the estimate of problem in different contexts (e.g., Tyre et al. 2003; Gu and
detection. You then divide M by this detection estimate to Swihart 2004). Theories and models of occupancy subse-
obtain N. This approach assumes that there is no ingress or quently developed as an attempt to determine the probability
egress in the studied population and that there is equivalent that a site, habitat patch, or other sampling unit was
survivorship among marked and unmarked individuals “occupied” or “unoccupied” by the species of interest. With
between the period of capture and recapture. Originally models developed by a number of scientists, as well as
employed as a method of estimating abundance of fish computer software that permits flexibility in assumptions
(Petersen 1896), use of these equations was expanded to associated with sampling, investigators now possess tools
applications of estimating abundance of waterfowl based on that enable precise calculation of the probability of occu-
band returns supplied by hunters (Lincoln 1930), and is still pancy and detection of a given species, ranging from deter-
widely used for this purpose. mining probability of detection for a single site visit to the
The Lincoln-Petersen Index (or Lincoln-Petersen Estima- probability of detecting the species over the course of multi-
tor, as it is also called) attempts to determine the “true” ple, repeated visits to the same site. For example, in a study of
number of individuals in a population and is one means of amphibian use of forest pools in the US state of Illinois, the
estimating numbers of undetected individuals. For this rea- authors determined, through repeated visits to the same pools
son, it is still widely used and applied. However, conserva- over a 3-year period, that the probability of detecting the
tion biologists today often want management decisions presence of the northern leopard frog (Lithobates pipiens)
informed either by an accurate measure of relative abundance on any single visit was 0.22, but the probability of detecting
of individuals, or, if this is not possible, an accurate estima- that species at a given site through repeated visits over a
tion of whether a species is present in a particular area. 3-year period was 0.98 (Van Dyke et al. 2017).
Presence is easier to estimate than abundance, and sometimes Accurate information about occupancy requires reliable
is the only population indicator that can be used when estimation of spatial variation (in number of animals) and
individuals are extremely rare or difficult to detect. detectability (how hard is it to know if the animal is there).
Differences in one or both of these variables in different areas
will produce different probabilities of detection (Fig. 6.14).
6.3.2 Occupancy Theory and Modeling Because true “censuses” of any species or area are rare
(because they are usually unaffordable in expense, time,
Given the importance of detection to science in general and and effort), investigators must choose sampling units to rep-
conservation biology in particular, efforts have continued to resent the entire area. When they do, it is critical that they
solve the problem of detectability and occupancy, and have select units that permit them to not only make inferences
increased in sophistication, leading to the development of a about the sites they have selected in their sample, but also
new branch of conservation science known as occupancy about sites they have not selected. Once the choice of sam-
estimation and modeling. The development of occupancy pling units has been made, measuring the probability of
modeling and related occupancy theory was first stimulated detection is critical because some animals will go undetected
by the need to solve problems associated with detection of even in units that are sampled.
amphibians in the US Geological Survey’s Amphibian Let’s work out the logic of estimating detectability this
Research and Modeling Initiative (ARMI), started in the way. Let Nit represent the true or actual number of animals
1990s. ARMI’s goal was to make comprehensive national of the species of interest in a particular sampling unit or area, i,
surveys and inventories of amphibian species in the United at a particular time or date, t. Let Cit represent the “count
States, but ARMI researchers soon realized that it was impos- statistic” (for example, the number of animals actually
sible to estimate absolute changes in amphibian abundance counted) in the survey of those individuals in area I at time t.
over large areas covering different time periods, and so Assume that C is a random variable whose expected value,
turned to simpler measures of detecting site-specific presence E (Cit) (i.e., its average value if we conducted the same survey
or absence. It became apparent that a target species would not under the same conditions at the same site many times), will be
always be detected on a single site visit. Therefore, analysis equal to the total number of individuals actually present (N)
and interpretation of data would have to account and correct times the detection probability, Pit (the probability of detecting
for imperfect detection. Such estimation required repeated an individual when it is actually there, or, put another way, the
conservation biologist wants to intelligently determine

which areas to protect. This leads us to further examination
of the problem of site occupancy. The probability that a site is
occupied, ψ, is the ratio of occupied sites (x) to the total
number of sites surveyed (s), or Ψ ¼ x/s (MacKenzie et al.
2006). We will have more to say about this problem in the
following chapter (Chap. 7) on habitat conservation, but, for
the moment, we will examine some applications of
estimating occupancy and detectability and the technologies
that can be used to that apply directly to make these
estimates.
6.3.3 Developing Technology and Applications

in Occupancy Modeling
Detection probability cannot be improved, or even deter-

mined, by “more field studies” (a common and essentially
Fig. 6.14 Illustration of two critical aspects of sampling animal worthless recommendation by authors of publications that
populations, spatial variation and detectability. The blue region present unreliable data). Detection probability is improved
represents the area or population of interest, with squares representing by better design of sampling methods in existing field studies
selected sampling locations. At each sampling location, animals are
detected (filled circles) or undetected (hollow circles) during a survey and by use of computer software specifically designed for this
or count. (Reprinted by permission from Elsevier, from MacKenzie, D. type of analysis. One of the most widely used computer
I., Nichols, J.D., Royle, J.A., Pollock, K.H., Bailey, L.L., Hines, J.E., applications for determining probability of occupancy and
2006. Occupancy Estimation and Modeling: Inferring Patterns and detection was developed by MacKenzie et al. (2003) in
Dynamics of Species Occurrence. Academic Press. # 2006 Elsevier
Inc. Modified by K. DeVoss) their program PRESENCE (the program and supporting
files are available at https://www.mbr-pwrc.usgs.gov/soft
ware/presence.html, a site maintained by the US Geological
probability that an individual that is present will actually be Survey’s Patuxent Wildlife Research Center in Maryland
included in the count statistic, Cit). This is the basic conceptual (USA)). Beginning with estimation of the so-called “naive
problem for every census technique that is based on counting detection rate” (the proportion of sampling events (surveys)
individuals. Symbolically, then, in which the species is detected given that it is actually
present at the site), PRESENCE and other similar models
E ðCit Þ ¼ N it Pit can then adjust for seasonal effects of detectability (for
example, increased vocalization or activity in a breeding
Now we could solve for N (MacKenzie et al. 2006), if only season), as well as calculate estimates of the species initial
we knew the probability of detection. Investigators past and occupancy (proportion of sites occupied at the beginning of
present sometimes have tried to get around the detection the study), colonization (rate at which unoccupied sites
problem by standardizing sampling methods, training all become occupied between years), extinction (rate at which
observers the same way, and sampling only under a carefully occupied sites become unoccupied between years), and
specified set of “ideal” conditions. Such actions represent single-visit detection probability (probability of detecting a
attempts to equalize detection probability on every survey species during a sampling event if it is present) (Van Dyke
by reducing methodological, observational, and environmen- et al. 2017, Table 6.2). Detection probability is itself a vari-
tal biases. But, in spite of these efforts, detection probabilities able that can be affected by multiple factors. For example,
will not be equal because there are always factors affecting Murn and Holloway (2016) evaluated factors affecting detec-
detection that are not accounted for, including the behavior of tion probability in surveys of the cryptic, elusive, and IUCN
the animal itself, and unexplained variation will influence Critically Endangered raptor, the white-headed vulture
detectability. Therefore, estimation of detection probability (Trigonoceps occipitalis) (Fig. 6.15), a native of
remains a real problem that must be solved in any serious sub-Saharan Africa. They determined that, as was the case
conservation effort. in our previous example of the leopard frog, probability of
Detection is not only important for accurate estimation of detection increased as number of visits to the same site
numbers of individuals, but also for estimating their range increased, but they also discovered that variability of detec-
and distribution, which is critical information if a tion probability decreased (Fig. 6.16), demonstrating that
Table 6.2 Detection probabilities (p) of amphibian species based on repeated surveys during breeding seasons in northern Illinois (USA) forest
preserves, 2008–2010
Species Naïve P for single survey X number of surveysa Probability of detection during studyb
Eastern tiger salamander 0.149 67.4 1.000
American toad 0.148 46.7 1.000
Cope’s treefrog 0.108 35.7 1.000
Gray treefrog 0.960 5.3 1.000
American bullfrog 0.464 17.9 1.000
Green frog 0.368 35.9 1.000
Northern leopard frog 0.214 16.0 0.983
Eastern newt 0.188 67.4 1.000
Spring peeper 0.477 54.6 1.000
Western chorus frog 0.644 34.5 1.000
The “naïve detection rate” refers to the proportion of sampling events (surveys) in which the species is detected given that it is actually present at the
site. A “true-” or “model-based” probability of single-visit detection can then be determined through additional analysis and calculation, which is
normally greater than the naïve detection rate. Through additional analysis, conservationists can determine the probability of detecting a species, if
present, over the course of the entire study. Where sampling effort is sufficient (i.e. repeated), “study detection probability” will often be 1.0 (certain
detection), increasing credibility of estimation of population presence or absence. Through the determination of such probabilities, conservationists
obtain improved precision in their estimation of population size and site occupancy
Reprinted by permission from Wiley, from Van Dyke, F., Berthel, A., Harju, S.M., Lamb, R.L., Thompson, D., Ryan, J., Pyne, E., Dreyer, G., 2017.
Amphibians in forest pools: Does habitat clustering affect community diversity and dynamics? Ecosphere 8, e01671. # 2017 The Authors
a
Average number of surveys conducted during each species breeding season over the course of the study (i.e., 3 years)
b
Model-based probability of detecting a species during the study period, given that it was present, for the average number of surveys conducted at a
site during each species breeding season (note: Assumes geographic closure among years – no local extinction/colonization)
(Fig. 6.17). The slower the method of travel, the higher the
probability of detection. As a result, Murn and Holloway
concluded that taking more time to conduct surveys for the
white-headed vulture and other species with similar
characteristics lowered the number of surveys (replication
effort) needed to achieve high probabilities of detection
with lower variation (Murn and Holloway 2016).
Increased precision in estimating the probability of
detecting a species at a given location adds insight to deter-
mining what the species needs for its conservation, including
needs associated with habitat use, landscape distribution, and
home range. Thus, improved reliability in estimation of
detection and occupancy leads to better decisions about
which sites to protect to conserve specific populations, spe-
cies and communities.
Because targeted species do not always stay put, accurate
assessment of changes in habitat occupancy is now one of the
most important measurements needed to estimate dispersal
(Driscoll et al. 2014). Occupancy detection is not only impor-
Fig. 6.15 The white-headed vulture (Trigonoceps occipitalis), an tant in making estimates of dispersal and dispersal-influenced
IUCN Critically Endangered species native to sub-Saharan Africa that population dynamics but is now also one of the most impor-
is cryptic and difficult to detect through surveys, and so is an important
tant metrics used to estimate immigration, colonization, and
species for determining detection probability and the factors that affect
it. (Photo courtesy of D. Irbis, reprinted under CC BY-SA-3.0) local site extinction (site-specific extirpation). The value of
occupancy theory in conservation has increased even more
estimates of detection probability were not only higher with because it can now be used in models evaluating multiple-
repeated surveys, but more reliable. They also found that species and multiple-seasons as well as in single-species,
survey method, whether on foot or through a combination single-season studies. The importance of occupancy estima-
of foot and vehicle travel, also affected detection probability tion will continue to grow in conservation biology, and its
as a function of the speed at which the observer was traveling uses will be more and more widely applied.
Fig. 6.16 Cumulative probability of detecting a white-headed vulture (Reprinted by permission under CC BY 4.0 International, from Murn,
in an occupied breeding territory as a function of the number of repeated C., Holloway, G.J., 2016. Using areas of known occupancy to identify
visits to that territory. Note that, as the number of site visits increase, not sources of variation in detection probability of raptors: taking time
only does probability of detection increase, but variation associated with lowers replication effort for surveys. Royal Society Open Science
probability of detection decreases, indicating increasing reliability of the 3, 160368. # 2016 The Authors)
detection estimate. Vertical bars represent upper and lower 95% CIs.
Fig. 6.17 Effect of increasing

average speed of observer
(influenced by traveling only on
foot or by a combination of foot
and vehicle) during a survey on
the probability of detecting white-
headed vultures in occupied
breeding territories. (Reprinted by
permission under CC BY 4.0
International, from Murn, C.,
Holloway, G.J., 2016. Using areas
of known occupancy to identify
sources of variation in detection
probability of raptors: taking time
lowers replication effort for
surveys. Royal Society Open
Science 3, 160368. # 2016 The
Authors)
6.4 Minimum Viable Populations 233
6.4 Minimum Viable Populations
6.4.1 General Considerations
Recall that genetic considerations were the original source of

estimates of minimum viable populations (MVPs) in conser-
vation biology, beginning with Franklin’s (1980) suggestion
that effective population sizes of 50 were needed to prevent
deleterious effects of inbreeding, and a minimum effective
size of 500 was required to maintain sufficient genetic varia-
tion to offset effects of genetic drift (Chap. 5). We have
already noted that such “rules” were too general to be useful
to specific populations. In most cases, much larger numbers
are required. Of greater concern is the fact that such an
estimate of MVP reflects only genetic considerations.
Today population viability analysis also considers demo-
graphic and environmental characteristics of a population,
shifting emphasis from minimum population size to the
probability of the population’s persistence through time.
What approaches can help estimate persistence and extinc-
tion probabilities based on accurate understanding of causes
of population decline?
6.4.2 Trend Analysis and Factor Resolution: Fig. 6.18 Logical flow of six sequential steps in population conserva-
tion to move a threatened population to a state of recovery and demo-
Systematic Approaches for Identifying graphic stability. (Figure concept by F. Van Dyke, illustration by
Causes of Population Decline K. DeVoss)
and Strategies for Restoration
provide an index of the population’s status- they cannot tell
Population decline and potential restoration can be us the prospects for the population’s persistence or the causes
investigated through long-term studies and associated models of its decline. Instead, we must advance from simple surveys
of population viability analysis, but small populations them- to demographic monitoring. Demographic monitoring
selves often must be saved through quick-witted adaptive follows the fates of individuals in a population over time
management strategies that can rescue a species on the brink and, like the presence-absence surveys previously discussed,
of extinction. Can we identify, with relatively short-term field makes repeated on-site visits to gain direct demographic
experiments, factors leading to a population’s decline and measurements of the population. Two important monitoring
management treatments that could be most effective in its tools that provide a means to gain insights into potential
restoration? To do these things, we must consider how to causes of and solutions to population decline are trend analy-
design experiments targeting specific demographic factors sis and factor resolution (Pavlik 1994).
and their potential impact on population restoration efforts. Trend analysis calculates the value of specific demo-
Restoring individual populations of plants or animals nor- graphic variables in one or more populations and, from
mally addresses one of two conditions. One is the restoration such calculations, determines if the population is growing,
of threatened or endangered populations. The other is the stable, or declining. Trend analysis alone cannot identify the
restoration of more common species on sites from which cause of the population’s low numbers or decline, but it can
they have been exterminated or to sites where they were not lead to intelligent guesses (hypotheses) about what factors to
historically present but are expected to do well. Both efforts investigate. For example, in plant populations, some general,
usually follow a six-part process depicted in Fig. 6.18. non-mathematical indicators of population stability can be
We have already seen (Sect. 6.3) the importance of used in trend analysis to make an initial assessment of
repeated site-specific surveys to estimate the detectability of whether an introduced plant population is stable and likely
populations, as well as their presence and abundance. Persis- to persist at its new site (Table 6.3). For example, a relatively
tent monitoring is also critical to determine the status of an short-lived, herbaceous species experiencing high rates of
endangered population, but although such surveys can annual adult survivorship but low rates of seed germination
Table 6.3 Demographic parameters that serve as general indicators of population stability in analysis of trends in endangered plant populations
Parameter Life Form Population Stable if
Survivorship Annual Mortality inflection point on survivorship curve (type I) follows onset of seed production.
Perennial The number of individuals in a new cohort equals or exceeds the number of established individuals
after inflection point on survivorship curve (type III).
Seed bank All Density of viable seeds in soil prior to season of germination far exceeds the average density of
established individuals.
Annual and Year-to-year changes in density of viable seed are not correlated with changes in the density of
herbaceous perennial established, reproductive individuals.
Seed production All Seed production per individual of an endangered taxon equals or exceeds that of a nonendangered
relative with similar life form.
Age structure Perennial Number of established, reproductive individuals is less than the number of established juveniles
and/or the number of recruited seedlings.
Frequency of Annual Frequency of establishment is less than the half-life of seeds in the seed bank.
establishment Perennial Frequency of establishment is less than the half-life of established, reproductive plants.
Design by M. J. Bigelow, based on data and concepts from Harper (1977) as summarized by Pavlik (1994)
suggests that environmental variables affecting seeds, such as the fiddleneck to California grasslands where it was no longer
soil moisture, seed predation, or soil temperature, might have present. In natural environments, some competition between
more to do with the decline than factors affecting adult plants, the fiddleneck and other plants would be inevitable.
such as ambient temperatures, herbivory, or competition. For Conservationists wanted to achieve restoration of the
example, trend analysis for the large-flowered fiddleneck fiddleneck in grassland communities of indigenous species.
(Amsinckia grandiflora), an endangered annual forb (Family But could it also persist with non-native grasses?
Boraginaceae) native to dry grasslands of California (US), To answer this question using a factor analysis approach,
suggested that competition with other plant species was the investigators measured the performance of Amsinckia gran-
primary limiting factor in population recovery (Pavlik 1994). diflora in grasslands dominated by native Sandberg bluegrass
A logical strategy then was to remove competing species (Poa secunda) compared to grasslands where non-native
from sites where the fiddleneck occurred. The most com- grasses were present at varying densities. Using the number
monly used treatments to remove competitors are herbicides of inflorescences (flower heads) on individual fiddlenecks as
and prescribed burning. Both treatments remove competitors, an index to reproductive performance, they found that both
albeit in different ways. Figure 6.19 shows the experimental types of grasslands reduced numbers of fiddleneck
design derived from trend analysis that Bruce Pavlik and his inflorescences at high grass densities, but native grasses
colleagues used to identify which treatment had greatest reduced fiddleneck inflorescences less at low and intermedi-
effect. Table 6.4 shows the results of the experiment. ate densities (Carlsen et al. 2000) (Fig. 6.20). They
Based on these results, we can see how trend analysis can hypothesized that these differences might be due to
guide the second phase of demographic monitoring, factor differences in growth forms in the grasses. Whereas non-
resolution, by helping investigators formulate specific native species tended to form a solid mat with few openings,
hypotheses in which experimental tests of one or more factors Sandberg’s bluegrass and other native grasses grew in well-
suspected of limiting population growth are conducted in the defined clumps, with openings between clumps, especially at
field. The results help to determine which factors limit popu- low and intermediate densities, providing more opportunity
lation growth and allow managers to identify which variables for the large-flowered fiddleneck to grow in competitor-free
to manipulate for the best chance of population recovery. space. Based on these findings, the investigators concluded
A second problem in population restoration is how to that “restored native perennial grasslands of intermediate
create new populations on sites where they do not exist. To densities have a high habitat value for the potential establish-
do this, the manager can begin with factor resolution and ment of the native annual A. grandiflora” (Carlsen et al.
proceed to trend analysis. Experimental tests would be 2000:18). Together, these studies show how well-designed
conducted to identify factors limiting the growth of the pop- experiments used in trend analysis can inform field
ulation. The results would then be used to determine the investigations using an approach of factor resolution,
trajectory of population growth and what ought to be done incorporating both to inform management decisions likely
to enhance it. Studies of the large-flowered fiddleneck again to lead to successful establishment of A. grandiflora.
offer a good example. Here the long-term goal was to restore
Fig. 6.19 Experimental design used to create a new population of the (Fusilade) treated) or a control in order to measurehe effects of compe-
endangered plant, Amsinckia grandiflora, within its historic range. Each tition from non-native grasses. (Design by M. J. Bigelow, based on
of the 20 plots was either a treatment (burned, hand clipped, or herbicide Pavlik 1994)
Table 6.4 Results of experimental treatments on germination, population size, survivorship, plant size and nutlet (seed) production of the
endangered plant Amsinckia grandiflora
Population Size Survivorship (% of Mean maximum Nutlet (Fruit)
Treatment Germination (Reproductive Plants/plot) Germination) to Reproduction Plant Size (cm) Production (No./Plant)
Control 55.4 5.2a 38.6 15.8a 42.7 16.5a 26.0 3.1a 15.1 10.1a
Burn 55.4 9.9a 67.2 19.8a 75.3 11.6b 33.7 5.3b 29.1 14.4a
Clip 54.1 4.8a 57.8 16.5a 63.1 12.0a 23.1 3.7a 6.6 5.6a
Herbicide 54.0 8.1a 56.4 15.6a 64.4 10.8a 40.5 4.1b 53.5 16.5b
Values (mean SD) in a column followed by the same letter are not statistically different (P < 0.05, ANOVA)
Note how burning significantly increased survivorship and plant size, while herbicide treatment increased plant size and fruit production per plant.
Design by M. J. Bigelow, based on data from Pavlik et al. (1993)

Reintroduction of endangered species is essential for their
Consider the data in Table 6.4. All three management
conservation, but success can be elusive despite repeated
treatments are designed to achieve the same result –
effort. Nowhere is this more evident than in attempts to
increased populations of Amsinckia grandiflora
restore populations of endangered carnivores which, even
through elimination of competing species – but by
under ideal conditions, may have always had low numbers,
different means. Considering the results, which treat-
and may be perceived negatively by humans. To understand
ment or combination of treatments would you use to
the complexity of problems confronting such an effort, we
achieve the largest population?
examine the ongoing restoration of one endangered
Fig. 6.20 Relationship of Amsinckia grandiflora total inflorescence

Fig. 6.21 An adult female black-footed ferret (Mustela nigripes) in a
number per plot (measured 12 April 1994) to final dry grass biomass
“ring reader” at the BLM-40 release site in Montana, USA photographed
(measured 1 May 1994), showing predicted means (middle lines) and
in September 2006. The ring reader is a device used to identify individ-
95% confidence intervals (outer lines). (Reprinted by permission from
ual ferrets released into the wild from a captive breeding program. Each
Wiley, from Carlsen, T.M., Menke, J.W., Pavlik, B.M., 2000. Reducing
released ferret is subcutaneously implanted with a Personal Identifica-
Competitive Suppression of a Rare Annual Forb by Restoring Native
tion Tag (PIT tag). Biologists observed ferrets at night using spotlights
California Perennial Grasslands. Restoration Ecology 8, 18–29. # 2000
and place the ring reader over the burrow into which the ferret descends
Society for Ecological Restoration)
when approached. The reader receives a signal from the PIT tag unique
to that individual. The black-footed ferret, one of the world’s rarest and
carnivore, the black-footed ferret (Mustela nigripes), a spe- most specialized carnivores, provides an exemplary case study for
understanding the problems, pitfalls, and potential for success in restor-
cies that has required an unprecedented breadth of interdisci- ing an extinct wild population using captive-bred individuals and
plinary expertise to save it from extinction. perseverant, repeated, long-term reintroduction efforts at multiple and
geographically widespread sites. (Photo courtesy of Dean Biggins/US
Geological Survey)
6.4.3 Saving a Population from Extinction: The
Case of the Black-Footed Ferret of hectares, creating a net reduction in available land for
grazing and agriculture, and burrow openings themselves
6.4.3.1 Population Recovery: Risk Taking, Creative create risk of injury for domestic livestock, as well as for
Breeding, and Reintroduction horse-mounted riders who care for the livestock. Given the
The black-footed ferret (Fig. 6.21) is one of the world’s most habitat changes and potential hazards created by prairie dogs,
specialized carnivores. Black-footed ferrets spend 90% of landowners have poisoned, trapped, and shot them persis-
their time underground (Biggins et al. 1986), preying selec- tently and in large numbers, often with the aid of government
tively on prairie dogs (Cynomys spp.), burrowing rodents money and expertise.
native to western North America. Ferrets exploit prairie Since the 1930s, prairie dogs also have been devastated by
dogs not only as a food source, but also for habitat. Ferrets sylvatic plague (Yersinia pestis), a bacterium introduced to
do not dig their own burrows but live in those found in prairie the United States in 1900 as a result of trade between east
dog colonies. To support a population of ferrets, a colony of Asia and the US West Coast. Following its arrival, sylvatic
prairie dogs must be relatively large, not only to provide plague spread through 17 western US states, and prairie dogs
sufficient prey and habitat, but because ferrets are territorial have shown no immunity to it (Eads and Biggins 2015). This
and avoid sharing space with other ferrets of the same sex combination of habitat loss, direct extermination, and disease
(Eads et al. 2014). have reduced prairie dog numbers and distribution to the
Unfortunately for black-footed ferrets, prairie dog point that today they occupy less than 5% of their historic
colonies have been systematically eliminated throughout range. Ferrets can contract the plague from prairie dogs.
their historic range because they occupy habitat that is or With the decline in prairie dogs and introduction of
could be used for livestock grazing or agriculture. Prairie plague, black-footed ferrets declined precipitously. By the
dogs clip herbaceous vegetation in and around their colonies early 1960s, the ferret was considered extinct in the wild,
in ways that shift vegetation communities toward greater but one population was discovered in 1964 in South Dakota
abundance of forb (non-grass) species, reducing forage for (US). As part of the recovery plan mandated by the US
domestic cattle, horses, and sheep. Extensive and contiguous Endangered Species Act, a breeding program was initiated
colonies and their burrow complexes can take up thousands in the 1970s, but the last ferret in the program died in 1979
(Biggins 2012). By this time, the South Dakota population early commitment to freezing semen from males (some of
had disappeared. In 1981, however, another ferret population which later died) and developing more effective methods of
was discovered in Wyoming, but within 2 months that popu- artificial insemination (AI) instead of relying solely on con-
lation had declined to 16 animals as a result of infection with ventional breeding methods (Holt 2016). Over the course of
sylvatic plague and canine distemper (contracted from a the captive breeding effort, semen up to 20 years old succes-
domestic dog). Six ferrets were caught for captive breeding. sively produced pregnancies and subsequent litters in captive
All died. Six more were captured in 1985–1986. At that females (Table 6.5), the first time a combination of frozen
point, the world’s entire black-footed ferret population semen technology and AI techniques had been fully and
consisted of six captive animals and four wild survivors successfully integrated into a captive breeding program
(Biggins et al. 2006). (Holt 2016). Scientists managing the breeding effort followed
a minimization of mean kinship (mk) strategy to mate selec-
6.4.3.2 Beginning the Recovery Process: Captive tion which minimized relatedness of breeding individuals
Breeding and Planned Release (Chap. 5), producing 139 kits with higher levels of gene
In 1986, there were no captive births, but the four wild ferrets diversity (GD, the probability that two alleles from the
produced two litters and a total of six “kits.” At this point, the same loci are not identical, Chap. 5) and lower mk that
Black-footed Ferret Recovery Team (BFRT) decided to cap- other offspring produced by conventional breeding methods
ture all remaining ferrets and bring them into the captive (Fig. 6.23) (Howard et al. 2016).This approach not only
breeding program. Of these, seven ultimately formed the increased the number of captive-bred ferrets produced, but
founding population for ferret recovery. reduced loss of genetic diversity from the now extinct wild
Skilled applications of captive-breeding techniques, veter- population. By 1991, captive production had achieved suffi-
inary medicine, and genetic management were essential to cient success to begin reintroduction efforts.
saving the ferret and preserving its genetic diversity. The The Black-footed Ferret Recovery Implementation Team
program was beginning to produce significant numbers of (BFRIT), formed in 1996 from the earlier BFRT, created
ferrets by the late 1980s (Fig. 6.22), in part because of an criteria for site selection for re-introduction efforts called
the “adaptive allocation matrix” to guide decision making
on site choice (Table 6.6). Primary factors were habitat qual-
Number of kits production in
500
ity, occurrence and current status of sylvatic plague outbreaks
400 and, after initial release, documented kit production and adult
captivity
300 survivorship (Jachowski and Lockhart 2009). After initial

200
direct releases into wild environments in 1991 (“hard”
release), biologists modified their release strategy by first
100
keeping released ferrets in semi-natural conditions in outdoor
0 pens containing prairie dog burrow systems, complete with
live prairie dogs, so captive-raised ferrets could refine their
87
90
93
96
99
02
05
08
19
19
19
19
19
20
20
20
Year
predatory skills (“soft” release). Through 2008, 147 ferrets
had been released at 18 locations on both public and private
Fig. 6.22 Annual production of black-footed ferret kits in captive breed- land (Table 6.7). Some efforts were failures. Male survivor-
ing facilities from seven founder animals beginning in 1987. Note, through ship was especially poor. Predation on captive-raised ferrets
the first 4 years of the captive breeding effort (1990), less than 100 kits had
been produced. After 1990, captive breeding output improved dramati-
was high, especially by coyotes (Canis latrans). Some
cally. (Reprinted by permission from IUCN Small Carnivore Specialist re-introduced populations were devastated by plague, such
Group, from Jachowski, D.S., Lockhart, J.M., 2009. Reintroducing the as the Conata Basin (South Dakota, US) population, which
Black-footed Ferret Mustela nigripes to the Great Plains of North America. declined from 335 animals in 2007 to 32 by 2013 (Eads and
Small Carnivore Conservation 41, 8: 58–61)
Biggins 2015).
Table 6.5 Performance of frozen spermatozoa of black-footed ferrets in terms of motile sperm, pregnancies, and offspring produced (kits) in
captive female ferrets inseminated with sperm 10–20 years old
AI No. of females Range (year) of Mean ( SEM) no. total motile Range no. total motile No. of No. of
year inseminated semen storage sperm/AI per female sperm/AI per female pregnancies kits
2008 4 10–11 5.7 0.8 4.7–6.2 2 2
2009 5 11–20 5.2 0.3 2.1–13.2 2 4
2010 5 13–20 8.5 2.0 2.1–8.7 1 2
2011 4 14–15 4.6 1.2 1.2–7.7 0 0
Note that sperm of all ages in this interval successfully impregnated female ferrets. SEM, standard error of the mean. Reprinted by permission from
Wiley, from Howard, J.G., Lynch, C., Santymire, R.M., Marinari, P.E., Wildt, D.E., 2016. Recovery of gene diversity using long-term
cryopreserved spermatozoa and artificial insemination in the endangered black-footed ferret. Animal Conservation 19, 102–111. # 2015 The
Zoological Society of London
Fig. 6.23 Effect of using artificial insemination (AI) with frozen- breeding, causing a consistent increase in the GD of the total captive
thawed spermatozoa from male black-footed ferrets (Mustela nigripes) population. (Reprinted by permission from Wiley, from Howard, J.G.,
deceased for 10–20 years on gene diversity (GD) in a captive black- Lynch, C., Santymire, R.M., Marinari, P.E., Wildt, D.E., 2016. Recov-
footed ferret population. Note that, through managed mating following a ery of gene diversity using long-term cryopreserved spermatozoa and
minimization of mean kinship (mk) strategy, GD of offspring produced artificial insemination in the endangered black-footed ferret. Animal
from AI was consistently higher that those produced by conventional Conservation 19, 102–111. # 2015 The Zoological Society of London)
Table 6.6 Factors used for prioritizing numbers of black-footed ferrets to potential reintroduction sites (“adaptive allocation matrix”). For each site,
factors are given a score of 1–5, with primary factors weighted by 4, secondary factors by 2, and tertiary factors not weighted
Primary factorsa Secondary factors Tertiary factors
Habitat suitability Long-range site conservation Pre-conditioning capabilities
Plague status Ferret monitoring Contingency planning
Documented kit production Habitat monitoring Veterinary and husbandry support
Ferret survivorship Disease monitoring/management Reintroduction proposal quality
Research benefits Project resource availability
Reprinted by permission from IUCN Small Carnivore Specialist Group, from Jachowski, D.S., Lockhart, J.M., 2009. Reintroducing the Black-
footed Ferret Mustela nigripes to the Great Plains of North America. Small Carnivore Conservation 41, 8: 58–61
a
Alternative primary factors used in place of documented kit production and Ferret survivorship when considering new sites are (1) proposed project
benefits and (2) management/legal status
Despite these discouraging setbacks, the Implementation different prairie dog densities as long as densities were
Team persevered. By 2010, there were an estimated 700–800 greater than 12 prairie dogs/ha (Ayers et al. 2014). Such
ferrets at 19 sites in eight US states, as well as Canada and studies made clear that overall habitat quality (reflected in
Mexico (Biggins 2012; Howard et al. 2016). Further estab- the total area occupied by prairie dogs), not prairie dog
lishment of new populations was no longer dependent on density, was the best predictor of site-specific success.
captive-bred individuals but could now be accomplished by
translocations of wild ferrets from site to site. By 2016, the 6.4.3.3 What Can We Learn from the Black-Footed
captive breeding program had produced more than 8000 Ferret?
ferrets, over 4100 of which had been released into the wild Despite the growth of populations on some sites, as well as
at 28 sites (Howard et al. 2016, Fig. 6.24). increased success in captive breeding, the black-footed ferret
Comparisons of successful versus unsuccessful sites remains an Endangered Species. Yet biologists engaged in
revealed that the most important predictor of success was this effort remain confident that recovery is attainable. To
having a site with at least 4300 ha occupied by prairie dogs combat plague, a vaccine was developed specifically for
(Jachowski et al. 2011). Prairie dog density, considered with- ferrets (Rocke et al. 2004), but must be given by direct
out respect to the area occupied by prairie dogs, was not injection, a difficult and time-consuming procedure (Biggins
different between successful and unsuccessful sites, nor was 2012). There is more hope in a vaccine developed for prairie
incidence of plague, number of ferrets released or number of dogs which can be delivered in bait and thus distributed over
repeated releases, or presence of different species of prairie large areas (Abbott et al. 2012). Prairie dog and ferret
dogs (Jachowski et al. 2011). Additionally, there was no populations are also limited by negative attitudes of private
difference in litter sizes of female ferrets living on sites with landowners. The Wyoming Farm Bureau, for example,
Table 6.7 Sites and results of reintroduction of black-footed ferrets on 18 sites in the United States and Mexico 1991–2008. Note that 9 (50%) of
the 18 sites had less than 20 individuals to this point
Year
Site reintroduction Total number of Estimated current
number Reintroduction site began Land management status Ferrets releaseda populationb
1 Shirley Basin, Wyoming 1991 Private and Bureau of Land 518 239c
Management
2 Badlands National Park, South 1994 National Park Service 244 22
Dakota
3 UL Bend National Wildlife 1994 US Fish and Wildlife 229 10
Refuge, Montana Service
4 Conata Basin, South Dakota 1996 US Forest Service 167 239
5 Aubrey Valley, Arizona 1996 Private 306 66d
6 Fort Belknap Indian Reservation, 1997 Tribal 167 0
Montana
7 Coyote Basin/Snake John, Utah 1999 Bureau of Land 332 11
Management
8 Cheyenne River Indian 2000 Tribal 185 75e
Reservation, South Dakota
9 Wolf Creek, Colorado 2001 Bureau of Land 239 13
Management
10 40-Complex, Montana 2001 Bureau of Land 95 0
Management
11 Janos, Chihuahua, Mexico 2001 Private 257 17
12 Rosebud Indian Reservation, 2004 Tribal 139 28d
South Dakota
13 Lower Brule Indian Reservation, 2006 Tribal 62 26
South Dakota
14 Wind Cave National Park, South 2007 National Park Service 49 18
Dakota
15 Espee Ranch, Arizona 2007 Private 51 20d
16 Logan County, Kansas 2007 Private 74 15
17 Northern Cheyenne Indian 2007 Tribal 38 8d
Reservation, Montana
18 Vermejo Ranch, New Mexico 2008 Private 89f 17
Reprinted by permission from IUCN Small Carnivore Specialist Group, from Jachowski, D.S., Lockhart, J.M., 2009. Reintroducing the Black-
footed Ferret Mustela nigripes to the Great Plains of North America. Small Carnivore Conservation 41, 8: 58–61
a
Combination of captive-born releases and wild-born translocations
b
Minimum number known alive through annual surveys as of December 2008 (unless otherwise noted)
c
Based on monitoring only 15% of habitat
d
Based on 2007 estimate because 2008 monitoring was not performed
e
Based on monitoring only 45% of habitat
f
Total number of Ferrets released that were not removed for translocation to other sites during the same year
complained that, in their state, ferrets receive too much pro- but scientists must learn from failure as well as success if the
tection and ranchers too little (Biggins et al. 2006). Such restoration effort is to ultimately establish wild populations.
attitudes are widespread in the western US. (4) True interdisciplinary cooperation involving veterinarians,
Five lessons can be learned from the restoration of the black- field biologists, ecologists, epidemiologists and scientists from
footed ferret. (1) Populations recently considered extinct might other disciplines, as well as broad stakeholder representation
not be. Perseverance in searching for extant populations of from government agencies, non-governmental conservation
recently “extinct” species is sometimes rewarded. (2) A captive organizations and private interests, is essential to restoration.
breeding plan must be fully integrated with the reintroduction (5) Public and political support for reintroducing an
effort, and be prepared to use all available reproductive endangered species are important elements of the recovery
technologies, not only to maximize the number of individuals process (Chap. 12). Without appreciation of the intrinsic
produced, but to retain gene diversity and, as a result, reduce value of an endangered species (Chap. 10) and provision for
effects of inbreeding depression. (3) Restoring populations its needs, restoration efforts will be limited in their effective-
through captive-bred individuals requires persistent ness, especially on private lands where needs of the endangered
introductions at many sites and repeated releases at the same species compete directly with the well-being and economic
site. Such efforts will experience many site-specific failures, interests of landowners.
1000 years. Today the belief that some discrete minimum

viable population can be calculated for individual species has
been largely abandoned in conservation biology, replaced
with projections of the persistence likelihood of a population,
a concept that has been refined with the development of a
technique called population viability analysis, or PVA, in
which analytical or simulation models make precise
estimates of the probability of species persistence within a
defined time period at a given level of probability (i.e.,
uncertainty). PVA models, especially models for individual
species, quantitatively evaluate extinction risks and estimate
of probabilities of stochastic events in populations (Groom
and Pascual 1998).
PVA is a specific application of the broader science and
techniques of risk assessment, molded into a specific appli-
cation for identifying risks to endangered species. As Mark
Shaffer, one of the originators of the MVP concept, explained
nearly 30 years ago, “Like the more usual forms of risk
analysis applied to issues of public health and safety, PVA
attempts to assess the likelihood of future events based on
Fig. 6.24 Sites (28) of re-introductions of black-footed ferrets (Mustela currently available data and theory, both of which contain
nigripes) in the western United States (26 sites), Canada (one site), and
Mexico (one site), 1991–2016. (Map courtesy of Dean Biggins/US some degree of uncertainty” (Shaffer 1981:39). The goal of
Geological Survey) PVA is to identify and evaluate threats to a species and
estimate the probability of that species persisting for a
given time into the future. More precisely, PVA is the esti-
mation of extinction probabilities by analyses that incorpo-
6.5 Population Viability Analysis rate identifiable threats to population survival into models of
the extinction process (Lacy 1993). PVA is, in fact, an
6.5.1 Conceptual Foundations integrated form of the previously discussed methodology of
trend analysis. In PVA, however, the multiple demographic
The critical problem associated with the conservation of variables identified and measured in trend analysis are now
endangered species like the black-footed ferret inevitably used to estimate the probability of a population of a specific
returns to determining their minimum viable population. size persisting for a specified period of time.
Specifically, what is the minimum number of individuals To accomplish, measure, and communicate the goals of
needed in a population to ensure its persistence? Although evaluating extinction risks and estimating population persis-
past attempts to answer this question were rules of thumb tence, PVAs generate “viability measures” that quantify dif-
based on genetic considerations (Franklin 1980), later ferent characteristics of a studied population essential for its
investigators found that greater threats to small populations persistence. The most commonly generated viability measure
lay in problems associated with processes like random varia- is the probability of extinction, but other measures can be
tion in birth rates, death rates, and other demographic generated for other applications (Table 6.8). Today PVAs are
variables (demographic stochasticity) and the effects of ran- applied to a wide range of conservation problems, including
dom environmental variation on the population’s rate of identifying relative importance of factors affecting popula-
increase (environmental stochasticity). tion dynamics and viability, assessing extinction risk,
Such discoveries motivated the search for more precise identifying critical habitats, weighing ecological and socio-
and comprehensive estimates of minimum population economic tradeoffs, prioritizing management alternatives,
thresholds needed for population persistence. Out of this communicating conservation problems to stakeholders, and
need arose the concept of the minimum viable population identifying information gaps to direct further research (Pe’er
(Shaffer 1981), or MVP, as an estimate of the minimum et al. 2013). To do all these things, PVA-based models often
number of individuals needed for the population to survive demand large quantities of data, require thorough understand-
for a given period of time with a specified probability of ing of the life stages of the species of interest, and must be
persistence. One common convention that originally emerged supported by careful population modeling. PVA models
for MVPs was the minimum population size needed to assume that enough is known about the population’s ecology,
achieve a 95% probability of persistence for 100 or dispersal, demography, genetics, and distribution to make an
6.5 Population Viability Analysis 241
Table 6.8 Viability measures Viability measure Number of cases

provided by PVAs generated in
Probability of extinction 39
78 studies covering 82 species
Population size at a given time 19
Time to extinctiona 18
Probability of quasi-extinction 7
Occupancy 7
Probability of decline 6
Growth rate 6
Time to quasi-extinction 5
Minimum viable population (MVP) 5
Minimum area requirement (MAR) 2
Relative population size 2
Expected minimum total abundance 1
Minimum patch number 1
Mean density 1
Mean number of breeding individuals per year per flock 1
Other measures 7
Reprinted by permission from Wiley, from Pe’er, G., Matsinos, Y.G., Johst, K., Franz, K.W., Turlure, C.,
Radchuk, V., Malinowska, A.H., Curtis, J.M. r., Naujokaitis-Lewis, I., Wintle, B.A., Henle, K., 2013. A
Protocol for Better Design, Application, and Communication of Population Viability Analyses. Conservation
Biology 27, 644–656. # 2013 Society for Conservation Biology
a
Including 4 cases reporting the intrinsic mean time to extinction
accurate estimate of the probability of persistence or extinc- reach a level of zero (actual extinction), but rather modelers
tion. This assumption is not always true. Nevertheless, PVA specify a lower limit below which extinction would be nearly
models still attempt to predict patterns of population change certain. This “quasi-extinction threshold” is often built into
over time as well as estimate the probability of population the model as a practical threshold, below which the popula-
persistence under specified conditions. In addition, tion would not be expected to recover.
PVA-based models have value in their capacity to explore The output of a PVA can take different forms. If the quasi-
potential causes of population decline and potential routes to extinction threshold and the time horizon are both allowed to
recovery, estimate the relative strength of different threats to vary, the output takes the form of a three-dimensional “quasi-
population persistence, and discover the importance of extinction surface” in which time to threshold, extinction
neglected aspects of population demography. threshold, and probability of dropping to or below the thresh-
PVA models can be especially helpful when managing old are plotted simultaneously. If time is fixed, the output
small populations common to threatened and endangered spe- produced is a two-dimensional “quasi-extinction curve” in
cies because direct experimental manipulation would have low which different numerical thresholds are plotted against differ-
statistical power (insufficient numbers of individuals and ent quasi-extinction probabilities. Or, if the quasi-extinction
groups) and unacceptable risk (high probability of losing threshold is fixed (for example, the manager determines that
some individuals of an already small population). However, the population must never drop below 200 individuals), then
field studies remain a necessary preliminary step in developing the output generated is a probability distribution of different
a PVA because they are essential for an accurate understand- extinction times. This kind of output is used to find the mean
ing of the demography of small populations as well as for their or median value of the probability distribution. A fourth
effective management. Field studies are also needed after the approach to output is to run multiple simulations with defined
initial PVA model has been developed, because models cannot thresholds, defined times, and defined probabilities. Such an
replace field studies and experiments needed to test hypotheses approach produces a series of curves known as “quasi-extinc-
generated by the model itself. tion contours” that reveal, for a given quasi-extinction proba-
PVA models can be broadly classed as either determin- bility, a combination of time and threshold associated with it
istic (model elements are not determined by random pro- (Fig. 6.25) (Groom and Pascual 1998).
cesses) or stochastic (model elements are affected by An “ideal” PVA combines features thus far described,
random processes). Stochastic models are harder to build beginning with an accurate identification of limiting factors
and interpret, but more accurately reflect population behavior constraining the population (Fig. 6.26). In one way or
because demographic processes are inherently driven by another, the model establishes connections between the pop-
random processes. Many deterministic and stochastic PVAs ulation and its environment and couples those interactions to
are intentionally constructed to not allow a population to their effects on age-specific reproduction and survival rates,
Fig. 6.25 An example of quasi-extinction contours generated from a

Population Viability Analysis (PVA). Each point on a given curved line
(contour) represents a combination of time and population size having a
given probability of quasi-extinction (reduction of the population to
such a low level that extinction is nearly certain) such that all points
on the same line have equal extinction probabilities. (Design by
M. J. Bigelow, based on concepts from Ginzburg et al. 1982)
which are treated as stochastic events. Genetic variation and

effects of inbreeding also can be modeled and coupled to Fig. 6.26 Simplified representation of an “ideal” Population Viability
measures of reproduction and survival. The model’s output is Analysis (PVA) model, including interaction between population
then an estimate of the probability of extinction of the popu- structure and environmental variation to affect demography, population
growth rates, and probability of extinction. The population’s age
lation under different scenarios of environmental, demo-
structure, sex ratio, behavioral interactions, distribution, physiological
graphic, and genetic events (Ralls et al. 2002). status, and age specific birth and death rates are modeled as components
of population structure. Arrows within boxes indicate increase or
decrease. (Reprinted by permission from the American Association
for the Advancement of Science, from Soulé, M.E., Mills, L.S., 1998.
6.5.2 Developing a Conservation PVA– The
No Need to Isolate Genetics. Science 282, 1658–1659. # 1998
Western Prairie Fringed Orchid AAAS)
6.5.2.1 General Considerations

Many species of plants and animals have life cycles with To understand how a stage-based PVA can be
distinct ages, life history stages, or sizes that influence popu- constructed, we will use an approach developed by popula-
lation growth. In some cases, reproduction occurs only when tion biologist Carolyn Sieg of the US Forest Service and her
organisms reach a certain age. The same may be true for colleagues in building a PVA model for the western prairie
mortality. In other species, demographic events are structured fringed orchid (Platanthera praeclara), a threatened species
by body size classes or distinct growth forms or life “stages” in the US in which long-term census and demographic data
with unique, stage-specific demographic rates. Further, the permit development of an accurate and detailed species PVA
probability of remaining in the same stage (stasis) or trans- (Sieg and King 1995; Samson 2002). Individuals pass
ferring to another stage strongly affects the observed pattern through and among different life stages which will be defined
of population growth. A generalized way of understanding quantitatively in the model (Sieg et al. 2003a, b, c, d). The
such stage-based transitions can be seen in Fig. 6.27, where orchid is a wetland species once common in the US west of
all of life history patterns are shown as variations on a the Mississippi River in the tallgrass prairie biome (Fig. 6.28)
common theme. To attempt population viability analysis of (USFWS 1996). With settlement of this region, more than
organisms with this pattern of life history, we must create a 80% of native prairie was converted to cropland or otherwise
model in which age-, size-, sex- or stage-specific demogra- developed (Klopatek et al. 1979), and many of the region’s
phy and transition rates are known and properly related to one wetlands were drained or altered (Dahl 1990). As a result, the
another. This requires the construction of a “stage-based” orchid disappeared from nearly 75% of its known range, and
model. in 1989 was listed as a threatened species (USFWS 1989).
3-lobed and fringed, hence the orchid’s common name

(Fig. 6.29b). When successful pollination occurs, flowering
plants may produce thousands of dust-like seeds (Hof et al.
1999). The next growing season, the seeds develop into an
underground structure called a protocorm which relies on
mycorrhizae for its sustenance (Fig. 6.29c). In time, the
protocorm develops into a seedling and, as the plant emerges
from the ground, begins to photosynthesize. Germination,
protocorm development and transition to a seedling can
occur within a single growing season.
6.5.2.2 A Stage-Based Deterministic Model

A Lefkovitch, or stage-based model (Lefkovitch 1965) can be
used to understand the fringed orchids development stages
and has been applied in PVA’s for conservation of many
species of plants, as well animals, especially fishes and
invertebrates, whose demographic rates are better related
with developmental stage than age. The first step in develop-
ing a stage-based model is to identify the life-history stages of
the species and the pathways of transition among them.
Transition probabilities can then be calculated from field
data collected over several years. Matrix algebra is used to
calculate several useful statistics, such as lambda (λ), the
discrete time rate of increase. For organisms with yearly
Fig. 6.27 (a) Age-structured life cycle of a species with transitions cycles, λ is the ratio of the population in year 2 to the
(probabilities of survival) from one age to the next with return transitions
population in year 1 (λ ¼ N2/N1). A population in which
to age 1, which are reproductive contributions; ω is the final age class
and no individuals survive past this age. (b) Life cycle of an insect; the estimated λ ¼ 1.0 is stable. When λ is >1.0, the popula-
during a given time period, individuals may remain in a stage or transfer tion is increasing. When λ is <1.0, the population is declin-
to the next stage. (c) Life-cycle of a species whose demographic events ing. The value of λ provides a measure of the rate of increase
are structured by size classes showing some of the possible transfers
or decline. For example, a population with λ ¼ 1.12 is
including shrinking to a smaller size and skipping a size class.
(Reprinted by permission from Elsevier, from Ebert, T.A., 1999. Plant growing at a rate of 12% per year, whereas a population
and animal populations: methods in demography. Academic Press, San with λ ¼ 0.97 is decreasing at a rate of 3% per year. The
Diego) discrete growth rate (λ) and the continuous growth rate (r,
used earlier in the chapter) are the most common measures of
population growth. Their conversions are r ¼ ln(λ) and
Only a portion of the area where orchid populations persist
λ ¼ er, provided that λ is measured over an interval where
has been managed for orchid protection. Multiple land use
growth has occurred.
activities, including surface water and groundwater diver-
sion, livestock grazing, prescribed burning, production of
6.5.2.3 Constructing the Model and Matrices
hay, and wetland drainage are common practices that are
expected to continue where the orchid occurs. Some land
Life-History Stages and their Parameters
management activities may be beneficial to the orchid by
removing competing vegetation. Others, such as wetland
drainage, are not. An accurate understanding of orchid’s life
history is therefore essential to exploring possible future Vegetative
Seeds Plant
population trajectories, as well as evaluating relative
differences among effects that land use activities might
have on population persistence.
The life history of the orchid includes two aboveground
stages. Vegetative plants are short (<15 cm) and have only Flowering Plant
one or two leaves (Fig. 6.29a). Flowering plants grow up to
1.2 m tall, producing a branched flowering stalk with numer- The generalized life stages of an annual plant might include
ous cream-colored flowers in which the lower petal is deeply vegetative plant, flowering plant, and seeds, which could be
Fig. 6.28 Historic distribution of the western prairie fringed orchid (solid dots) of greater than 3000 plants each. (Map courtesy of US
(Platanthera praeclara) in the central United States and southern Fish and Wildlife Service, public domain)
Canada (green area), and the 3 largest remaining metapopulations
displayed as above. In the box below you can see how one Suppose we have data from 650 vegetative plants that
could arrange probabilities for within-stage transitions. Here were marked and monitored in the field for 3 years. Of
P values represent within-stage transition, or the probability these, 320 flowered. Now suppose we measured seed produc-
that the plant will remain in the same stage. G values repre- tion of 50 flowering plants, and over the same 3 years,
sent transition probabilities from one stage to another, and average seed production per plant was 23, of which 45%
F is the number of seeds produced by a flowering plant. were viable. We might not have data on germination rates
Subscripts represent the column (first number) and row (sec- of seeds, but let us assume that 10% germinate, and 20% of
ond number) and transitions reflect the order of the stages in the seeds remain viable in the soil seedbank. Using these
the population vector (seed, vegetative plant, flowering plant data, we could calculate values for our transition matrix,
in the example below). Thus, P11 is the probability that a seed replacing general expressions previously used with real
will remain viable in the soil. G12 is the probability that a seed probabilities. Some cells might have no values because
will germinate and become a vegetative plant, G23 is the some transitions never occur (for example, vegetative plants
probability that a vegetative plant will flower. F3 is the do not revert to seeds). But, where transitions do occur, the
number of seeds a flowering plant produces. Let’s show transition matrix would look like this.
these in a matrix summarizing life stages and transitions.
Present life stage
Present life stage Next life stage Seeds Vegetative plant Flowering plant
Next life stage Seeds Vegetative plant Flowering plant Seeds (0.2) – 23(0.45) ¼ 10.35
Seeds P11 – F3 Vegetative plant (0.1) – –
Vegetative plant G12 – – Flowering plant – 320/650 ¼0 .49 –
Flowering plant – G23 –
Fig. 6.29 Life stages of the

western prairie fringed orchid
(Platanthera praeclara) including
vegetative stage (a), reproductive
flowering adult (b), and
protocorm and seedling stage (c).
(Photos courtesy of C. Hull Sieg/
USFS (a and b) and V. J. Nicholas
(c))
Fig. 6.30 Visual representation

of the life-history stages and
transitions of the western prairie
fringed orchid (Platanthera
praeclara). P variables represent
probabilities of remaining in the
same stage (stasis probabilities),
G variables represent transition
rates between life stages, and F5
represents the fecundity rate (seed
production) of flowering plants.
(Concept by F. Van Dyke,
drawing by K. DeVoss)
Constructing Transition Matrices for the Western stages of the western prairie fringed orchid, drawing arrows
Prairie Fringed Orchid on the diagram to indicate transitions. It would look like
To identify all the transitions in the life history of the orchid, Fig. 6.30. The protocorm/seedling stages are combined into
additional information is needed. The western prairie fringed one matrix element because growth from protocorm to seed-
orchid is a perennial plant that may persist for several years in ling can occur within one growing season.
some locations. A vegetative plant in year 1 may remain Using the diagram as a guide, we could construct the
vegetative in year 2, become a flowering plant, or disappear. complete transition matrix. For example, the probability
A flowering plant in year 1 may be vegetative or flower in that a seedling in 1 year becomes a vegetative plant in the
year 2, or it may disappear. Plants that disappear may be next year would be designated as G23. As already noted,
dormant, or they may be dead. One could now construct a some cells in the matrix will have no values because some
diagram similar to the one displayed earlier showing the life transitions never occur. From this generalized format, we
could now add real, empirical data gathered from in situ Sieg and King did not have data on seed viability in the
studies. To complete the transition matrix, we would need soil, but arbitrarily assumed that 50% of the seeds produced
detailed data on both aboveground and belowground remain viable. The resulting matrix is:
transitions. For aboveground data, we will use information
Status this year
collected over 5 years on 16 sites on the Sheyenne National
Status next
Grassland in southeastern North Dakota, USA (adapted from year Seeds Seedling Vegetative Dormant Flowering
Sieg and King 1995). Beginning in 1990, vegetative and Seeds 0.5 – – – F5 ¼ 13,749
flowering plants in 16 belt transects were permanently Seedling 0.0015 0.00 – – –
marked and numbered. Each year the status of previously Vegetative – 0.0301 0.2806 0.0815 0.2106
marked plants was recorded, and new plants were marked. Dormant – – 0.5783 0.1015 0.6968
With this data, we can calculate the transition probabilities Flowering – 0.0099 0.1411 0.0299 0.1025
for a vegetative plant becoming a flowering plant in the
following year and calculate the overall average for the We have worked through this example slowly to allow
4 years. you to re-create the intellectual process through which a
population viability analysis could actually begin to be cre-
Number of Number of Probability of a ated, beginning with determination or estimation of all the
vegetative vegetative plants vegetative plant
transition probabilities associated with the life history of the
plants in year that flowered in year flowering in year
one two two species. There are many ways of creating a PVA, but conser-
1990–1991 74 0 0.00 vation biologists who develop PVAs or attempt to interpret
1991–1992 54 10 0.185 their results need a thorough understanding of this process for
1992–1993 154 53 0.344 three reasons. First, in order to conceive of how to do empir-
1993–1994 361 12 0.033 ical research on populations that will yield the kind of data
Average 0.141 ¼ G35 needed for population viability analysis. Second, to know
how to determine transition probabilities for a population
Estimating Fruit Set model from empirical data. And third, to be able to under-
Sieg and King estimated fruit set by permanently marking stand and interpret the results of any PVA, such as one might
635 flowering plants between 1995 and 1998 (Sieg and King find in a scientific journal.
1995). At the end of each growing season, they recorded the
number of plants that produced viable fruits and the number
of viable fruits per plant. If we assumed that each flowering 6.5.3 Incorporating Stochasticity
plant produced an average of 1.2 fruits per plant and each
fruit produced an average of 21,618 seeds, of which an In deterministic models, transition probabilities are held con-
average of 53% were viable (Hof et al. 1999), we could stant throughout the projected time period and populations
calculate the average number of viable seeds produced per are allowed to increase without bounds. Deterministic models
flowering plant as (1.2) (21,618) (.53) ¼13,749 ¼ F5, and we are mathematically convenient, but biologically unrealistic.
could now complete the matrix. The other aboveground data Because most biological processes are stochastic, stochastic
could be calculated similarly. Sieg and King also estimated models, properly constructed, more accurately represent real
the probability of seedlings becoming vegetative and population dynamics.
flowering plants, based on appearances of new plants. To incorporate the array of variation in nature that can
Using data from seed packets buried in the ground and influence population growth, stochastic models contain
retrieved the next year, they approximated transition elements that deterministic models do not. For example,
probabilities for seed germination rates and development stochastic models can incorporate the variability inherent in
into protocorms and seedlings (Hof et al. 1999). With this annual rates of reproduction and death (demographic
information, we can now complete the matrix. stochasticity) as well as climatic changes, such as wet and
Present Status dry years (environmental stochasticity), which in turn influ-
Status next ence demographic rates. Further, stochastic models often
year Seeds Seedling Vegetative Dormant Flowering incorporate the concept that habitats have a maximum carry-
Seeds P11 – – – F5 ¼ 13,749 ing capacity – that is, there is a limit to the number of
– – –
Seedling G12 P22
individuals that they can support. A “population ceiling”
Vegetative – 0.0301 0.2806 0.0815 0.2106
may be invoked in the model to designate an upper numerical
Dormant – – 0.5783 0.1015 0.6968
limit beyond which the population cannot grow (Burgman
Flowering – 0.0099 0.1411 0.0299 0.1025
et al. 1993). Stochastic models can also incorporate
probabilities for uncommon environmental events
(catastrophes) that may have disproportionately severe 6.5.5 Applications for Animal Populations –
effects on population numbers. In the western prairie fringed Bonelli’s Eagle in Western Europe
orchid, we could incorporate standard deviations associated
with aboveground demographic transitions, allowing us to We introduced the procedure for building a stage-based
place confidence intervals around our population projections. matrix model to develop a PVA with the example of a plant
In addition, we could impose a ceiling on the maximum because plant growth and development, in most species,
population of flowering orchids and use 1000 iterations of offers clear differences in stages of life history and clear
the model so that an average percentage of population boundaries for transitions. Although not always as obvious,
projections of <50 individuals could be calculated. Assuming animal development, and the effect of transition rates to
that populations of less than 50 individuals have a low different stages of development, can also be understood in
probability of persisting, this estimate would provide us the same way. For example, Antonio Hernández-Matías at
with a measure of the probability of “quasi-extinction,” a the University of Barcelona (Spain) and his colleagues devel-
valuable concept discussed earlier representing a level oped a PVA for Bonelli’s eagle (Aquila fasciata) (Fig. 6.31)
below which we would not want the population to drop based on the eagle’s life stages, with calculation of appropri-
(Groom and Pascual 1998). Finally, to make the model ate transition rates, shown in Fig. 6.32. Developing a PVA
even more realistic, we could incorporate the occurrence of with accurate understanding of the eagle’s life history, com-
three climatic scenarios (average, wet and dry) to project bined with 20 years of demographic data from 12 populations,
population growth rate and persistence under different envi- Hernández-Matías and his colleagues gained insights critical
ronmental conditions. Outcomes would differ in each case to the eagle’s current status and future conservation. First,
(Table 6.9). their PVA revealed that a single large population on the
Iberian Peninsula was the primary source for other
populations which had lower rates of survival or natality, a
6.5.4 Evaluating Elasticity classic source-sink population configuration described earlier
(Sect. 6.2.2). This means that, in prioritizing conservation of
Understanding how to translate PVA results into meaningful populations, the source population was most critical for the
management actions requires that we know which variables eagle’s persistence in this region. Second, elasticity analyses
most affect the viability of the population and, if possible, revealed that eagle population growth was most sensitive to
why. Large changes in some variables might have little or no adult survival (Fig. 6.33). Therefore, managers should priori-
effect on population persistence, while small changes in other tize actions that will increase adult survival rates. This means,
variables might have large effects on population persistence. initially, that, to conserve populations of eagles over the
We can determine which variables have the greatest impact entire area, managers should identify populations with the
on model outcomes by conducting an elasticity analysis, a highest levels of adult mortality, analyze the causes of mor-
specific type of sensitivity analysis that determines the effect tality specific to those populations, and then implement man-
of proportional changes in each variable on model outcomes. agement actions that decrease such mortality and improve
The variable or variables that have the greatest proportional adult survivorship in these groups (Hernández-Matías et al.
effect on λ (i.e., those with the greatest elasticity) are those 2013).
that should be most intensely managed.
Elasticity analyses can provide insights on identifying
potential management strategies for threatened populations 6.5.6 Evaluating a PVA
because they can identify which matrix element to change
that could provide the quickest route to population recovery. A good PVA, regardless of species or approach, will have
They are also vital when the value of key model parameters is certain consistent characteristics, and so do good uses of a
uncertain because they tell the modeler how much such PVA. The most important are that (1) the PVA is informed by
uncertainty matters. If we find variables with high elasticities real data on the species of interest, ideally collected over an
that are also tractable to management actions, we can then extended time period in a variety of conditions;
focus on changing these variables in ways that increase the (2) assumptions of the PVA are explicit, clear, and reasonable
likelihood of population persistence. In contrast, if we find relative to what is known about the species; (3) initial perfor-
variables with high elasticities but have no idea why they are mance (model “runs”) of the PVA are subject to sensitivity
affecting population persistence as strongly as they are, we analyses of model variables; (4) the PVA identifies and
must organize future research efforts around examining these addresses uncertainties; (5) modelers reporting results of the
variables to gain a better understanding of their role in the PVA identify parameters of the model that can be
population’s growth. manipulated directly by management actions; and (6) the
248
Table 6.9 Output of a PVA of the western prairie fringed orchid (Platanthera praeclara), showing changes in numbers of individuals over time under average, wet, and dry climate conditions
Average Wet Dry
Percent of iterations Percent of iterations Percent of iterations
Time Flowering Flowering below 50 flowering Time Flowering Flowering below 50 flowering Time Flowering Flowering below 50 flowering
step plants plants λ plants step plants plants λ plants step plants plants λ plants
0 250 0 250 0 250
1 158 0.63 0 1 382 1.53 0 1 20 8.08E-02 100
2 98 0.62 0.1 2 313 0.82 0 2 10 0.49 100
3 111 1.13 0 3 310 0.99 0 3 39 3.93 100
4 121 1.09 0 4 413 1.33 0 4 22 0.57 100
5 116 0.95 0 5 549 1.33 0 5 13 0.56 100
6 110 0.95 0 6 664 1.21 0 6 12 0.92 100
7 108 0.98 0 7 780 1.18 0 7 9 0.76 100
8 106 0.98 0 8 934 1.20 0 8 6 0.69 100
9 103 0.97 0 9 1136 1.22 0 9 5 0.76 100
10 100 0.97 0 10 1382 1.22 0 10 4 0.76 100
6
11 98 0.98 0 11 1674 1.21 0 11 3 0.74 100

12 95 0.97 0 12 2014 1.20 0 12 2 0.74 100
13 93 0.97 0 13 2424 1.20 0 13 1 0.75 100
14 91 0.98 0 14 2933 1.21 0 14 1 0.75 100
15 88 0.98 0 15 3562 1.21 0 15 1 0.74 100
16 86 0.98 0 16 4310 1.21 0 16 1 0.75 100
17 84 0.98 0 17 5226 1.21 0 17 1 0.75 100
18 82 0.97 0.1 18 6283 1.20 0 18 1 0.75 100
19 80 0.98 0.1 19 7614 1.21 0 19 1 0.75 100
20 78 0.97 0.7 20 9198 1.21 0 20 1 0.75 100
21 76 0.97 0.7 21 11,037 1.20 0 21 1 0.75 100
22 74 0.98 1.5 22 13,359 1.21 0 22 1 0.75 100
Table format by M. J. Bigelow
The Conservation of Populations: Theory, Analysis, Application
6.6 Applying PVA Results in Conservation Management 249
PVA evaluates and ranks outcomes of different management should, ideally, also help to prescribe real population levels
scenarios based on their effects on one or more viability necessary for species persistence, which then serve as
measures of the species; and (7) Modelers reporting output guidelines for recovery criteria for that species. Is PVA
of the PVA tie such output directly to recommendations for being used effectively for this purpose?
management action.
It is not easy to predict extinctions, but that is what PVA is
supposed to do. Obstacles to accurate predictions arise from 6.6 Applying PVA Results in Conservation
two sources. First, information may be inadequate, especially Management
for endangered species about which little is known. Second,
characteristics of the model may create their own problems. Today PVA is widely used and applied for animal
The model must often make long-term predictions using populations, for example in IUCN’s Species Survival Plans
short-term databases. Population change will unfold in a for endangered animal species. PVA remains under utilized in
dynamic environment, but most PVAs do not incorporate the conservation of endangered plants. Ziegler et al. (2013),
considerations of such change. A good PVA must inform for example, examined 258 recovery plans for 642 plants
both absolute and relative recommendations for conservation listed under the US ESA and found that only 154 species
management. That is, relative recommendations may only be (24%) had recovery plans that used or recommended PVA.
able to rank relative outcomes of different management Even in plans incorporating PVA, most did not conduct a
scenarios, but using absolute recommendations, a PVA PVA prior to or during the writing of the recovery plan, and
most did not use PVA to set recovery criteria (Fig. 6.34).
Most PVAs also failed to incorporate stochastic variation,
genetics, or density dependence, and most had less than
5 years of data from which to make their long-term future
projections (Ziegler et al. 2013).
Even when PVAs are developed early in a recovery effort,
problems remain, and fall into three categories. These are
problems in design of the PVA, problems in application of
the PVA, and problems in communicating results of the
PVA. To address these problems, Guy Pe’er of Germany’s
Helmholtz Centre for Environmental Research and his
colleagues designed a protocol for creating a “good” PVA
by identifying essential steps in both the design of the under-
lying population model and essential elements of the
associated PVA (Table 6.10). If all protocols are followed
Fig. 6.31 Bonelli’s eagle (Aquila fasciata), a species whose regional and all recommended elements included, the resulting PVA
populations in western Europe are of conservation concern. (Photo
courtesy of A. Marmasse) will have greater value in conserving the targeted population.
Fig. 6.32 Life stages of Bonelli’s eagle (Aquila fasciata) as conceived by permission from Wiley, from Hernández-Matías, A., Real, J.,
by Hernández-Matías et al. (2013) in creating a stage-based PVA to Moleón, M., Palma, L., Sánchez-Zapata, J.A., Pradel, R., Carrete, M.,
serve as a basis for the eagle’s conservation. Solid lines represent Gil-Sánchez, J.M., Beja, P., Balbontín, J., Vincent-Martin, N.,
transitions between age classes while dashed lines represent transitions Ravayrol, A., Benítez, J.R., Arroyo, B., Fernández, C., Ferreiro, E.,
between age classes through additions via reproduction. SR represents García, J., 2013. From local monitoring to a broad-scale viability
sex ratio, S represents survival rate, and F represents fertility for specific assessment: a case study for the Bonelli’s Eagle in western Europe.
age classes. Numbers after S or F reflect age in year(s) of eagles in that Ecological Monographs 83, 239–261. # 2013 Ecological Society of
age class. A represents adult birds 5 years of age or older. R is the America)
proportion of birds establishing territories as first year adults. (Reprinted
Fig. 6.33 (a) Sensitivity and elasticity analyses of PVA of Bonelli’s growth rates. (Reprinted by permission from Wiley, from Hernández-
eagle (Aquiloa fasciata) where SA represents adult survival, S123 Matías, A., Real, J., Moleón, M., Palma, L., Sánchez-Zapata, J.A.,
subadult survival, FA adult fertility, and K carrying capacity for a Pradel, R., Carrete, M., Gil-Sánchez, J.M., Beja, P., Balbontín, J.,
local population. (b) Absolute changes in population growth rates Vincent-Martin, N., Ravayrol, A., Benítez, J.R., Arroyo, B., Fernández,
(lambda) in relative to relative changes in vital rates. SA ¼ solid line, C., Ferreiro, E., García, J., 2013. From local monitoring to a broad-scale
S123 ¼ dash-dotted line, FA ¼ dotted line and K ¼ dashed line. Note viability assessment: a case study for the Bonelli’s Eagle in western
the steep slope of the SA line, indicating that small changes in adult Europe. Ecological Monographs 83, 239–261. # 2013 Ecological
survival rates cause disproportionately large changes in population Society of America)
Fig. 6.34 Context under which

population viability analysis
(PVA) is used or recommended in
final recovery plans for 642 plant
species listed under the
US Endangered Species Act. Note
how little PVA was used in early
stages of recovery planning or in
actual recovery criteria in these
recovery plans. (Reprinted by
from Zeigler, S.L., Che-Castaldo,
J.P., Neel, M.C., 2013. Actual and
Potential Use of Population
Viability Analyses in Recovery of
Plant Species Listed under the
US Endangered Species Act.
27, 1265–1278. # 2013 Society
for Conservation Biology)
6.7 From Population Viability Analysis to Population Viability Management 251
Table 6.10 Elements of a good population model (after Schmolke et al. 2010) and elements of a useful PVA delineated along the steps of
model design, application and communication
Elements of a good model Elements of a good PVA
During design Includes stakeholders Includes stakeholders in model design, validation, and interpretation
Builds on (long-term) high-quality data
Formulates objectives; justifies choice of Performs and justifies a careful model selection
model approach & complexity
During Careful parameterization Includes relevant parameters based on knowledge of the system and the
application literature and in consideration of gaps
Applies careful parameter estimation and parameterization
Calibration, verification, validation Performs calibration, verification, and validation or directs further
monitoring and validation efforts
Quantification of uncertainties Performs sensitivity analyses and addresses uncertainty in a systematic and
transparent way
Applies multiple models Compares the outcomes of alternative models where possible
Differentiates among parameters affecting the model, the real world, and
those that are management relevant
Ranks management scenarios to support decision making
During Formulates assumptions Communicates the entire modeling cycle and justifies decisions and
communication assumptions along the way
Effective documentation and transparency Reports all inputs and outputs systematically to allow repeatability
Uses carefully selected time horizon and viability measures and reports
using consistent units to allow comparability
Demonstrates that the PVA serves its purpose by, e.g., leading to on-the-
ground actions
Enhances collective learning and potential generalization
Peer reviewed Both the model (design, code, application) and the report are peer reviewed
Reprinted by permission from Wiley, from Pe’er, G., Matsinos, Y.G., Johst, K., Franz, K.W., Turlure, C., Radchuk, V., Malinowska, A.H., Curtis, J.
M., Naujokaitis-Lewis, I., Wintle, B.A., Henle, K., 2013. A Protocol for Better Design, Application, and Communication of Population Viability
Analyses. Conservation Biology 27, 644-656. # 2013 Society for Conservation Biology
evaluates expected measurement error in relation to intensity

6.7 From Population Viability Analysis of monitoring effort associated with each approach. PVM
to Population Viability Management then ties monitoring data to population viability by simulta-
neously following the effectiveness of the monitoring system
Some critics of PVA go beyond advocacy for revision, like and the species’ population dynamics.
Ziegler and Pe’er, to advocacy for alternatives. One of these Bakker and Doak (2009) used a PVM approach to formu-
is Population Viability Management (PVM). PVM attempts late a recovery plan for the endangered island fox (Urocyon
to create stronger links between model results and actual littoralis), a species found only on the Channel Islands off the
species recovery than traditional PVA models by placing (US) California coast (Fig. 6.36). Originally threatened
greater stress on ongoing population monitoring (determin- because of its restricted habitat, the island fox has more
ing how much and what kind of information is necessary) and recently faced mounting threats from predation by golden
on effects of management actions (What is the full range of eagles (Aquila chrysaetos). Managers developed a PVA that
possible actions a manager could take toward a population, identified adult survival as the most important demographic
and what effect would they have?). The overall approach of variable to population persistence, as well as determining a
PVM and its incorporation of the results of PVA is illustrated maximum allowable adult mortality rate of 20% for
in Fig. 6.35. Specifically, PVM incorporates monitoring and maintaining stable populations (Coonan 2003). However, a
management directly into PVA, using extinction risk (proba- PVA cannot state how to translate insights about demography
bility of extinction) of each management option as a measure into specific management actions. Seeing the need to do so,
of its conservation value. PVM models also estimate the Baker and Doak built their PVM analysis around evaluation
degree of uncertainty of outcome associated with different and prediction of extinction risk under different management
management actions, their efficacy at reducing specific strategies.
impacts on a population, and the range of conditions under Baker and Doak first defined an acceptable recovery stan-
which management actions could be implemented. In addi- dard of adult population size and rates of adult mortality
tion, PVM simulates different kinds and degrees of monitor- related to different levels of extinction risk. Monitoring
ing systems that inform the management decisions, and then methods were adapted to emphasize getting estimates of
Fig. 6.35 A comparison of a Population Viability Analysis (PVA)

Population Viability Analysis
(PVA) (a) to Population Viability Generalized
Management (PVM) (b). Management management
Adaptive management decisions decisions and recommendations
(blue) based on monitoring data Management actions
(green) are aimed to reduce threats efficacy One-time PVA
to at-risk populations and ensure Predicted extinction
their long-term viability (red). risk
PVA (yellow) has often been a Threats
single, one-time exercise yielding
generalized recommendations
without specific accounting for Monitoring
Survival and
uncertainties in data and
reproduction data
generating management decisions Demographic
based on trends in species’ rate
abundance or demographic One-time
monitoring Observation
parameters. (Reprinted by error monitoring
Abundance
permission from Wiley, planning
Abundance
from Bakker, V.J., Doak, D.F.,
monitoring
2009. Population viability
management: ecological standards
to guide adaptive management for b Population Viability Management (PVM)
rare species. Frontiers in Ecology
and the Environment 7, 158–165. Management
# 2009 Ecological Society of decisions and
America) actions
P(ext) = Start/stop management?
Management Management P(ext) = Change monitoring?
efficacy
Planning
Iterative PVA
Predicted extinction risk
given monitoring data, observation
Threats error, management and its efficacy
Monitoring
Survival and planning Monitoring
reproduction Demographic data
rate
monitoring Observation
Abundance error
Abundance
monitoring
these two variables. Monitoring frequency was structured to capture multiple eagles at once, rather than individual, trap-
produce the most reliable estimates, but adaptable to changes based capture methods (Bakker and Doak 2009). In this
in management priorities and budgets over time. Bakker and effort, a PVM approach created more functional relationships
Doak then related model elements associated with fox popu- between researchers and managers and better connected
lation size and mortality to different approaches to managing model results to management actions.
eagles, which in this case meant evaluating effort required to
capture and remove eagles from islands after eagle predation
on foxes was detected. Practically, this meant modeling 6.8 The Problem of Recovery: Protecting
changes in numbers and mortality rates of foxes in the pres- Conservation – Reliant Species
ence of different numbers of eagles. Such modeling revealed
that the level of management (removal) of eagles necessary to 6.8.1 What Is a Conservation–Reliant Species?
lower extinction probability of the island fox to less than
50 percent could only be achieved if the current removal The goal of managing endangered populations is their recov-
rate of eagles was doubled. Managers responded by not ery. That is why management plans for endangered species
only increasing intensity of removal, but employing more are called “recovery plans.” In the United States, the goal of
efficient methods, such as using net gunning, which could the US Endangered Species Act (ESA) is to bring listed
6.8 The Problem of Recovery: Protecting Conservation – Reliant Species 253
Fig. 6.37 The Kirtland’s warbler (Setophaga kirtlandii), an exemplary

representative of the increasing number of “conservation-reliant spe-
cies” that require continued management effort, even after recovery, to
avoid returning to endangered status. (Photo courtesy of J. Trick and the
US Fish and Wildlife Service, public domain)
6.8.2 The Kirtland’s Warbler: A “Success Story”

Fig. 6.36 The island fox (Urocyon littoralis), found only on one group
of islands, the Channel Islands, off the California coast of the United
of Conservation-Reliance
States. The island fox has been the subject of an intensive effort in
Population Viability Management due to its decline as a result of 6.8.2.1 Three-Dimensional Rarity: Endemism,
predation by the golden eagle (Aquila chrysaetos). (Photo courtesy of Specialization, and Low Populations
S.F. Lamb)
As we noted in our examination of biodiversity (Chap. 2), the
Kirtland’s warbler (Setophaga kirtlandii) (Fig. 6.37) can be
considered rare in each of the three categories of rarity. Until
recently, it has been an example of extreme endemism,
breeding in only five counties in northern lower Michigan
species to “the point at which the measures provided [by the] (USA), as well as an example of extreme habitat specializa-
Act are no longer necessary” (ESA § 3 (4); Chap. 12). tion (beta rarity), using only homogeneous stands of 5- to
Although there have been failures (eleven species have 20-year old jack pine (Pinus banksiana). It also exemplifies
become extinct under the “protection” of the ESA), a review demographic rarity, with its population reduced to less than
of the ESA’s success through March 2020 found that 99% of 500 adults by the late 1970s.
listed species still survived, 42 formerly endangered species Conservation measures used in the warbler’s recovery
had been reclassified as threatened and were moving toward plan employed a five-part strategy of (1) creating more suit-
recovery, and 6 more species were under active considerable habitat (young, even-aged jack pine), (2) increasing
ation for reclassification (USFWS 2020). In spite of this recruitment of warblers by controlling an avian nest parasite,
record of effective conservation effort, only 58 of the the brown-headed cowbird (Molothrus ater), (3) monitoring
2532 ESA-listed species have been designated as recovered populations, (4) conducting research, and (5) developing a
and subsequently removed (delisted) from the Endangered program of public education about the Kirtland’s warbler and
Species List. The low rate of delisting is due to the fact that, its needs. The first two elements were the most specific and
even when the population of an endangered species actionable. Thousands of hectares of young jack pine habitat
increases, most listed species continue to depend on ongoing was created within the Kirtland’s historic range through
management action to maintain their trajectory toward recov- planting and prescribed burning. Aided in recent years by
ery. Such cases are examples of conservation reliant species, large natural fires occurring in breeding areas, the amount of
a descriptor that can be accurately applied to more than 80% suitable habitat increased dramatically. The second element
of species protected under the US ESA (Scott et al. 2010). To was accomplished by trapping and removing (killing)
understand the problems associated with conservation-reliant cowbirds in warbler breeding areas.
species, and their potential solutions, we will examine one These efforts contributed to a steady increase in numbers
case exemplifying such conservation “success.” of Kirtland’s warblers, beginning in the 1990’s (Fig. 6.38).
Fig. 6.38 Number of singing male Kirtland’s warblers detected in Conservation Management Agreements to Secure Postrecovery Perpet-
decennial then annual censuses conducted throughout the species’ uation of Conservation-Reliant Species: The Kirtland’s Warbler as a
range from 1981 to 2011. Note that in every year from 2001 on, numbers Case Study. BioScience 62, 874–879 # 2012 American Institute of
exceed recovery goal. (Reprinted by permission from Oxford University Biological Sciences)
Press, from Bocetti, C.I., Goble, D.D., Scott, J.M., 2012. Using
Since 2001, Kirtland’s warbler populations have exceeded Forest Service, with the latter having jurisdiction over many
their recovery target of 1000 adult singing males (determined areas warblers use as breeding habitat), the state government
by an annual breeding season survey) every year. Given that (the Michigan Department of Natural Resources, which has
recovery criteria have been met annually for nearly two taken a leading role in creating new warbler habitat), private,
decades, the US Fish and Wildlife Service proposed delisting non-governmental conservation organizations (NGOs) with
the warbler in April 2018. But with delisting comes loss of vested interests in the recovery of the Kirtland’s warbler (like
legal protection for the species and loss of federal govern- The Audubon Society, as well as smaller regional
ment funds designated for it, funds used for habitat creation organizations, whose staff and volunteers have made
and cowbird removal. The mechanism used to replace both contributions to population monitoring and public educa-
legal protection and funding for the Kirtland’s warbler is that tion), private landowners (who control access to breeding
of a Conservation Management Agreement (CMA). areas) and private industry (especially the wood products
industry, which manages large areas of jack pine forests
6.8.2.2 Conservation Management Agreements which represent potential breeding habitat for the Kirtland’s
for Conservation-Reliant Species warbler). In this case, the KWRT plans to create a nonprofit
The most recent threat analysis for the Kirtland’s warbler conservation organization dedicated to continued manage-
determined that termination of cowbird control and loss of ment of the warbler. The organization will be governed by
habitat creation and management posed the biggest, post- its own Board of Trustees who will work with an advisory
delisting threats to this species’ persistence. The Kirtland’s board of experts from public and private sectors, especially
Warbler Recovery Team (KWRT) also concluded that addi- those who have previously served on the Kirtland’s warbler
tional elements of population monitoring, research, and pub- recovery team. These boards will form the functional part-
lic education would remain essential to maintain warbler nership that will oversee the continuing recovery of the
populations after delisting (Bocetti et al. 2012). The CMA warbler.
for the Kirtland’s warbler, like any CMA, requires four The Partners primary responsibilities will be to oversee a
elements to be effective. These are (1) a partnership agree- management plan developed cooperatively by the US Fish
ment that continues to enable effective management actions; and Wildlife Service, US Forest Service, and Michigan
(2) a management plan informed by understanding critical Department of Natural Resources, the government agencies
elements of the species’ recovery; (3) sufficient funding; and which have been the core players in management efforts to
(4) legal enforcement (Fig. 6.39). this point. The respective roles and responsibilities of the
In this case, the Partners will include the US federal different agencies, formalized under an official Memorandum
government (the US Fish and Wildlife Service and the US of Understanding (MOU), will include specific assignments
6.8 The Problem of Recovery: Protecting Conservation – Reliant Species 255
Fig. 6.39 Schematic

representation of the elements
needed for an effective
Conservation Management
Agreement (CMA) as exemplified
in elements specific for continued
management of a conservation-
reliant species, the Kirtland’s
warbler (Setophaga kirtlandii)
after delisting from the
US Endangered Species List.
(Illustration by K. DeVoss)
related to writing, developing, and executing the new man- become true for more and more species worldwide in the
agement plan. Partner agencies will take leading roles in years ahead.
habitat creation and management, cowbird control, research,
population monitoring, and public education, the five critical
elements of the original recovery plan.
Given the increasing number of formerly endangered
To carry out a management plan, a CMA must have
species that remain conservation-reliant even after
adequate financial resources. Here, the Partners envision the
“recovery” and delisting, should the US Endangered
development of a private trust fund dedicated to conservation
Species Act be amended, or a new law passed, that
of the Kirtland’s warbler. Not all government funding for
would give special consideration and status to species
conservation of the Kirtland’s warbler will end when the
formally classified as “conservation reliant”? What
species is delisted, but the amount of funding will be less.
would be the advantages and disadvantages of this
The trust fund will need contributions from conservation
change?
NGOs, as well as from private industry, to enable manage-
ment actions to be carried out.
Finally, the CMA must have authority to makes its man-
agement plan and management actions legally enforceable.
Synthesis
This legal authority must take the form of an enforceable
Theories and examples presented in this chapter are
contract or, on private land, a conservation easement giving
intended to bring us to the point that we can create a
power to regulate the kinds of use and activity that can take
path toward effective restoration of small and declining
place on lands used as breeding areas by the Kirtland’s
populations. There are no certain solutions for success
warbler. In this case, government management agencies
in this difficult and complex process, but we will risk
involved are expected to develop, agree to and sign legally
offering a logical stepwise approach to evaluating and
binding memoranda of agreement (MOAs), with the US
solving problems associated with population
Forest Service taking the lead in ensuring that such MOAs
restoration.
are enforced.
Step One: Determine the Cause of the Decline –
In October 2019, the Kirtland's warbler was offi-
Populations may decline for many reasons. The cause
cially delisted, formally initiating a post-delisting monitoring
of decline may appear obvious, but the “obvious” cause
plan (USFWS 2019). This case history is limited to a single
might, in fact, have little to do with the observed
species, and burdened with imperfection in present efforts
decline. To determine the real reason for the decline,
and uncertainty about future results. Nevertheless, the
examine available information that compares the pop-
Kirtland’s warbler CMA provides us with one approach that
ulation in its present state to the same population in the
can be used for other species that “recover” from endanger-
past, particularly if records exist that describe the
ment yet remain “reliant” on conservation management to
persist in a human-dominated world, a condition that will (continued)
population when it was larger. Comparisons should analysis. These factors can be examined through PVA
focus on critical variables of interest: simulation models, but also through other kinds of
analyses.
1. Geographic range – What are the differences
between the past and present geographic ranges? 1. Environmental stochasticity – What level of envi-
2. Habitat use – What are the differences between past ronmental variation is present and how does it affect
and present habitat use? population numbers?
3. Competitors, predators, parasites, and disease – 2. Demographic stochasticity – What is the current
Are there differences in the types, species, or status of the population’s demography?
intensities of competitors, predators, parasites, or 3. Genetic constraints – What is the population’s cur-
disease that interact with this population now than rent level of inbreeding and heterozygosity? Does
in the past? the population show signs of inbreeding, such as
4. Environmental conditions – Are environmental deformities, sterility, or abnormal juvenile
conditions for the population today the same as or mortality?
different than those faced by this population in the 4. Susceptibility to natural catastrophes – Is the popu-
past? lation susceptible to any types of natural disasters?
5. Integrative comparisons – If data are available to If so, what kind, and at what frequency and sever-
answer the first four questions, the same data can be ity? Is it feasible to protect the population from such
used to answer more complex questions. Some of disasters?
these will be:
(i) How do distributions of competitors, Step Two: Formulate a Hypothesis About the Cause
predators, parasites, and disease organisms of Population Decline – Even after a careful examina-
compare with present and historic distributions tion of available data, facts alone will not unequivo-
of the population? Are these obvious overlaps cally reveal the cause of the population’s decline, nor
or disjunctions in particular interactions? will they spontaneously arrange themselves into a man-
(ii) How has the availability of preferred habitat agement plan. These elements require human insight.
changed in the population’s geographic range Following an adaptive management approach,
over time? managers, informed by the data they possess, should
(iii) Are changes in range and habitat use, if any, formulate one or more hypotheses about the nature and
associated with changes in environmental causes of the population decline. The hypotheses and
conditions experienced by the population? their predictions should then be incorporated into man-
agement actions executed within careful experimental
In addition to comparisons to historic conditions, designs, such as those used in approaches like trend
an examination of present conditions should deter- analysis and factor resolution (Sect. 6.4.2).
mine the status of two human activities that, if
ongoing, should be stopped if the population is to Step Three: Determine Potential Avenues for Increas-
survive. ing Population Size – Whether experimental manipula-
tion is possible or not, managers will have to try
6. Direct exploitation- Does the population experience something to lead a declining population toward recov-
direct exploitation by humans, legally or illegally? ery. Management actions to increase the size and persis-
7. Habitat destruction – Is the critical habitat of the tence of small populations fall into one of three categories:
population, especially breeding habitat, stable in
quantity and quality? If not, what human activities 1. Intensive ecological and environmental manage-
are creating a net loss of critical habitat for the ment of the species in its natural habitat. Using
species? this strategy, managers depend primarily on natural
reproductive capabilities and adaptations for survi-
Important factors regarding present conditions – vorship of a wild population, but enhance the envi-
Historical data are not always available and, if they ronment in such a way as to maximize favorable
are, there are dimensions of a population’s current environmental conditions, minimize detrimental
status that merit examination independent of historical environmental variation, and optimize population
demography toward maximum growth through If there is no captive population, the decision to start
removing competitors, predators, and parasites; one means that some animals must be taken from a wild
controlling disease; creating favorable habitat; population to do it. Some animals will die from
increasing the quality of available habitat; improv- trapping, handling, or transport. Some will not survive
ing opportunity for migration and movement, and, in captivity. The wild population will become smaller.
in some cases, translocating animals and providing
supplemental resources for them. 3. Removal of all remaining wild individuals to pre-
serve the population in captivity – This approach is
This kind of effort is sometimes formally described a more extreme version of the previous strategy, but
as the optimal niche gestalt because of its focus on has been necessary in many cases, such as that of the
habitat management. An optimal niche gestalt strategy black-footed ferret (Sect. 6.4.3). The goal is to even-
assumes there are structural features of an environment tually release captive-bred individuals back into the
that allow a species to thrive over and above those that wild, but, if this strategy is employed, the restoration
allow it to merely persist (James et al. 2001). Managers process may be lengthy and fraught with setbacks.
should first identify such features by determining
correlations between environmental characteristics Managers must assess which of these strategies car-
and high-density populations or subpopulations, exper- ries greatest potential for recovery. Small populations
imentally test hypotheses about underlying causes that leave little room for error. Managers must examine data
lead to these correlations, and then, informed by the carefully, formulate clear hypotheses regarding causes
results, manage selected sites intensively to favor proof population decline, and make testable predictions
cesses and structures that create features associated that can be evaluated in management actions if they
with these high density populations (James et al. are to be effective in restoring small and declining
2001). This approach is exemplified in the creation of populations. Risk of failure is high, but scientists who
young, even-aged jack pine stands for the endangered lead restoration efforts must assess and choose which
Kirtland’s warbler, a major factor contributing to the risks they will take because management leading to
successful recovery of this species. successful restoration will continue to be one of the
most important tasks of conservation biology.
2. Supplementing wild populations through additions
of captive-reared individuals – To employ this strat-
egy, managers must have or create a captive-bred
population, the offspring of which they release into
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The Conservation of Terrestrial Habitat
and Landscape 7
Conservation is a geographic problem because one of the greatest threats to biodiversity is habitat loss and
fragmentation.
Scott M. Pearson (2002)
Keywords 7.1 A Foundational Understanding

Habitat scale · Habitat complexity · Habitat heterogeneity · of Habitat
Habitat suitability model · Habitat fragmentation and loss ·
Patch dynamics · Edge effects · Neutral landscape models 7.1.1 What Is Habitat?
· Percolation theory · GAP analysis
Habitat can be defined as the physical and biological sur-
roundings of an organism (Bolen and Robinson 1995). More
precisely, habitats are sites having appropriate levels of the
Overview biotic and abiotic features required by a species for survival
In this chapter you will learn: and reproduction (Pearson 2002), or “all locations that pro-
vide resources for an organism’s survival” (Tscharntke et al.
1. Concepts and definitions of habitat and landscape 2012:603). In other words, habitats are arrangements of
and the role of habitat conservation in conservation resources that meet the needs of individual species. Such
biology. definitions can be more fully appreciated if we clarify what
2. Definitions of habitat heterogeneity and patch we mean by habitat structure, which can be defined as the
dynamics. amount, composition, and three-dimensional arrangement of
3. Specific mechanisms through which habitat loss, biotic and abiotic elements where an animal lives as a specific
fragmentation, and isolation threaten biodiversity. time and location (He et al. 2019). Because resources needed
4. Principles of reserve design and their role in habitat for survival and reproduction differ in every species, habitat
conservation. and habitat structure are, properly understood, highly
5. Means of reducing the impacts of human distur- species-specific concepts. Our definition of habitat structure
bance to conserve habitat and habitat-dependent is important because it identifies the three fundamental
populations in non-reserve environments. aspects of any given habitat that we will study and apply
throughout this chapter: (1) h, which refers to the amount of
spatial area of the habitat in relation to the size of the refer-
ence landscape; (2) habitat complexity, which is the relation-

262 7 The Conservation of Terrestrial Habitat and Landscape
ship of the scale to amount of physical components (for threat to wetland-dependent species because some wetland
example, food resources) in the habitat; and (3) habitat het- habitats disappear. Environmental variation can never be
erogeneity, which is the relative abundance of different kinds eliminated in natural environments, but preservation of habi-
of components in the habitat at a given spatial scale (He et al. tat, at landscape, regional, or global scales can reduce adverse
2019). A habitat in which there was only one kind of habitat effects of variation and conserve essential, habitat-specific
component would have zero heterogeneity. resources needed by every species.
Scientists and managers often “name” habitats according Conservationists have long considered physical habitat
to the dominant vegetation present in them, (for example, alteration one of the greatest threats to species and
“sagebrush habitat,” “grassland habitat,” or “forest habitat”). ecosystems worldwide (Soulé 1991; Noss and Cooperrider
However, the vegetation communities used to label a particu- 1994), a view supported by trends in habitat change
lar kind of habitat are not the same as the habitats themselves. documented in the Millennium Ecosystem Assessment
Habitat-related threats to particular species arise from (MEA 2005). Habitat alteration includes physical conversion
naturally-induced changes, such as plant succession, or of habitat to unusable non-habitat (habitat loss), breaking
changes associated with human use and modification of large, contiguous blocks of habitat into smaller patches (hab-
land surfaces that create negative consequences in a species’ itat fragmentation), increasing separation of blocks of habitat
physiology, behavior, or interactions with other species from one another (habitat isolation), and changes in habitat
(Fischer and Lindenmayer 2007). that affect composition, structure, or function (habitat degra-
Habitats have traditionally been conceived as occurring in dation) (Noss et al. 1997).
patches. Patches are contiguous regions of the same kind of
habitat (Pearson 2002) or sites where the habitat conditions
of a species are realized. Patches often exist in networks in 7.1.2 How Do We Measure Habitat Use?
which a collection of spatially distinct patches is connected
by linear elements (structural definition) and linked by a flow 7.1.2.1 Species-Specific Habitat Choices
of individuals from patch to patch (functional definition) We have already stressed that “habitat” is a species-specific
(Opdam 2002:318). If this is what we mean by “habitat,” concept. Examine what that means in a species such as moose
what do we mean by “landscape?” (Alces alces). Faced with an array of habitats in a particular
Landscapes can be defined as large areas (measured at landscape, a mobile species, such as moose, can choose
spatial scales of km2 or higher) that comprise more than one which habitats to use. Figure 7.1 gives a summary of these
type of habitat distributed in numerous patches (Danielson choices, using a sample of radio-collared moose from a larger
1991). If we permit the definition to include human influence, population in the US state of Montana (Van Dyke et al.
a landscape is an aggregate of different but interacting 1995). Habitat classification for this population may use
elements, sometimes united by a cultural [human-induced] arbitrary categories, named, for convenience, by their domi-
attribute (e.g., an agricultural landscape of cultivated fields, nant vegetation type, as noted previously. The categories
pastures, stockponds, and hedgerows) (Bolen and Robinson themselves do not perfectly describe the habitats, but by
1995). These definitions are important because the funda- creating categories we can analyze their distribution in a
mental goal of conservation biology – the preservation of matrix of complex vegetation and then see how animals use
biodiversity – rests largely upon the conservation, preserva- them.
tion, and management of habitat and landscape. Habitat dis- We see in this example that moose, like most species, use
tribution is a primary determinant of species’ abundance, and more than one kind of habitat (Fig. 7.1). This is because a
it is often the degree of habitat specificity that determines single habitat seldom provides all necessary resources for a
whether a species is common or rare (Rabinowitz et al. 1986, species to accomplish various essential activities over its
Chap. 2). Therefore, habitat and landscape considerations are lifespan, including survival and reproduction. Note that habi-
fundamental to all forms of geographic-based approaches to tat use is often different from habitat availability. Some
biodiversity conservation. A geographic-based approach to habitats are used at rates greater than expected, whereas
biodiversity conservation, as opposed to a population-based others are used less than expected relative to their distribution
approach, focuses less on the dynamics of a particular popu- in a landscape. These differences can lead conservationists to
lation and more on the qualities of habitat and landscape that make well-informed hypotheses about why patterns of appar-
sustain it and other populations. The role of habitat and ent habitat preference or avoidance appear in particular spe-
landscape are vital to population persistence because cies. If these hypotheses are tested through careful
populations become more susceptible to extinction in the experimental design and manipulation, the results of the
face of environmental variation to which they are not previ- experiments can provide insight about why animals choose
ously adapted. For example, prolonged drought can pose a habitats, and about the resources particular habitats provide
7.1 A Foundational Understanding of Habitat 263
Fig. 7.1 Seasonal habitat

selection by three male and ten
female moose in the Fiddler and
Fishtail Creek drainages, south-
central Montana, USA,
1989–93. Numbers indicate
percentages. Symbols in
parentheses indicate selection for
(+), selection against (), or no
selection (0). P < 0.01
(simultaneous confidence
intervals, Marcum and
Loftsgaarden 1980) for all cases
of selection and for differences
between sexes, except where
noted. (Figure based on data from
Van Dyke et al. 1995)
to the animals. But first, we need to determine methods proportionately much higher than availability, as in our
through which we can quantify habitat use and detect prefer- example, suggesting preference for that habitat. Alterna-
ence for habitats that might have disproportionately high tively, it will approach a value of negative one when use is
conservation value for an individual species. far less than availability, suggesting avoidance of the habitat.
If many samples of habitat use are made from multiple
7.1.2.2 How Do We Measure Habitat Selection individuals in the same area, standard statistical techniques
and Preference? can be used to see if the average value of the selection index
There are many algorithms that can employ these kinds of is different from zero (Neu et al. 1974; Marcum and
data for making assessments of such differential use of Loftsgaarden 1980).
habitats by individuals and populations. One used for To acquire data on the availability of habitat in an area, the
identifying the strength of selection for habitats is Ivlev’s first step is to determine the target population relative to the
Selection Index (SI) (Ivlev 1961). Originally developed to habitat, because habitat is often, functionally, species-
assess selection for food items, the SI can be effectively specific. Once we are clear about which population or species
applied to measure selection for habitat as well (Yeo and we want to investigate relative to habitat use, we then make
Peek 1992). The SI for a given type of habitat is determined sure that our selection of sampled sites truly represents
as habitats used by that population. One could then use existing
vegetation maps, aerial photos, or other geographic informa-
SI ¼ ðU AÞ=U þ A: tion systems (GIS) data to create a complete picture of the
habitat that is truly “available” to the target population. If
where U is equal to the proportion the animal’s use of none of these sources exist, availability can be determined
the habitat out of all use and A is the proportion of the from sampling numerous random points within the area used
same habitat available in the landscape. For example, sup- by the target population until an appropriate number of
pose a moose spends 80% of all its time in stands of quaking samples have been made. Use of habitats by an animal can
aspen (Populus tremuloides) trees (U ¼ 80) within a home be determined from direct observation, radio telemetry, sys-
range in which quaking aspen stands make up 10% of the tematic trapping, or locations of vocalizations (“songs”),
landscape (A ¼ 10). Then, tracks, or sign (scat or markings). Such data provide a
means to obtain an estimate of habitat use relative to avail-
SI ¼ ð80 10Þ=ð80 þ 10Þ ¼ 70=90 ¼ 0:78: ability, and then using such information to make a more
informed decision about which habitats to preserve for par-
In Ivlev’s formula, SI will be zero whenever the animal ticular species.
uses the habitat in the same proportion as its availability These data also demonstrate the principle that habitat use
(no selection). It will approach the value of 1.0 when use is changes over time (in this example, seasonally). Such
changes should be expected because the availability of 7.1.2.3 Accounting for Spatial Scale in Habitat
resources in habitats also changes. This reality has often Selection
been overlooked in the design of parks and nature reserves, Our perception of habitat preference in a species will change
to the detriment of the reserve’s conservation objectives. If with the scale of examination and measurement. Most
the full array of habitat structure that a species requires in animals that consistently use the same area or “home range”
different seasons does not exist within the reserve, then do not use all parts of that area equally. Most use normally
individuals must leave the reserve to find appropriate habitat, takes place in a “core use” area that may contribute only a
and, in doing so, face many threats the reserve was intended small fraction of the total area within the home range. For
to protect them from, or perish in the reserve without their example, in this population, individual moose, especially
needed habitat. This is a problem that cannot be solved “once females, concentrated their use in areas with optimal foraging
and for all” in reserve design, because habitats, like characteristics. Moose are browsers, feeding primarily on
organisms, have “lifespans” and therefore provision must woody plants, so, as you can see in Table 7.1, it is not
be made to create new habitat as well as to preserve existing surprising that, despite individual differences and use of
habitat. But habitat management and conservation are further spatially independent areas, all of the female core areas had
complicated because habitat use differs among individuals. similar (and high!) levels of shrub coverage in the core area
Sex- or age-related differences in body size, social organiza- plant communities (Van Dyke et al. 1995). We can under-
tion, or investment in reproductive effort and parental care stand the problem better and in greater detail if we consider
can lead individuals of the same population to select different an even more precise examination of scale-specific habitat
habitats during the same period of time. Failure to consider selection in another species, the giant panda (Ailuropoda
these differences can lead to inappropriate choices of which melanoleuca).
habitats to conserve and result in ineffective conservation The giant panda (Fig. 7.2) is one of conservation’s most
efforts. iconic species. It has also proven one of the most challenging
Table 7.1 Plant community characteristics of core foraging areas used by five female moose in south-central Montana (USA), 1989–1992. Two-
letter column headings are designations for different core foraging areas. MC Molly’s Canyon, EF East Fiddler, MB McDonald Basin, LR Little
Rocky, BR Bates Ranch
Core Area
Characteristic MC EF MB LR BR Combined
Plant coverage 71.5a (1.7) 98.1b (1.2) 94.7 (3.0) 90.3 (2.2) 93.3 (1.8) 88.5 (5.1)
(%)
Shrub coverage 53.5b (3.9) 45.3 (3.6) 53.3 (4.2) 42.5 (3.4) 49.1 (2.5) 48.7 (8.6)
(%)
Forb coverage 5.6a (2.1) 24.0 (4.4) 26.0 (2.8) 32.7 (2.9) 24.8b (2.5) 22.3 (7.1)
(%)
Graminoid 12.8c (1.9) 17.0 (4.5) 15.2 (1.9) 14.1 (1.5) 20.5 (2.5) 16.2 (6.2)
coverage (%)
Plant volume (m3/ 3943.2a (515.8) 13838.7d (706.3) 12751.6d (520.5) 6773.2a (433.0) 9773.2a (620.9) 8786.6 (1484.8)
ha)
Shrub volume 3645.9a (576.7) 10981.1d (911.7) 11731.4d (556.0) 5817.3a (430.8) 8132.3a (560.3) 7512.6 (1511.2)
(m3/ha)
Forb volume (m3/ 56.6a (16.6) 643.1 (242.3) 542.2 (87.7) 497.1 (98.4) 755.3 (134.9) 502.9 (316.2)
ha)
Graminoid 191.2e (67.1) 866.7 (307.7) 457.9 (58.9) 422.7 (44.0) 878.9b (183.8) 571.1 (390.7)
volume (m3/ha)
Numbers in columns represent means with Standard Errors in parentheses. All differences are P < 0.05. Analysis of Variance, Fisher’s Least
Significant Differences Test. Reprinted under CC BY 3.0, from Van Dyke, F., Probert, B.L., Beek’, G.M.V., 1995. Seasonal Habitat Use
Characteristics Of Moose In South- Central Montana. Alces 31, 15–26
a
Different from all other core areas
b
Different from LR
c
Different from BR
d
Different from BR, LR, and MC
e
Different from BR and EF
7.1 A Foundational Understanding of Habitat 265
to conserve, given its diet specialization for bamboo and

associated habitat preference for bamboo forests. Like most
species, giant panda use different selection criteria at differ-
ent spatial scales. Dunwu Qi and his colleagues conducted a
careful analysis of the problem of habitat selection at differ-
ent spatial scales by examining habitat use of giant pandas in
the Liangshan Mountains in China’s Sichuan Province. They
evaluated habitat use associated with (1) habitat found within
the home range of individual giant pandas; (2) the core area
of individual home areas (giant pandas often concentrate use
in as little as 10% of their total home range); (3) daily move-
ment (habitat characteristics found along the daily movement
path of a giant panda); and (4) seasonal elevational migration
(habitat selected at different elevations in different seasons).
Using this approach of multiple scales of measurement of
habitat preference, Qi and his colleagues were able to develop
indices of habitat suitability (i.e., a “habitat suitability
model” or HSM) for giant pandas to guide decisions on
which areas were most important to conserve (Fig. 7.3; Qi
et al. 2012). This approach and others like it are foundational
Fig. 7.2 The giant panda (Ailuropoda melanoleuca), an iconic species to creating HSMs that allow conservationists to evaluate
in world conservation, which, like many species, displays different potential reserve areas in terms of value for individual
habitat preferences at different spatial scales in a landscape. (Photo by species.
A. Lucidon/White House, reprinted under CC BY 3.0)
Fig. 7.3 Habitat quality maps for a b

giant panda (Ailuropoda
melanoleuca) in the Liangshan
Mountains, Sichuan Province,
China, derived from
measurements of habitat use by
pandas at different spatial scales,
including (a) the study area
examined using an ecological
niche factor analysis based on a c
spatial scale similar to distances
traveled by giant pandas during
seasonal elevational migrations.
The encircled area on the large
map shows habitat suitability
derived from analysis of the (b)
core range, (c) daily movement,
(d) home range and (e) seasonal d
elevational migration. Dark green
areas denote optimal habitat,
green denotes suitable habitat,
blue denotes marginal habitat, and
beige represents unsuitable
habitat. (Reprinted by permission
from Wiley, from Qi, D., Zhang,
S., Zhang, Z., Hu, Y., Yang, X., e
Wang, H., Wei, F., 2012.
Measures of giant panda habitat N
selection across multiple spatial Habitat Suitability
Optimal
scales for species conservation. Suitable
Marginal
The Journal of Wildlife Unsuitable
Management 76, 1092–1100. 0 30
# 2012 The Wildlife Society) Km
7.1.2.4 Improving Habitat Assessment Through high-resolution LiDAR data globally from space (NASA and
Advanced Technology and Modeling UMD 2020)), conservationists will be able to construct
Until recently, the conventional source of data for HSMs was HSMs of increasing accuracy and resolution.
digital cartography, which might employ characteristics of
land use, geology, or climate conditions as their foundational
attribute. Such an approach, no matter how rigorously 7.2 Heterogeneity, Landscape Gradients
applied, may overlook ecological structures and features and Patch Dynamics
important for wildlife. The development of Light Detection
and Ranging (LiDAR) technology, which employs laser 7.2.1 Habitat Heterogeneity, Gradients,
scanning techniques gathering data from sensors mounted and Patchiness
on airplanes or drones that can fly over areas of interest, can
accurately measure attributes of height and stratification of It is not enough to know which habitats a species might prefer
vegetation, canopy structure, quantity of woody biomass and to use. We must also know how habitats are distributed. A
even numbers of trees per area (Tattoni et al. 2012). Clara persistent characteristic of habitats is that they are variable in
Tattoni and her colleagues employed LiDAR technology in space and time, a quality referred to as habitat heterogeneity.
northern Italy to develop HSMs for birds of conservation Habitat heterogeneity can be defined as any form of variation
concern in that region – including the corncrake (Crex crex) in the environment, including physical and biotic
(Fig. 7.4), red-backed shrike (Lanius collurio), and Eurasian components. Such variation may appear as spatial or tempo-
green woodpecker (Picus viridis) – and then compared ral patterns (Ostfeld et al. 1997). Broadly, there are primarily
models constructed with and without LiDAR data. LiDAR two types of habitat models that explain habitat patterns in
variables including height of trees, percentage of trees, and landscapes. Patch models assume that heterogeneity exists
length of ecotone (area of gradation from one habitat type to because habitats occur in patches, which can be defined as an
another) proved significant for these species and classified area, smaller than a landscape, that contains only one type of
habitat more accurately than models that did not incorporate habitat (Danielson 1991). Patchiness, a description of habitat
LiDAR variables. Models created without LiDAR variables arrangement, can be defined as a form of spatial heterogene-
contained high levels of “omission errors.” That is, they more ity in which boundaries may be discerned. Patchy heteroge-
often mistakenly classified points where these species were neity appears as contrasting discrete states of physical or
present as points where they were absent. LiDAR models biotic phenomena. (Ostfeld et al. 1997). This so-called “patch
also displayed greater spatial specificity and resolution, as model” of landscape heterogeneity is appropriate to describe
can be seen in Fig. 7.5 which compares habitat suitability the spatial structure of some ecological systems, especially
models for the corncrake with and without LiDAR variables those in which there are clearly defined boundaries or edges
(Tattoni et al. 2012). As LiDAR and other technologies with between different kinds of habitat. The patch model is more
applications for assessing habitat suitability advance and accurate and more applicable at larger landscape scales. In
evolve (e.g., new US NASA missions like GEDI, collecting landscapes with habitat arrangements that fit the patch model,
species richness and species diversity will be correlates of
habitat heterogeneity and patchiness.
In many systems, however, a gradient model, in which
boundaries between different habitats are not clearly defined
and environmental conditions (variables) change slowly and
gradually at fine spatial scales, provides a more accurate
description of ecological reality. Gradient models are often
most applicable in environments where the distribution of
organisms, particularly plants, is strongly affected by one or
more continuously varying environmental variables, such as
moisture, elevation, temperature, or the concentration of a
particular soil nutrient or soil type. When this is the case,
habitat “patches” may not be easily discernible, and the
distribution of organisms, as well as community biodiversity,
is better modeled and more easily explained by tracking and
Fig. 7.4 The corncrake (Crex crex), a bird of conservation priority in predicting changes in the controlling variables. Gradient
northern Italy whose available habitat has been assessed using Habitat
Suitability Models (HSMs) employing Light Detection and Ranging
models can be especially useful at smaller, microhabitat
(LiDAR) data. (Photo courtesy of G. Thoburn (garytsphotos.zenfolio. scales, but in some ecosystems, they are also applicable at
com) larger scales, such as in some northern temperate forest
7.2 Heterogeneity, Landscape Gradients and Patch Dynamics 267
Fig. 7.5 Different spatial models of habitat suitability (orange areas) identifying habitat suitability at a landscape scale for this species when
for the corncrake (Crex crex) in northern Italy using Light Detection and LiDAR data are included. Black dots are points of species presence.
Ranging (LiDAR) data incorporating specific parameters of vegetation (Reprinted by permission from Elsevier, from Tattoni , C., Rizzolli, F.,
type and height (a) compared to models constructed without LiDAR Pedrini, P., 2012. Can LiDAR data improve bird habitat suitability
data (b). Note the greater specificity and resolution achieved in models? Ecological Modelling 245, 103–110. # 2012 Elsevier B.V.)
systems, in which the distribution of vegetation and 7.2.2 Habitats and Landscapes: Measuring
associated habitat is strongly influenced by gradients in tem- Scales of Space and Time
perature and moisture (Ohmann and Spies 1998).
The realities of heterogeneity, patchiness, and gradients Habitats, whether consisting of discrete patches or of
can cover a wide spectrum of conditions. At one extreme, gradients with indiscernible boundaries, are ultimately het-
patches of habitat can be entirely discrete and discernible erogeneous. Perceiving their heterogeneity depends on the
from one another. In the opposite condition, habitat, although spatial and temporal scales used to measure it. As we have
heterogeneous, may have subtle and almost imperceptible already noted, habitat and landscape structure are not con-
gradations from one kind of patch to another. A patchy stant but change over time and space. The higher the rate of
landscape possesses a rich internal structure of different change, the lower the probability of regional population
habitats, although sometimes with only gradual differences survival (Fahrig and Merriam 1995). Environmental hetero-
in environmental conditions across spatial scales. In any form geneity associated with landscape is a function of spatial and
or at any scale, patchiness and gradients create a discontinuity temporal heterogeneity. Spatial and temporal heterogeneity,
of resource distribution that provides a variety of niches for in turn, are functions of the intensity and frequency of land-
different species to exploit. Thus, heterogeneity, patchiness, scape disturbance.
and environmental gradients are dominant influences in hab- Habitats may be classified at multiple scales of heteroge-
itat composition and landscape structure, as well as landscape neity, but organisms respond to habitats at particular scales,
biodiversity. as we have already seen in the example of the giant panda.
Begin with a scale of time. To a casual observer, habitats give

an impression of being permanent landscape features. They
are not. Just as we speak of “population dynamics,” it is
appropriate to speak of “habitat dynamics.” In the context
of habitat dynamics, temporal scale can be defined as habitat
lifespan relative to the generation time of the organism
(Fahrig 1992). If the lifespan of the habitat is short relative
to that of the organism, organisms must enter periods of
dormancy to endure non-habitat environments until the habi-
tat reappears or employ nomadic behavior, or high rates and
distances of dispersal, to reach another suitable habitat. If the
lifespan of the habitat is long relative to the organism, seden-
tary behavior and more limited dispersal are favored.
To appreciate and analyze effects of spatial and temporal
scale independently, Lenore Fahrig created a stochastic
model of habitat-organism interactions that evaluated the
relative importance of each effect on population persistence.
Imagine a two-dimensional model universe of habitat “cells” Fig. 7.6 The effect of spatial scale on mean population size at four
of two categories, “useable” habitat cells in which organisms different levels of habitat persistence. Note that as habitat persistence
can survive and reproduce, and “non-usable” habitat cells (temporal scale) increases, average population size rises. (Reprinted by
permission from Elsevier, from L. Fahrig, 1992. Relative importance of
that are inhospitable. Usable cells are clustered as “habitat spatial and temporal scales in a patchy environment, Theoretical Popu-
patches.” They are not only clustered but transient, existing lation Biology 41, 300–314. # 1992 Academic Press, Inc)
for a limited time. New habitat patches appear at random at
discrete time intervals (“steps”). Fahrig’s model is simple, appearing unpredictably and disappearing frequently?
containing only six constants (fraction of grid in useable Because temporal scale may be more important than spatial
habitat, per capita birth rate per time step, death rate per scale, size of reserve fragments may be less important than
time step, age of organism at maturity, maximum population their persistence. Fahrig goes on to recommend that “since
per habitat cell, and fraction of population dispersing per time habitat continually undergoes modification due to processes
step) and three variables (lifespan of patches L, size of such as disturbance. . .more attention should be placed on the
patches S, and mean dispersal distance Z ) (Fahrig 1992). In duration of reserve fragments as habitat for particular species
each time step, four events occur in sequence: (1) habitat of interest” (Fahrig 1992). Good advice. Can we follow it?
patches are “born,” (2) some organisms survive and repro- Yes, but only if we can predict changes in habitat along a
duce while others die, (3) some survivors disperse to new temporal scale.
patches, and (4) some patches “die.”
By holding the amount of habitat constant but varying
values of patch lifespan, patch size and species dispersal 7.2.3 How Do We Predict Habitat Change?
distance, Fahrig was able to independently evaluate the
effects of temporal and spatial scale on population size. The 7.2.3.1 Predicting Habitat Transitions Using
model’s outcomes showed that populations increased with a Markov Model
increasing temporal scale (i.e., densities of organisms We have already examined how to build models that evaluate
increased the longer a given patch of habitat persisted) and the viability of individual populations through time (popula-
decreased with increasing spatial scale (i.e., as the ratio of tion viability analysis (PVA) models, Chap. 6). Can we also
distances between patches to the organism’s average dis- construct useful models that tell us something about how
persal distance increased, densities declined). In relative habitat and associated landscape structure might change
importance, temporal scale had a far greater effect on popu- over time? The answer is yes, and in much the same way.
lation stability than spatial scale (Fahrig 1992). At any spatial Let us begin with a simple approach that requires us to follow
scale, populations were larger when habitat patches persisted five steps in explicating the characteristics of a landscape.
longer (Fig. 7.6). We’ll use a hypothetical distribution of forest stand types
Important insights emerge from this model. If the goal of a common in mountainous areas of the western United States to
conservation effort is to preserve a particular species, illustrate our procedure at each step.
conservationists must determine the temporal scale of its To make our model work, we will need to know what
preferred habitat. Is its favorite habitat persistent relative to habitats or cover types are present at specific locations today
the life of the organism or are preferred habitats ephemeral, and what habitat or cover types were present in the same
7.2 Heterogeneity, Landscape Gradients and Patch Dynamics 269
Table 7.2 The transition matrix of a Markov model that permits predictions of transitions in habitat states from time t to time t + 1
t+1
t Aspen Lodgepole Spruce Douglas Fir Meadow Total
Aspen 2 14 2 0 2 20
Lodgepole 4 14 0 1 1 20
Spruce 2 10 7 0 1 20
Douglas fir 1 4 1 13 1 20
Meadow 5 3 6 3 3 20
Total 14 45 16 17 8 100
In this matrix, the total number (tally) of cells classified as “aspen” is 20. In 14 of the 20 cells “aspen” has been replaced by “lodgepole” and 6 remain
“aspen.” The row total for aspen is 14 + 6 ¼ 20 and 14/20 ¼ 0.7. Thus, 0.7 is the value of pij ( paspen to lodgepole) in this transition matrix, the
probability that aspen will be replaced by lodgepole over the observed time span. The diagonal elements in the matrix represent the proportion of
cells of each cover type that did not change (stasis rates). Design by F. Van Dyke
places at some time in the past. Let m represent the number of stage-based population model (Chap. 6). Both use the same
cover types. We might get that number using maps, structure, the Markov model. A Markov model assumes that
photographs, satellite images, vegetation surveys, or to predict the state of the system at time t + 1 one need only
other Geographic Information Systems (GIS) data. Let’s know the state of the system at time t (Usher 1992). The heart
select a point in the past (a point where we still have reliable of a Markov model is the transition matrix P, which
and comparable records) to provide us with a distribution of summarizes the probability that a cell in cover type i will
past cover types, and call this past point time t, and call our change to cover type j during a single time step, whether we
present point in time t + 1, which we’ll use as the point to are talking about habitats or organisms. The time step is the
estimate present distribution of forest cover types. Table 7.2 interval over which the data were observed to change. Mar-
will represent the transition matrix, showing the changes that kov models, although simple, can be solved by iteration to
occur from time t to t + 1. The first step needed to create this project the state of the system in the future. The predictions of
matrix is to divide the landscape into cells and identify the habitat change generated by the Markov model are important
habitat or cover type in each one of them. Second, tally all the considerations in determining an optimal conservation strat-
cells in which cover type in the cell changes from time t to egy because extinctions of individual populations are com-
time t + 1. Third, summarize all the tallies of all m cover types mon in habitat patches. Recolonization of populations
in an m m matrix (Table 7.2). Fourth, note that an individ- following extinction depends upon the distances between
ual element of the matrix, nij, (the number in the cell specified occupied and unoccupied patches (landscape spatial struc-
by the intersection of row i and column j), represents the ture), the rates and distances of dispersal by the organism
number of cells that changed from cover type i to cover type (dispersal characteristics), and the length of time that habitat
j over the studied time interval. In the fifth step, divide each patches persist (temporal changes). The stability of the com-
matrix element, such as nij, by its row total to generate the munity will depend on whether a species is likely to replace
transition matrix P in which a given matrix element, such as itself after a disturbance or whether it is likely to be replaced
pij of P, summarizes the proportion of cells that changed by another species. If changes in landscape structure occur at
from cover type i to cover type j during the studied time unnaturally high levels (as with some anthropogenic changes
interval. For example, suppose that in the matrix at time t, the described in Chap. 3), neither dispersal nor adaptation can
total number (tally) of cells classified as “aspen” is 20. Now keep up. The result is a high rate of local and regional
suppose that on 14 of the 20 cells “aspen” has been replaced extinction.
by “lodgepole” and 2 remain “aspen,” two are replaced by The interaction between rate of change in landscape spa-
“spruce” and two by “meadow.” The row total is 14 + 6 ¼ 20 tial structure and rate of change in dispersal behavior
and 14/20 ¼ 0.7. Thus, 0.7 is the value of pij in this transition determines the probability of a species’ regional survival.
matrix, which is the probability that aspen will be replaced by As long as the rate of change in dispersal behavior is greater
lodgepole from t to t + 1 (Table 7.2). The diagonal elements than the rate of change in landscape spatial structure, it is
in the matrix, pii, represent the proportion of cells of each possible for the organism to survive in the changing land-
cover type that did not change (stasis rates). We can repeat scape by moving around in it and integrating its resources
this procedure for every habitat or cover type, and thus over space. For example, in unaltered habitats of woods or
determine a transition rate for every habitat transition that brush in the eastern US, the white-footed mouse (Peromyscus
occurs over the time interval (Urban and Wallin 2002). leucopus) uses home ranges of less than 0.5 ha. When agri-
Notice the similarity of this transition matrix, and the cultural clearing fragments woodland habitat, home ranges
procedure to create it, to that of a transition matrix for a increase to tens of ha and mice may move hundreds of meters
in one night (Merriam and Lanoue 1990; Wegner and example, LANDIS-II was recently used to model potential
Merriam 1990). Similarly, red fox (Vulpes vulpes) disperse restoration success of the once widespread American chest-
farther in fragmented urban habitats than in less fragmented nut (Castanea dentata) –a species exterminated in the wild
boreal forests (Lindstrom 1989; Hansson 1991). Although by a disease, the chestnut blight, spread by a fungus
organisms like mice and foxes can adapt their dispersal (Cryphonectria parasitica) – in the center of its former
distances to changes in landscape structure, each species range in the central Appalachian Mountains of the United
has a maximum possible rate of change in dispersal behavior. States, incorporating expected effects of climate change on
If landscape structure changes faster than they can change the species physiology. The model demonstrated that such
their ability to disperse, species will be unable to recolonize change would favor the chestnut because of its optimal tem-
areas where local extinctions have occurred at a sufficient perature range (which would be likely under a warming
rate, and the regional population will become extinct (Fahrig climate regime) and relative drought resistance (Gustafson
and Merriam 1995). et al. 2018). This model-based evaluation of the potential
Both traditional and contemporary efforts in conservation restoration success of the American chestnut in its former
have traditionally focused on issues of spatial scale (i.e., range in a future of climate warning is one among a growing
preserving local populations in local places). But such efforts, number of examples demonstrating that process-oriented sto-
without considerations of temporal scale, may not conserve chastic models like LANDIS-II can provide an alternative
habitats or populations. Because rates of change in habitat approach to traditional Markov models, producing more reli-
and landscape structure have larger effects on populations able projections of future landscapes under the novel
than arrangements and distances between habitats, preserving conditions of the future (Chap. 3).
habitats and species alone is not a sufficient condition for Regardless of the types of modeling tools managers might
long-term population and habitat persistence. Only conserva- use to predict habitat change in landscapes, managing such
tion efforts that incorporate temporal, as well as spatial, change over time is an exercise in managing succession.
management of landscape processes will be effective in the Succession in a habitat can be managed toward the goals of
long run. This means that conservation biologists must be conservation in four ways. First, managers can alter the
able to manage patterns of habitat change through time. That frequency, extent, and intensity of disturbance events by
is, they must be able to manage patterns of ecological applying agents like fire, flooding, cutting, herbicides, or
succession. mechanical methods of vegetation removal at varying
intervals. The second way is to manage succession by alter-
7.2.3.2 Habitat Transition – Managing ing plant and animal interactions. One means to this end is to
Successional Processes manipulate the density of herbivores on the site following a
Changes in habitat predicted in the Markov model are often disturbance. Manipulations can range from no control, which
expressions of ecological succession, a pattern of continu- often leads to heavy use of vegetation by herbivores and
ous, directional, and non-seasonal change (replacement) of significant impacts on the amount and composition of vege-
plant populations on a site over time. A shortcoming of tation, to total exclusion of herbivores, a strategy that often
Markov models is that they are deterministic rather than produces communities of high plant biomass but low net
stochastic, and therefore assume that the past is a reliable productivity. A third way is to change the availability of
predictor of the future. Aside from the generic problems species that can invade a disturbed site. For example, range
inherent in deterministic models which we have explored managers restoring a degraded prairie might first burn the site
previously (Chap. 6), an additional problem of Markov and then seed it with native species that represent the system
models is that their assumption of making the past a predictor that will function best on that site, or that are of particular
of the future may not be true under current dynamics of value for conservation. In a forest, sites disturbed by logging
global climate change (Chap. 4). Therefore, more sophisti- may be arranged close to abundant seed sources of desired
cated process-oriented stochastic models simulating land- species, or such species may be planted directly on the site
scape dynamics, such as the LANDIS-II model (Scheller when logging is completed. Similarly, wetlands created by
et al. 2007), which has been used to predict change in forest deliberate flooding may be seeded with species of value to
landscapes over time, may be more appropriate and realistic conservation or have value as food and cover for wetland
for predicting succession. LANDIS-II, for example, is a wildlife. The same approach can be employed, in a negative
spatially explicit forest landscape disturbance and succession way, through species-specific herbicide application after the
model that can independently simulate multiple ecological disturbance event, thus altering the successional pattern. A
and anthropogenic processes and effects (Mladneoff 2004, fourth way of managing succession is to manipulate the
Gustafson et al. 2010). LANDIS-II can be linked to global availability of resources at a site, and so alter the interactions
circulation models so that global changes in climate can be of plants with the environment and with other species.
linked with landscape processes in the model simulation. For Adding specific nutrients and fertilizers to logged sites in
7.3 Dimensions of Destruction: Understanding Habitat Loss, Fragmentation, Isolation. . . 271
forests or to cleared or burned areas in prairies to favor the habitat conservation and species conservation, which are
establishment of particular species is a common management intimately related, require management of succession.
practice. Removing undesirable species from sites after dis-
turbance may be done to favor species considered more
beneficial to system function, more valuable for animals, or
How would the perspectives and activities of a reserve
of greater value in conservation. Leaving resources in place,
manager change if he or she changed the goal from
such as snags, following a logging operation, can increase the
achieving a state (e.g., a proportional abundance of a
availability of nest sites for cavity-nesting birds.
desired habitat or a targeted population level of a par-
Can a conservation biologist employ multiple methods of
ticular species) to enhancing a process like ecological
managing succession simultaneously to attain a conservation
succession? What specific changes can you imagine in
objective? Yes, but to do so requires ingenuity. One example
allocations of time, questions of research interest, and
of an initiative that used multiple approaches in succession
desired outcomes if such a change occurred?
management is Johnson and Leopold’s (1998) effort to man-
age habitat for the endangered eastern massasauga rattlesnake
(Sistrurus catenatus catenatus). The massasauga uses
non-forested wetlands throughout its range in the eastern
United States and Canada, preferring early successional
communities. It suffers habitat loss as succession proceeds, 7.3 Dimensions of Destruction:
especially from encroachment of woody plants into wet Understanding Habitat Loss,
meadows. Fragmentation, Isolation
Johnson and Leopold applied the first approach, directly and Degradation
altering the type, frequency, and intensity of disturbance
events, by cutting and burning all woody vegetation in 7.3.1 Defining Our Terms
selected experimental plots in the Cicero Swamp Wildlife
Management Area in US state of New York, a peatland Habitat degradation can be defined as the gradual deteriora-
complex inhabited by massasaugas. They then made use of tion of habitat quality (Fischer and Lindenmayer 2007).
the second method, altering plant and animal interactions, by Many kinds of disturbances, such as fire, create habitat
enclosing selected plots in wire fences, thus excluding needed by species that are themselves disturbance-
herbivores (primarily white-tailed deer, Odocoileus dependent. And many kinds of intentional habitat disturbance
virginianus). Finally, by applying herbicides specific for efforts are employed in the work of conservation as a means
woody vegetation combined with cutting, Johnson and of creating habitat for particular species. Human activities,
Leopold employed the third process, and altered the pattern however, also often degrade habitat by modifying it in a
of plant invasion subsequent to the disturbance. variety of ways along a continuum of increasing unplanned
Over 3 years, Johnson and Leopold were successful, to anthropogenic disturbance that can result in decreasing con-
varying degrees, in altering patterns of succession through nectivity between landscape elements (Fig. 7.7). Today
these strategies, particularly in achieving reductions in the humans are estimated to have modified over 50% of the
density, basal area, and height of shrubs in treated plots Earth’s surface area in some manner. Such modification can
compared to untreated areas, and, with herbicides, increasing be described in four categories. Intact habitats and
the rate of mortality and decreasing the rate of re-sprouting landscapes are characterized by contiguous natural vegeta-
among woody species. Although response of the massasauga tion covering more than 90% of their land surface area. In
to treatments was inconclusive and limited by a small sample these environments, human modification may create
size (nine radio-marked massasaugas), massasaugas did use variegated habitats and landscapes in which human alter-
treated areas at more than four times their availability (10.1% ation reduces the amount of natural habitat and introduces a
of locations in treated areas that made up only 2.5% of the greater degree of landscape heterogeneity and variety of
total area). vegetation. More intensive human modification can lead to
Johnson and Leopold’s effort illustrates the role and value fragmented habitats and landscapes in which most original
of managing succession in conservation. Foundationally, this vegetation cover is removed, and remaining natural vegeta-
approach is often effective because ecological models based tion exists in smaller, non-contiguous “pieces” or
on habitat succession have proven more accurate at “fragments” separated by matrix habitat which differs, to
predicting the distribution of organisms than null models varying degrees, from habitats characteristic of natural vege-
(models based on random environmental fluctuation) or iso- tation. The most extreme and extensive human modifications
lation models (models based on island biogeography theory in which natural vegetation associations are reduced to
and metapopulation theory) (Kareiva et al. 1997). Both relictual habitats in which only minimal (<10%) natural
vegetation remains, and this exists in fragments separated by

sharp boundaries adjacent to highly modified, more inhospi-
table habitat (matrix habitat that is so degraded it may be
referred to as “non-habitat”).
In landscapes in which the modified matrix habitat
becomes the predominant landscape element, managing the
matrix between fragments of natural or semi-natural habitat
may be more important than managing the fragments, and
more decisive in achieving desirable conservation outcomes.
Species that depend on occurrence and arrangement of spe-
cific kinds of habitat patches (“patch-dependent” or “frag-
ment-dependent” species) are most influenced by three
primary matrix-related impacts. These are the effects of the
matrix habitat on movement and dispersal, the availability of
essential resources, and the abiotic environment of the
matrix. Such impacts are modified by variation in the matrix
habitat, the spatial and temporal scale of that variation, and
the adaptability of the species attempting to use, pass
through, or reside in the matrix (Driscoll et al. 2013; Fig. 7.8).
The process and effects of habitat destruction and degra-
dation often follow a predictable sequence. (1) A block of
contiguous habitat is reduced in size by conversion to
non-habitat (habitat loss) and increasingly broken into
numerous smaller patches (habitat fragmentation).
(2) Remaining patches decrease in size and number because
Fig. 7.7 A conceptual model of four states of increasing landscape of increased vulnerability to disturbance and invasion by
modification that affect habitat quality. (Reprinted by permission from
other species and increased ease of alteration or removal by
Wiley, from Fischer, J., Lindenmayer, D.B., 2007. Landscape modifica-
tion and habitat fragmentation: a synthesis. Global Ecology and Bioge- humans. (3) Distance between patches increases (habitat
ography 16, 265–280. # 2007 The Authors) isolation). (4) Quality of the patches decreases (degradation)
Fig. 7.8 Five dimensions of

influence of matrix habitat on
patch-dependent species. Three
core effects (dispersal, resource
provision, and abiotic
environment of the matrix), affect
species through the spatial
variation, spatial scale, temporal
variation, and temporal scale of
the matrix, mediated through the
adaptability and plasticity of the
species. (Reprinted by permission
from Elsevier, from Driscoll, D.
A., Banks, S.C., Barton, P.S.,
Lindenmayer, D.B., Smith, A.L.,
2013. Conceptual domain of the
matrix in fragmented landscapes.
Trends in Ecology & Evolution
28, 605–613. # 2013 Elsevier
Ltd)
(Oksanen and Schneider 1995). Although loss, fragmenta-

tion, isolation and degradation can occur concurrently, they
must first be analyzed separately to be clearly understood.
Habitat loss refers to physical destruction of habitat
(which often accompanies fragmentation), such that, in a
given area, less habitat remains after a destructive event.
Habitat fragmentation refers to a single block of contiguous
habitat being broken up into small “pieces” that are no longer
directly connected or adjacent to one another. It is possible
for habitat loss to occur without habitat fragmentation, but it
is not possible for habitat fragmentation to occur without
habitat loss. Habitat isolation, which often follows fragmen-
tation, refers to a condition where remaining fragments
become separated from one another by increasing distance
or by increasing resistance of matrix habitat between them to
Fig. 7.9 The effect of habitat loss on amounts of edge and interior
movement of individuals from one fragment to another. habitat. When a 40% portion of habitat is removed, the amount of
Habitat degradation, which can co-occur with habitat loss, interior habitat decreases by approximately 60%. In this scenario, edge
fragmentation, or isolation, refers to habitat which has lost species do not experience a significant increase or decrease in habitat
critical resources or structures needed by individual species,
and therefore affects the amount of available habitat. Habitat
fragmentation and isolation, while potentially affecting habi- and 70% of world forests are within 1 km of an edge (Haddad
tat amount, also affect habitat configuration, the physical et al. 2015). As fragments are created through habitat distur-
spatial arrangement of habitat and the proximity and related- bance and alteration, the amount of edge (linear length of
ness of different parcels of habitat to one another in the same boundaries between different kinds of habitat) increases.
landscape. With such increase, there can be a corresponding increase
In conservation science, the two categories that are most in the strength of edge influences (EI), also called “edge
commonly confused or conflated are habitat loss and habitat effects.” Consider a forest in which 40% of original habitat
fragmentation, terms often used with careless interchange- in a single block is destroyed and the remainder exists as two
ability. Carelessness in expression can lead to confusion in blocks (Fig. 7.9). In this case, species adapted to edge
concept in deciding whether loss or fragmentation is the environments (“edge species”) suffer no loss of habitat, but
process causing the ecological effect and conservation con- species adapted to environments associated with the forest
sequence. As habitat scientist and modeler Lenore Fahrig put interior (“interior species”) lose 60% of their habitat. Given
it bluntly, “Habitat loss should be called habitat loss; it has the pronounced effect of habitat fragmentation on a habitat
important effects on biodiversity that are independent of any patch’s area-to-edge ratio, it is not surprising that some of the
effects of habitat fragmentation per se” (Fahrig 2003:509). most well-known and well-studied aspects of habitat frag-
Further, Fahrig has asserted that cross-extrapolation of effects mentation are edge effects. You can see just how devastating
of habitat fragmentation to effects of habitat loss (i.e., the even simple, “ordinary” disturbances can drastically reduce
assumption that habitat fragmentation creates effects similar the amount of interior habitat through the example in Infor-
to habitat loss) is unreliable (Fahrig et al. 2019). Can we mation Box 7.1.
separate effects of habitat loss from habitat fragmentation
and, in so doing, create better outcomes in habitat Information Box 7.1: A Road, a Powerline,
conservation? and the Creation of Edge
Begin with a 1 km2 (100 ha) block of contiguous
habitat (Information Box Fig. 7.1). Assume that
7.3.2 Isolating Consequences of Habitat changes associated with edges penetrate 100 m at
Fragmentation and Effects of Edge each border and are the same on all sides. This reduces
the amount of “core” habitat (habitat unaffected by
Today habitat fragmentation is a globally pervasive phenom- edge effects and processes) to an area 800 m long and
enon. A recent global analysis of forest fragmentation, for 800 m wide or 64 ha. Now bisect this block by one road
example, revealed that 20% of the world’s remaining forest is
within 100 m of an edge – a border with a non-forest habitat – (continued)
7.3.3 Habitat Alteration Through Effects

Information Box 7.1 (continued) of Edge: First Principles
and one power line. Assume that 10 m 100 m are lost
for the power line and its right of way (1000 m2), 10 m Edge effects are complex and vary according to time,
100 m are lost by the space taken up by the road variables studied, edge orientation, management history,
(1000 m2), and that edge effects continue to penetrate and other factors. Despite the great diversity of form and
100 m at each boundary. The habitat unaffected by effect that characterize edges, all edges create: (1) exchange
edge is now reduced from a single block of 64 ha to or flow of energy, material, and organisms across the bound-
4 separate blocks, each 290 m on each side, or 8.41 ha ary and (2) alterations in biophysical processes and ecosys-
in area. The four blocks now contribute only 33.64 ha tem composition and structure. Analytically, the degree of
of habitat unaffected by edge effects, little more than edge influence (EI) will be determined by three things. The
half (53%) of the unaffected habitat present in the first is a quality that is known as patch contrast. The greater
original block. Note that, as long as fragmentation the difference between the two habitats forming the edge, the
continues, edge habitat increases and interior habitat greater is likely to be the effect of edge influences on both
decreases. And notice also that, in the early stages of habitats. Flows of energy across an edge increase with greater
fragmentation, interior or “core” habitat is moved patch contrast, and such flows regulate the magnitude and
closer to edge habitat and to the processes and effects distance of edge influences into interior habitat. Habitat
associated with the edge environment. managers generally believe that edge influence can be
minimized by reducing the contrast between patches. For
example, in a forest, reducing contrasts between age or struc-
ture of adjacent forest stands has long been recommended to
reduce edge influences (Harris 1984).
A second determinant of EI is the rate of dissipation of the
edge effect over distance from the edge into the interior of the
habitat. This distance effect is sometimes incorporated into
models that attempt to simulate effects of edge influences, in
which case it is sometimes referred as decay value, K. Daolan
Zheng and Jiquan Chen developed a model for measuring
edge effects in forest environments in which they specified
decay values for different variables (Zheng and Chen 2000).
In their model, the decay value would determine the rate of
change from a clear-cut to the interior forest for a specific
variable such as, for example, wind speed. The larger the
decay value, the faster the variable changes per unit distance
from the edge. Large decay values are associated with rapid
change in the value of the variable over a short distance,
while low values would indicate little change over a long
distance (Fig. 7.10). Individual K values can change signifi-
cantly from one variable to another. For example, at the edge
of a forest clear-cut, wind speed might change from 40 kph at
the edge to 10 kph 20 m from the edge, a decay value that
could be expressed as 30 kph/20 m or 1.5 kph/m.
A third determinant of the effect of EI is that of magnitude
or strength of edge influence. To remain with our example of
wind effects, magnitude could be estimated as the speed of
the wind at the edge, which might vary from 40 kph at the
Information Box Fig. 7.1 The effect of habitat fragmentation edge of a forest – clear-cut boundary to <5 kph at a boundary
on edge and interior habitat. Note, in this scenario, when a road between two different kinds of forest of similar ages and
and power line intersect the habitat, edge species experience a net
structures. Decay and magnitude can change independently
habitat increase of over 80%, while interior species lose nearly
50% of their original habitat in the same variable and will not necessarily respond to edge
creation in the same way. The relationship between decay
and magnitude could be envisioned as a gradient that could
vary from steep and short (a variable of high magnitude that
increase in extent of edge influence over time (Fig. 7.11)

(Harper et al. 2005).
All edges are characterized by distinct abiotic and biotic
gradients associated with these effects. Flows across edges
driven by these gradients have been likened to movement
across a semipermeable membrane. Consider a common
direct effect of edge creation in a forest, tree damage. Tree
damage leads to reduced canopy cover and greater abundance
of snags and logs at edges. In this case, the primary process
response of tree mortality has greater magnitude and distance
of EI than the primary structural response. Following primary
edge responses, accentuation of abiotic gradients near edges
is the probable mechanism for secondary process vegetation
responses of regeneration, growth, and mortality. These sec-
ondary processes often have distance and magnitude similar
Fig. 7.10 Theoretical relationships between magnitude of individ- to or greater than primary responses, probably because pri-
ual edge factors (EFi) and distance from edge as functions of decay mary structural responses initiate secondary responses that
values (K ) in a forest landscape. Variables with high values of K are occur farther from the edge (Harper et al. 2005).
associated with edge effects that decay quickly over a short distance into There are situations in which EI is more pronounced and
the forest. Variables with low K values are effects whose influence
declines slowly over a long distance. (Reprinted by permis- more ecologically important, other conditions being equal.
sion from Elsevier, from Zheng, D., Chen, J., 2000. Edge effects in These are: (1) high mean annual (or growing season) air
fragmented landscapes: a generic model for delineating area of edge temperature, (2) low latitudes with high solar radiation,
influences (D-AEI). Ecological Modelling 132, 175–190. # 2000 (3) low mean annual (or growing season) cloud cover, (4) fre-
Elsevier Science B.V.)
quent, extreme winds, (5) edges facing the equator or into
prevailing winds, (6) shallow soil depth, (7) abrupt, open
loses its effect over a short distance) to shallow and long edges, (8) edges where patch contrast is maintained over
(a variable of low magnitude that retains its effects deep into time, (9) forests with tall, dense canopies, (10) closed-canopy
the interior of the habitat (Zheng and Chen 2000). (generally mid-successional or mature) stands, (11) regional
Edge effects that are influenced by patch contrast and can flora or fauna with many pioneer species, (12) regional flora
be measured by decay and magnitude can be classed as or fauna with many exotic and invasive species, (13) biomes
“primary” or “secondary” effects of edge influence, some- or forest types subject to infrequent stand-replacing
times referred to as “direct” or “indirect” effects. Primary disturbances, or (14) forest communities or landscapes with
responses are those resulting immediately and directly from low inherent heterogeneity in vegetation, topography, or
the effects of edge creation. To remain with our forest sce- soils. When managers work to conserve biodiversity under
nario, primary process responses could include damage to these conditions, they must be particularly concerned with
trees and other vegetation; disruption of the forest floor and how to mitigate EI.
soil; increased dispersal of pollen and seeds; changes in
evapotranspiration, nutrient cycling, decomposition, and
rates of energy exchange. These ecological processes are 7.3.4 Environmental Characteristics of Edges
then mechanisms responsible for primary structural
responses such as changes in forest structure (including can- Traditionally, edge effects have long been assumed to lead to
opy cover, tree density, downed wood, leaf area, and vegeta- habitat degradation, especially through pressures that change
tive biomass). Subsequently, secondary responses arise conditions in habitat blocks of reduced size (fragments).
because the primary responses change the original abiotic Certainly, once EI is in play, associated effects and processes
and biotic gradients associated with the edge. Secondary can create an environment very different than interior habitat.
processes to edge creation such as, in vegetation, regenera- Some differences are physical and obvious. Edges usually
tion, growth, reproduction, and mortality, reflect both edge- receive more direct insolation, and thus are typically warmer
related gradients and primary responses to them. As edges and have lower relative humidity than interior areas. Radia-
age, they may be subject to processes such as sealing, soften- tion and moisture fluxes are greater at edges. Increased radia-
ing, or expansion. “Sealing” refers to the development of tion and insolation at edges tend to increase soil temperatures
dense vegetation at the edge, an effect especially pronounced and affect rates of invertebrate and microbial activity (Klein
at maintained edges. “Softening” is the reduction of edge 1989; Parker 1989) as well as processes of nutrient decom-
influence at regenerating edges. “Expansion” refers to position. Increased soil temperatures may also reduce
Fig. 7.11 Conceptualized diagram of processes and responses follow- dense vegetation at maintained edges. Edge softening is the reduction of
ing edge creation including (a) a recently created edge and (b) an older edge influence at regenerating edges, and edge expansion is the increase
edge. Upward and downward arrows within boxes denote increases and in extent of edge influence over time. (Reprinted by permission from
decreases, respectively. Abiotic and biotic gradients (triangles, with Wiley, from Harper, K.A., Macdonald, S.E., Burton, P.J., Chen, J.,
height representing magnitude of edge influence (MEI) and length Brosofske, K.D., Saunders, S.C., Euskirchen, E.S., Roberts, D., Jaiteh,
representing distance of edge influence (DEI) are strong at newly created M.S., Esseen, P.-A., 2005. Edge Influence on Forest Structure and
edges and become steeper, weaker, or longer at older edges with edge Composition in Fragmented Landscapes. Conservation Biology 19,
sealing, softening, or expansion. Edge sealing is the development of 768–782. # 2005 Society for Conservation Biology)
retention of water in the soil and alter growth rates and can lead to increased evapotranspiration and desiccation.
phenology of vegetation. Secondary effects of wind at edges can include increased
Wind behaves differently at edges than in the interior of a transport of material into the interior from surrounding, but
habitat. As wind moves over a landscape and flows from one different, vegetation types, including soil, seeds, insects and
type of vegetation to another, the upper part of the wind dead organic matter.
profile retains the characteristics formed over the previous Habitat fragmentation and its associated creation of edge
vegetation type, while the lower portion takes on the profile can also alter local water regimes. Rates of interception and
characteristics of the new vegetation. The two profiles do not evapotranspiration are changed by removal of native vegeta-
fully equilibrate with one another for some distance, increas- tion and by modification of native vegetation at the edges of
ing turbulence and wind shear at their boundaries. The dis- remnants (Saunders et al. 1995). Replacement of deep-rooted
tance required for equilibration is normally 4–6 times the perennials, which are more typical of native vegetation, with
height of the vegetation. The effects of edge on wind behav- herbaceous crops, pasture, or non-vegetative surfaces at
ior have important consequences. Trees near an edge are edges leads to greater runoff and increased surface and
more susceptible to damage, wind pruning, uprooting, and groundwater flows, with accompanying increases in rates of
other forms of physical damage. Increased wind speeds also erosion and transport of particulate matter.
Other characteristics of edges are manifested as biological 7.3.5 What Lies Between? Managing Matrix
phenomena because biological interactions also change at Habitat
edges, usually to the detriment of species adapted to interior
habitats. Edge habitats create a variety of effects on interior The idea of habitat “fragmentation” implies that the habitat
species of both plants and animals, which can include remnants or “fragments” that remain after division of a con-
increased predation, increased parasitism and increased her- tiguous block of habitat are isolated by areas (the “matrix”)
bivory (Alverson et al. 1988; Harris 1988; Temple and Cary that are hostile environments to species within the remnants.
1988; Yahner 1988). For example, the presence of edges can This may not always be the case. For example, a meta-
alter and intensify interspecific interactions. Some generalist analysis of 63 wetland studies examining 330 population
mammalian predators, such as raccoon (Procyon lotor) and responses in 155 species determined that the amount of forest
red fox (Vulpes vulpes) are known to preferentially follow cover in a landscape was more important to amphibians than
edges rather than forage in interior habitat areas. Some birds the amount of wetland cover, suggesting that the quality of
common to edges, such as blue jays (Cyanocitta cristata) and the matrix may be more valuable than the quality of the
house wrens (Troglodytes aedon), may engage in nest preda- breeding and rearing habitat (Quesnelle et al. 2015). Whether
tion (blue jays) or the destruction of eggs of other species the matrix represents a threat or an asset to a given population
(house wrens). can often be a highly species-specific response, but
Increased edge results in less secure habitat for interior maintaining continuity of remnant groups as a functional
species because nest predators and parasites not only reduce population in a landscape of dispersed habitats requires either
their nesting success at edges but penetrate more deeply into some means of creating connectivity between the fragments
the interior of other habitats from edges (Temple 1986). For (which can be affected by managers) or the ability of a
example, increased rates of nest predation among forest birds species to develop sufficiently long-distance dispersal
may extend up to 600 m into the forest from the edge abilities enabling it to move from one fragment to another
(Wilcove 1985). Thus, there is less habitat in which resident (which is a trait intrinsic to the species, and not easily affected
species are unaffected by edge processes and effects. When by managers). The first question to answer is: how hostile is
habitat is fragmented and the ratio of edge-to-interior the matrix? Can the species move through it, benefit from it,
increases, species dependent on particular habitats become or even live in it?
more vulnerable to predation because they have little habitat Some classic studies shed light on these questions. For
left that edge predators cannot penetrate. In North America, example, Diffendorfer et al. (1995) examined the effect of
species of insectivorous songbirds with high vulnerability to habitat fragmentation on three species of North American
predation during the breeding season have declined signifi- grassland rodents, the hispid cotton rat (Sigmodon hispidus),
cantly in recent years, while those species with low vulnera- prairie vole (Microtus ochrogaster) and deer mouse
bility to predation during the breeding season have increased (Peromyscus maniculatus) by experimentally fragmenting
(Robinson et al. 1995). This finding has led some to conclude grassland habitat to varying degrees. The largest of these,
that habitat fragmentation and edge creation in temperate the cotton rat, had higher densities in contiguous habitat, but
forests, with associated increases in penetration depths of the two smaller species had higher densities in fragmented
predators, may be more important in songbird declines than habitat. The smallest species, the deer mouse, had the most
the more publicized problem of tropical deforestation. positive response to fragmentation, was most abundant in the
More recent reviews have criticized this traditional inter- smallest areas, and used the interstitial area (the matrix) that
pretation. For example, in a review of landscape-scale empir- was created to separate habitats. In other words, to the deer
ical studies, Fahrig (2017) found that ecological responses to mouse, there was no fragmentation. The matrix was simply
habitat fragmentation per se (fragmentation independent of more habitat. Differences in effects of fragmentation in these
habitat amount) were usually non-significant (>70% of species also arose from a combination of differences in habi-
responses) and that 76% of significant relationships were tat quality and changes in competitive interactions. Cotton
positive. Species abundance, occurrence, richness, and other rats suffered because smaller sites had insufficient resources
response variables examined in multiple studies increased for long-term survival, and its decline in fragmented habitats
rather than decreased with habitat fragmentation per se. It is created competitor-free space for the two smaller species.
important to note that positive effects of fragmentation were (Diffendorfer et al. 1995).
generally observed at landscape scales through mechanisms To the cotton rat, the habitat was fragmented. To the deer
such as increased habitat diversity, spreading of risk, and mouse, the habitat was variegated, a landscape of real (and
landscape complementation. In past studies, including those human-induced) differences, but not so different as to create
we have reviewed here, negative effects of habitat fragmen- barriers between habitat patches or any real loss of habitat.
tation, especially on interior species, have been most strongly This pattern of perception is not confined to rats and mice.
manifested at patch scales of habitat. McIntyre and Barrett (1992), for example, found that, in
Australian grasslands, human-induced changes create more

habitat variegation than habitat fragmentation. Among plants, Information Box 7.2 (continued)
they found native species that could grow over the whole
range of human habitat modifications found at their study
sites. “For these species,” they noted, “potential habitat forms
a continuum across the landscape” (McIntyre and Barrett
1992). Other plant species were intolerant to most or even
all forms of habitat modification, and for these the habitat was
truly fragmented. But most species fell between these
extremes. Such species could be conserved with well man-
aged human-induced habitat modifications. McIntyre and
Barrett note, from their own and other studies, that conservation
strategies often concentrate on managing fragments which
benefit species specially adapted and restricted to these
fragments but ignore species that can effectively use both the
fragments and intervening areas. Their insight: don’t simply
manage the fragments – manage the matrix, and more species Information Box Fig. 7.2 The eastern collared lizard
(Crotaphytus collaris), whose relict eastern subpopulations are
will benefit! This principle is well-illustrated in practice in one being restored through management (prescribed fire) of its matrix
species of reptile, the eastern collared lizard (Crotaphytus habitat (deciduous forests), permitting it to disperse more effec-
collarus), which requires effective and specific management tively to preferred but fragmented habitat (rock-covered forest
of its habitat matrix to survive (Information Box 7.2). openings). (Photo courtesy of J. Briggler, Missouri Department
of Conservation)
Information Box 7.2: The Eastern Collared Lizard

and the Management of Matrix Habitat
The collared lizard (Information Box Fig. 7.2) is native When it is not possible to make the matrix more hospitable
to and common in dry grasslands and deserts of the for dispersal, it may be wise to shift priority to preserving
southwestern United States, but its historic range habitat fragments in close proximity to one another. Some
extended further east. As this eastern part of its range species, like forest dwelling amphibians, may be “fragment
became wetter and more forested in recent centuries, specialists,” for whom no amount of matrix management
collared lizards declined, and their declines have been creates benefit. In these species, fragments themselves must
exacerbated by fire suppression in southeastern US be carefully managed, and priority of protection should be
forests (fires create openings that benefit collared given to clustered arrangements rather than to fragments
lizards) and increased conversion of grasslands to agri- isolated by distance from other fragments. But, to understand
cultural lands in the eastern portion of its range. Today the problem of habitat conservation more thoroughly, we
only a relict eastern population of collared lizards must separate and distinguish the effects of habitat loss
remains, confined to rocky openings (“glades”) in the from habitat fragmentation. A remarkably well designed
Ozark Mountains, primarily in the states of Arkansas field study of these processes on two species of spider
and Missouri. Relict populations had declined because shows us how.
collared lizards would not disperse through forests
surrounding the glades, a “matrix” they perceived as
inhospitable. Recently conservation managers in 7.3.6 Loss and Fragmentation: Experimental
Missouri reintroduced fire around the lizard’s glades Isolation of Separate Effects
through a program of prescribed burning. When burned
areas occur at increasing frequency and connect multi- Samuel Marshall, Sean Walker, and Ann Rypstra, biologists
ple glades, collared lizards dispersed from glade to at Miami University in Ohio (USA) found themselves
glade and began establishing stable metapopulations increasingly concerned that the effects of habitat loss, habitat
(Templeton et al. 2011; Driscoll et al. 2013). connectivity (fragmentation), and habitat degradation due to
EI were too often confounded in conservation studies, such
(continued)
that, even when a significant effect was observed on affected But how could one create habitats in which habitat loss,
populations, it was impossible to determine exactly which fragmentation, and isolation could be manipulated
process was causing the effect. As Marshall et al. noted, “Part independently?
of the challenge in studying the mechanisms behind the Marshall and his colleagues used a pre-existing experi-
influence of habitat fragmentation on species persistence is mental field formerly cultivated to grow soybeans. The field
that the spatial and ecological attributes of fragmented had been converted to conservation tillage, producing a habi-
landscapes have correlated effects. . . fragmented landscapes tat more like a native North American prairie, with increased
generally suffer a net loss of habitat in addition to increasing densities of vegetation, a change that produced an increase in
spatial subdivision leading to an overall decrease in connec- the densities of wolf spiders. Using arrays of 25 2.0 2.0 m
tivity and core habitats” (Marshall et al. 2006:241). Many habitat cells (“habitat islands”) in multiple field plots, they
studies of fragmentation have assumed that species with randomly selected either 5 (20%) or 20 (80%) of the cells in
certain ecological traits like large home range size, habitat each array for disturbance (which Marshall et al. referred to
specialization, and aversion to edges (i.e., “interior” species) as “destruction” because all vegetation was removed) with
will suffer more from habitat loss, fragmentation (connectiv- one array left undisturbed as a control. Then, in a separate
ity) and edge influence, and species with opposite traits will experiment to isolate the effect of fragmentation, they divided
suffer less, or even benefit from it. But the characterization of a 25 m2 area into two, four, and eight equally-spaced squares,
some species as “interior species” or “edge species” has often while keeping a separate single 25 m2 patch undivided to
been made with little empirical support or careful represent a least-fragmented habitat. Subdivision carried out
investigation. this way would keep the total amount of habitat constant but
To isolate these correlated but distinct effects, Marshall increase fragmentation. Finally, to isolate effects of edge
and his colleagues examined the effect of landscape pattern influences, they created three different landscape
on populations of two species of wolf spider, Pardosa configurations: (1) square, 5 5 m; (2) rectangle, 7.5
milvina and Hogna helluo (recently renamed Tigrosa helluo). 3.3 m; and (3) long rectangle, 10 2.5 m (Marshall et al.
Wolf spiders are vagrant, generalist invertebrate predators 2006:243–244) (Fig. 7.13). By creating habitats of different
that hunt their prey on the surface of soils. Because they shapes, Marshall et al. changed the length of edge without
dwell almost exclusively on the surface, they are sensitive changing the amount of habitat. In their own words, this is
to changes in soil-surface conditions, especially moisture and what happened.
temperature. The major determinants of soil moisture and “There was a strongly opposing response to habitat
temperature are the frequency of disturbance and accumula- destruction by the two wolf spider species. Population
tion of vegetative litter. Tigrosa helluo (Fig. 7.12) is larger, densities of Hogna [Tigrosa] declined by approximately
more habitat selective, a poor disperser, and averse to bare 75% with only a 20% reduction in area, while Pardosa
(i.e., disturbed) surfaces. In contrast, Pardosa milvina is numbers are almost twice as high in the 80% area reduction
smaller, more mobile, and makes use of barren surfaces. treatments as in the no-destruction controls” (Marshall et al.
2006:244). In other words, habitat loss, independent of isola-
tion or fragmentation, had greater effect on Tigrosa, the
“interior” species, than on Pardosa, the “edge” species. Fur-
ther, “There was a negative response by Hogna [Tigrosa]
populations to the increasing subdivision of the habitat. . .and
no significant response to habitat fragmentation by Pardosa.”
(Fig. 7.14) and “Hogna [Tigrosa] populations had a negative
response to increasing edge in the subplots. . ., whereas
Pardosa was apparently unaffected. . .” (Fig. 7.15) (Marshall
et al. 2006:245). Habitat loss, habitat fragmentation, and
increased edge independently had negative effects on
Tigrosa, but not on Pardosa.
From their results, Marshall et al. perceived that “. . .‘the
matrix’ is generally assumed to be inhospitable to the focal
species and if this assumption is false it will lead to misinter-
pretation of experimental results... the same barren habitats
that function as an aversive matrix for Hogna [Tigrosa] (i.e.,
Fig. 7.12 Tigrosa helluo, a large, habitat selective species of wolf
spider with poor dispersal ability and aversion to habitat disturbance,
functionally inhospitable) may be a useful, and even neces-
experiences population declines when subjected to habitat fragmenta- sary, landscape element for Pardosa. . .Hogna [Tigrosa] and
tion. (Photo courtesy of J. A. Godfrey) Pardosa present a revealing contrast of how two related,
Fig. 7.13 Aerial photograph of

the experimental fields near
Miami University, Ohio (USA)
used in the study of responses of
two species of wolf spider Tigris
helluo and Pardosa milvina to
habitat fragmentation. Each field
is 60 70 m and holds a replicate
of either an area reduction
experiment (fields with three large
subplots) of the area subdivision
experiment (fields holding four
smaller subplots). (Photo courtesy
of S. D. Marshall and reprinted by
Marshall, S.D., Walker, S.E.,
Rypstra, A.L., 2006. Two
ecologically-divergent generalist
predators have different responses
to landscape fragmentation. Oikos
114, 241–248. # 2006 Oikos)
syntopic species will respond to habitat fragmentation as a that occurred immediately following a fragmentation event,
result of their divergent ecologies” (Marshall et al. which would represent only temporary increases. But even
2006:247). Whether terms like “edge species” or “interior some longer-term studies with plant species did not show a
species” ultimately prove useful to science, Marshall et al. clear pattern.
succeeded in demonstrating that habitat loss and habitat Corridors enhance movement between fragments – Here
fragmentation have different effects on different species, results were species specific. Fragmentation usually reduced
even species that are closely related, and such effects can movement of individuals between fragments, but corridors
have important population consequences. did not always enhance it. Most studies found that corridors
increased movement between fragments for some species,
but not all (Debinski and Holt 2000).
7.3.7 A Larger Perspective: Long-Term Since Debinski and Holt’s review, more recent analyses
Studies of Habitat Loss have been more definitive. Nick Haddad and his colleagues
and Fragmentation examined results of the world’s longest running habitat
experiments examining, separately and together, effects of
The work of Samuel Marshall and his colleagues is valuable reduced area (habitat loss), increased habitat isolation, and
in understanding the distinctive effects of habitat loss and increased edge at multiple scales in multiple biomes spanning
habitat fragmentation, but it is one experiment. To better five continents and 35 years of research. In these longer-term
understand effects of habitat loss and fragmentation at a experiments, fragmentation consistently reduced species
broader scale, Diane Debinski and Robert Holt undertook a diversity (13–75%) and impaired ecosystem function by
comprehensive review of fragmentation experiments at the decreasing system biomass and altering nutrient cycles.
turn of the millennium, in 2000, and found only 20 studies in Reducing the area of habitat reduced not only species rich-
which fragmentation had or was being studied through direct ness but also reduced species persistence and altered commu-
experimental manipulation in the field. Those studies showed nity composition. Increased distance between habitat
the following trends. fragments (increased isolation) reduced species abundance,
Species richness increases with habitat area – Actual persistence and richness as well as movements between
manipulative experiments supported this hypothesis, espe- fragments. Increased edge, like reduced area, altered commu-
cially in arthropod species (like spiders, for example). Causes nity composition, and all three effects (increased loss, isola-
of decline with fragmentation included loss of specialist tion, and edge) negatively affected ecosystem processes like
species, increased mortality in specific life history stages in nutrient retention, succession rate, pollination, productivity
fragments, and reduced ability of some species to colonize and other ecological functions (Haddad et al. 2015)
smaller habitat patches relative to larger ones. (Fig. 7.16).
Species density or abundance increases with area – Here Although the results of these long-term experiments on
results offered less support, possibly due to crowding effects habitat loss, isolation, and fragmentation show consistently
7.4 Theories and Models of Loss and Fragmentation 281
Fig. 7.15 (a) Response of Tigris helluo (formerly Hogna helluo), a

species of wolf spider favoring specialized “interior” habitat conditions,
and (b) Pardosa milvina, a smaller, more mobile species of wolf spider
tolerant of disturbed conditions, to increasing ratios of edge to interior
habitat in an experimentally manipulated field environment. Error bars
(line) represent standard error of the mean (solid column). Means with
different letters are significantly different. Densities of Tigris helluo
declined with increasing amounts of habitat edge. Densities of Pardosa
milvina were not affected by amount of edge. (Reprinted by permission
Fig. 7.14 (a) Response of Tigris helluo (formerly Hogna helluo), a from Wiley, from S. D. Marshall, S.D., Walker, S.E., Rypstra, A.L.,
species of wolf spider favoring specialized “interior” habitat conditions, 2006. Two ecologically-divergent generalist predators have different
and (b) Pardosa milvina, a smaller, more mobile species of wolf spider responses to landscape fragmentation. Oikos 114, 241–248. # 2006
tolerant of disturbed conditions, to increasing habitat subdivision (frag- Oikos)
mentation) in an experimentally manipulated field environment. Error
bars (line) represent standard error of the mean (solid column). Means
with different letters are significantly different. Densities of Tigris helluo
declined with fragmentation. Densities of Pardosa milvina were not 7.4 Theories and Models of Loss
different in fragmented habitats. (Reprinted by permission from Wiley, and Fragmentation
from S. D. Marshall, S.D., Walker, S.E., Rypstra, A.L., 2006. Two
ecologically-divergent generalist predators have different responses to
landscape fragmentation. Oikos 114, 241–248. # 2006 Oikos) 7.4.1 Neutral Landscape Models
One approach to modeling the problem of habitat loss, frag-

mentation, isolation, and degradation is through neutral
landscape models in which arrangement of habitats is inde-
negative effects, species-specific and ecosystem-specific pendent of (i.e., “neutral” toward) biophysical processes that
differences must be considered in predicting effects of frag- shape landscapes. Neutral landscape models are attractive
mentation in specific landscapes, and likewise inform and because they can serve as “null” models or statistical
modify management decisions. Field experiments can be baselines for exploring the effects of various spatial patterns
complemented by theoretical investigations of habitat change of habitat in landscapes on population persistence and
that employ different models of habitat configuration. growth, and because they can be easily created.
Fig. 7.16 Fragmentation effects on species, community composition A., Holt, R.D., Lovejoy, T.E., Sexton, J.O., Austin, M.P., Collins, C.D.,
and ecosystem processes. For each variable, each dot represents the Cook, W.M., Damschen, E.I., Ewers, R.M., Foster, B.L., Jenkins, C.N.,
mean effect size (the ratio of the natural log (ln) mean in the more King, A.J., Laurance, W.F., Levey, D.J., Margules, C.R., Melbourne, B.
fragmented treatment/natural log (ln) mean in the non- or less A., Nicholls, A.O., Orrock, J.L., Song, D.-X., Townshend, J.R., 2015.
fragmented treatment). Horizontal bars represent range of response in Habitat fragmentation and its lasting impact on Earth’s ecosystems.
cases of multiple studies. (Reprinted under CC-BY-4.0 International, Science Advances 1, e1500052. # 2015 The Authors)
from Haddad, N.M., Brudvig, L.A., Clobert, J., Davies, K.F., Gonzalez,
In neutral landscape models, the landscape is represented fragmentation) than do those constrained to move only
as a grid (lattice) of area m2 in which some fraction ( p) of the through habitat cells. By creating movement rules using
grid cells (sites) are randomly “filled” or “occupied” with a dispersive abilities of individual species and their tolerance
type of habitat in which an organism can live and reproduce, to non-habitat (matrix) conditions, conservation biologists
thus being “neutral” to a specific structuring process. In can create models based on actual species’ perceptions of
contrast, “empty” or “matrix” cells of non-habitat are landscape structure. They can then begin to look at landscape
arranged among the “filled” habitat cells. How would an connectivity from the perspective of an individual species,
organism relate to this model landscape? To answer this and this approach permits the development of more effective
question, a neutral model can be used to provide input to a conservation strategies. As landscape ecologist Kimberly
dispersal model. In the particular case of agent-based dis- With puts it, “Determining the scale at which organisms are
persal models, the dispersal model assesses the connected- able to interact with the landscape pattern is thus the key to
ness between the model’s habitats, or, habitat connectivity of defining landscape connectivity” (With 2002a:109). One the-
the model, determined by the model’s neighborhood or oretical approach that offers a systemic understanding of the
movement rules, which specify the distance across which effects of habitat loss and fragmentation is called percolation
sites are accessible to organisms (the “agents” of the model) theory.
by virtue of their dispersal or gap-crossing abilities. Thus, the
critical threshold of habitat abundance ( pc) at which the
habitat connectedness is lost and the landscape becomes 7.4.2 Percolation Theory: Defining the Critical
“disconnected” shifts to increasingly lower levels of habitat Threshold of Fragmentation
abundance ( p) as the movement neighborhood becomes
larger. That is, pc approaches 0 as the “neighborhood size” How many red spots make a white cow red?
(the area that organisms can disperse to) approaches m2 (the How many clearings make a forest, prairie?
size of the landscape). Organisms with good dispersal A score? More? A coalescing core?
abilities should perceive landscapes as connected across a A threshold reached?
(Forman and Godron 1986)
greater range of habitat abundance (and levels of
Scientific textbooks are not usually known for good poetry,

yet this cryptic verse from Forman and Godron’s classic text, Landscape timber
Landscape Ecology, captures the essence of a critical conser- harvest pattern
vation question. At what point in habitat and landscape
fragmentation is a critical threshold reached, beyond which
a formerly contiguous habitat degrades and its species
decline? Can we predict where this point will occur and C
Maximum
prevent it?
Effect on
Over 30 years ago, Jerry Franklin of the US Forest Service patch size Forested Cutover
and the aforementioned Richard Forman of Harvard Univer- patches patches
Patch size (hectares)

sity offered a detailed analysis of the ecological
consequences of historic patterns of timber harvest in US
Pacific Northwest forests in the first issue of the (then) new
journal, Landscape Ecology. In their groundbreaking paper,
“Creating landscape patterns by forest cutting: ecological
consequences and principles,” Franklin and Forman exam-
ined projected long-term effects of the then-current manage-
ment practice of “staggered-setting clearcutting,” in which
10–20 ha patches of clear-cuts were interspersed with uncut 10
forest areas of at least equal size (Franklin and Forman 1987). 0
0 50 100
These scientists noted prophetically that “. . .the ecological
Cutover area in landscape (percent)
and economic appropriateness of this system should be
reexamined. . .evidence is accumulating that this Fig. 7.17 Changes in landscape characteristics along a timber cutting
system. . .increases the risk of some types of catastrophic gradient based on regularly distributed dispersed (checkerboard) patch
disturbance” (Franklin and Forman 1987:6). Franklin and cuttings (clear-cuts) of 10–20 ha. Note the precipitous decline in the
average patch size of uncut forests when the landscape reaches the 30%
Forman determined that “at about the 30% cutover point, cutover point (clear-cuts cover 30% of the forest). (Reprinted by per-
the average [uncut] forest patch size begins to drop mission from Springer Nature, from Franklin, J.F., Forman, R.T.T.,
sharply. . .because cuts coalesce into continuous lines of 1987. Creating landscape patterns by forest cutting: Ecological
patches dividing the previously continuous forests into consequences and principles. Landscape Ecology 1, 5–18. # 1987
SPB Academic Publishing)
sections” (Franklin and Forman 1987:8). Why does this
happen?
When less than 30% of the forest has been cut, cut areas problems of habitat fragmentation and began to incorporate it
are relatively small gaps in a contiguous forested landscape. into neutral models for describing landscape patterns
But when this critical cutover point is reached, the landscape (Gardner et al. 1987).
loses it contiguous character (Fig. 7.17). If dispersive abilities At a landscape level, percolation theory can be understood
of the organisms and integrity of ecological processes remain this way. Stochastic disturbances that disrupt linkages among
high relative to separation distances between habitats, the sites reduce the size of the percolation cluster until it
system remains “connected.” For example, in a model fragments and the system no longer percolates (flows cannot
depicting an undisturbed area, a single group of connected occur across the landscape). The level of disturbance (e.g.,
habitat patches, or “cluster,” might span the entire system the proportion of sites destroyed) at which the transition from
(i.e., there would be connectedness or “flow” between all a connected to a disconnected system occurs is called the
habitat cells). Here there is overall connectivity because critical or percolation threshold, denoted earlier as pc. The
flows can percolate across the entire system, and the span- percolation threshold is defined as the level of habitat abun-
ning cluster is therefore called the percolation cluster dance ( p) at which the probability of having a connected
(Fig. 7.18) (With 2002b). Percolation theory is the analysis landscape is less than or equal to 0.5. Above this value, the
of connectivity in spatially structured landscape systems. landscape is considered to be connected; below the threshold,
Percolation theory was first developed in the 1940s by the landscape is disconnected (With 2002b:214). Thus, a
physicists to describe physical properties of gels, polymers, percolation cluster is really a threshold at which the probabil-
and glassy materials, particularly as a means to understand ity of the landscape containing a spanning cluster goes from
the flow of liquids through material aggregates. In the 1980s near 1 (certainty) to near 0 (non-existent) over a very small
landscape ecologists Robert Gardner and Robert O’Neill of range of p. Above the threshold ( p > pc), the probability of
Tennessee’s (US) Oak Ridge National Laboratory and their having a percolation cluster is high and the largest cluster is
colleagues saw potential applications of percolation theory to the percolation cluster, which dominates the system. Below
Fig. 7.18 Percolation cluster of a

random, computer-generated
landscape, where pc, the critical
fragmentation threshold, is 0.6. As
long as 60% of the landscape
remains “habitat,” all parts of the
habitat in the landscape are
“connected,” and in sufficient
proximity to permit movement of
organisms from one portion to any
other. (Reprinted by permission
from Elsevier, from Munoz, F.,
Huth, G., Pitard, E., 2018.
Boundary constraints on
population dynamics in a
percolating habitat. Ecological
Complexity 36, 230–238.
# Elsevier B.V.)
the threshold ( p < p(c)), no percolation cluster occurs and the per cell (Flather et al. 2002). Flather et al. also defined
system consists of numerous small clusters (i.e., the system is movement rules permitting the birds’ offspring to move to
disconnected). Above this “percolation threshold,” habitat new habitat cells, but the probability of reaching a new cell
destruction results in habitat loss but not loss of habitat declined with distance from the natal cell.
connectedness. Below this threshold, additional habitat As long as habitat abundance was greater than 40%,
destruction results in increasing loss of habitat connectivity. amount of habitat had the greatest effect on population size,
accounting for 97% of the variation in abundance (Fig. 7.19a,
Table 7.3) and the probability of the population’s persistence
7.4.3 Can Percolation Theory Explain the Real approached 1.0 (Fig. 7.19b). Fragmentation levels, which
World? Models and Field Studies affected arrangement of habitat, were inconsequential. How-
ever, if the amount of habitat dropped to 40% or lower, the
7.4.3.1 Habitat-Population Models Support probability of the population’s persistence in the landscape
the Predictions of Percolation Theory dropped suddenly (Fig. 7.19b) and the degree of fragmenta-
Percolation theory predicts that transition from a connected to tion of remaining habitat became a major factor in population
an unconnected habitat will be abrupt when the amount of persistence (Fig. 7.19c). That is, the amount of habitat
habitat reaches some critical value. Is there evidence this remained the critical variable in determining the size of the
would really happen? US Forest Service scientists Curtis population at all levels of habitat availability, but the
Flather, Michael Bevers, and John Hof developed a model arrangement of habitat abruptly increased in importance if
to evaluate effects of fragmentation on a hypothetical “spe- there was less than 40% habitat in the landscape. In this
cies” of bird. Their landscape was composed of cells arranged model, it appears that pc ¼ 0.4.
in a 32 32 matrix (1024 cells) in which each cell was In models examining some of the same effects in amphib-
assigned a category of habitat or non-habitat. Flather and ian populations, Jackson and Fahrig (2014) demonstrated that
his colleagues allowed amount of habitat to vary from 10% it was proportion (i.e., amount) of habitat, not degree of
to 90%, changing in 10% increments, creating 9 levels of fragmentation, that had the largest effect on abundance (num-
habitat abundance. They also specified 9 different levels of ber of individuals), presence/absence of the population at a
fragmentation of the landscape, from highly fragmented to given site, and genetic diversity of the population. Regardless
highly aggregated, thus producing 81 different combinations of the level of fragmentation (from 0.1 to 0.9 proportion of
(99) of availability and fragmentation. Into this landscape habitat fragmented), model populations always increased in
they introduced a generic, computer-generated “bird species” genetic diversity (number of different alleles), presence/
that bred in habitat cells, defended a territory equal to the area absence, and abundance as habitat amount increased
of one cell and had a carrying capacity of one breeding pair (Fig. 7.20) (Jackson and Fahrig 2014).
Fig. 7.19 Population response of a hypothetical species to simulated persistence, with the persistence threshold (c, shaded) of the population
landscapes with specified amounts of habitat and levels of fragmenta- dropping sharply and suddenly at critical levels of fragmentation
tion. As long as 40% or more of the landscape remains in suitable (“stairstep” regions) in a landscape with reduced habitat. (Reprinted
habitat, variation in the amount of habitat accounts for 97% of the by permission from Springer Nature, from Flather, C.H., Bevers, M.,
variation in population size (a) (See Table 7.3). When the amount of Hof, J., 2002. Prescribing Habitat Layouts: Analysis of Optimal Place-
habitat in the landscape drops below 40%, probability of population ment for Landscape Planning, in: Gutzwiller, K.J. (Ed.), Applying
persistence declines rapidly. (b) Below this level of habitat, degree of Landscape Ecology in Biological Conservation. Springer New York,
fragmentation now becomes an important factor in population New York, NY, pp. 428–453. # 2002 Springer-Verlag New York, Inc)
Table 7.3 Summary Analysis of Variance findings for simulated landscapes evaluated in a 99 factorial experiment with nine levels of habitat
abundance (10–90%) and 9 levels of fragmentation (0.1–0.9) above and below a persistence threshold (see Fig. 7.19). DF ¼ degrees of freedom
Source of variation DF % of Total SS F P
Full experiment
Habitat amount 8 96.8 15,871.5 0.0001
Fragmentation 8 0.7 124.4 0.0001
Habitat amount fragmentation 64 0.6 13.3 0.0001
Error 2349 1.8 – –
Above threshold
Habitat amount 6 96.3 10,497.2 0.0001
Error 1653 2.5 – –
Below threshold
Habitat amount 3 30.3 122.3 0.0001
Error 696 57.4 – –
Reprinted by permission from Springer Nature, from Flather, C.H., Bevers, M., Hof, J., 2002. Prescribing Habitat Layouts: Analysis of Optimal
Placement for Landscape Planning, in: Gutzwiller, K.J. (Ed.), Applying Landscape Ecology in Biological Conservation. Springer New York,
New York, NY, pp. 428–453. # 2002 Springer-Verlag New York, Inc
These studies indicate that both habitat loss and fragmen- of habitat available. Based on the results of these and other
tation can affect populations, but that by far the more impor- studies, Lenore Fahrig has proposed the habitat amount
tant factor is habitat loss, i.e., it’s not the size or arrangement hypothesis – that species richness in equal-sized sample
of the habitat patches that matters as much as the total amount sites should increase with the total amount of area within an
Fig. 7.20 Mean local abundance (number of individuals), presence/ permission from Wiley, from Jackson, N.D., Fahrig, L., 2014. Land-
absence (or population) and allelic richness (number of diffent alleles) scape context affects genetic diversity at a much larger spatial extent
plotted against amount (proportion) of habitat at different levels of than population abundance. Ecology 95, 871–881. # 2014 Ecological
fragmentation (1 – H, 0.1–0.9) in a model landscape. (Reprinted by Society of America)
appropriate distance to the sample site (the “local landscape”) because arrangement doesn’t matter. Just measure the
regardless of the configuration of habitat within the land- amount of habitat!
scape (Fahrig 2013, 2017). If this hypothesis is true, patch There is support from field studies for Fahrig’s hypothesis.
size and patch isolation would no longer be variables that Many studies have shown that patch boundaries don’t limit
predict species diversity, and therefore need not be measured. movements of many populations, but that many animal spe-
It would also be unnecessary to distinguish between patch- cies move freely between them (Petranka and Holbrook
scale effects and landscape-scale effects or between patch- 2006; Roe et al. 2009; Schultz et al. 2012). And other studies
size effects and patch-isolation effects. And it would no have shown that the amount of site-specific species richness
longer be necessary to evaluate the configuration (arrange- increases with the amount of habitat in the local landscape
ment) of habitat independently of the amount of habitat (Laurance et al. 2002; Fahrig 2013). If this is true, there is
nothing special about patches of habitat that requires one to contrast, were, both scale- and species-specific, becoming
measure extinction-colonization dynamics of habitat at the the most important factor in one species, the North American
level of patches. Distinctions between patch-scale habitat veery (Catharus fuscescens), at small radii (500 m), but
effects and landscape-scale habitat effects on species diver- declining in importance at larger scales (Smith et al. 2011).
sity would not be necessary. These processes and their effects Such studies show us that species and populations are
need only be measured at the landscape level (Fahrig 2013). affected by habitat configuration and fragmentation, but
Nevertheless, as Fahrig herself admits, “habitat amount through what mechanisms? Peng He and his colleagues at
isn’t everything” (Fahrig 2013:1657). Many studies show the Max Planck Institute for Ornithology (Germany)
that the degree of connectivity between habitat patches hypothesized that habitat configuration works in part through
matters. Such connectivity may be most influenced by a social processes, interactions, and structures of animal
species’ dispersive abilities because population persistence populations. Distribution of habitat components affect
in fragmented landscapes depends on the ability to move where animals move by alternatively attracting or repelling
between habitat patches and that probability of successful them to specific sites and increasing or decreasing frequency
movement (immigration and emigration, Chap. 6) is a reflec- of social encounters (He et al. 2019). In a study on Australian
tion of patch connectivity (Ranius et al. 2010). Connectivity sleepy lizards (Tiliqua rugosa, Fig. 7.21), Leu et al. (2016)
reflects potential for dispersal, and ability to disperse manipulated the structure of the lizard habitats by placing
influences ability to persist. Here is one example. 100 8 m fences throughout one area but leaving another
The US city of Chicago and its surrounding suburban area unfenced. Using information obtained from GPS locators
is a highly modified environment with almost no natural attached to the lizards, Leu et al. determined that lizards in
vegetation remaining that is not protected in conservation fenced areas had higher social connectivity (more encounters
reserves. Within these forest preserves, native amphibian with other lizards) than those where there were no fences.
species make extensive use of permanent and seasonal forest They hypothesized that fences increased connectivity
pools for breeding. Most amphibian species, however, have because lizards moved around fences to resume their original
limited dispersal abilities, which makes it difficult for them to path, causing them to (1) cover greater distances in daily
move long distances to different pools even within the same movements and (2) follow similar paths between fences.
reserve, and the hostile nature of the environment between Leu et al. concluded that “level of habitat structural complex-
preserves (concrete and asphalt surfaces with heavy volumes ity can modulate the social structure of a nongregarious
of human vehicular traffic) make movement from one pre- species” (Leu et al. 2016:27).
serve to another not merely difficult but dangerous. In this Habitat configuration could interact with processes affect-
landscape, Van Dyke et al. (2017) investigated the effect of ing movement, social structure, social organization, and pop-
habitat clustering (the occurrence of high quality habitats in ulation dynamics (Fig. 7.22). Such a conceptual framework
close spatial proximity to one another) on the species diver- permits generation of predictions and hypotheses of how
sity of amphibian communities. In cases where pools were habitat configuration could affect habitat components with
clustered within 100 m of one another, species richness of the consequent expectations on social interactions and network
amphibian community was higher than in isolated pools structure (Table 7.4) (He et al. 2019). In conservation efforts,
(single pools more than 500 m from the nearest neighboring are there examples of habitat configuration, as well as habitat
pool). Even when differences in open water (breeding) area amount, interacting with social processes?
were accounted for between clusters and isolates (habitat
amount), differences remained. Amphibians were also more
likely to colonize clustered pools than isolated pools and less
likely to experience site-specific extinctions in clustered
pools. Species richness was also affected. All (10) local
amphibian species were resident in clustered pools, but only
four species in isolated pools. (Van Dyke et al. 2017). Con-
nectivity mattered.
Fahrig’s habitat amount hypothesis, as stated, only applies
to species diversity. Are other aspects of populations affected
by fragmentation? In a study of the relative importance of
habitat amount, habitat fragmentation, and matrix quality on
occurrence of forest birds in Ontario (Canada), habitat
Fig. 7.21 The Australian sleepy lizard (Tiliqua rugosa), a species
amount was more important than matrix quality or fragmen- whose social structure can be influenced by habitat complexity that
tation over a range of 500 m–10 km in radius and 1–300 km2 affects the frequency of encounters between individuals. (Photo cour-
in area (Smith et al. 2011). Effects of fragmentation, in tesy of B. Twist)
Fig. 7.22 A conceptual framework to illustrate the relationship structure and mating patterns which can drive changes in population
between habitat configuration, animal movement, social behavior and demography as well as alter ecological and evolutionary processes.
demography. Changes in amount and spatial arrangement of abiotic and (Reprinted under CC BY 4.0 International, from He, P., Maldonado-
biotic components of habitat can affect patterns of movement at different Chaparro, A.A., Farine, D.R., 2019. The role of habitat configuration in
spatial scales, including changes in patterns of habitat use, dispersal and shaping social structure: a gap in studies of animal social complexity.
migration, which can then facilitate or restrict social interactions among Behavioral Ecology and Sociobiology 73. # 2019 The Authors)
individuals, thus influencing key aspects of sociality such as social
Table 7.4 Predictions and hypotheses concerning how biotic and abiotic components of the habitat can shape animal movement, social
relationships among individuals, and social network structure
Expected effects on social network
Expected effects on movement Components Expected effect on social interactions structure
Behaviorally Attract Resources Higher rates of repeated social interactions More defined network communities and
shaping movement where (or when) resources are more higher modularity when resources are more
decisions (why and clustered clustered resulting in higher mean
when to move) association strength and density within
communities
Mates Fewer mates trigger more agonistic Stronger edge weight in male-male
interactions among males (or females) (or female-female) interaction networks
and higher density in sexual networks
Repel Competitors Higher levels of competition generate more Stronger edge weight in directed agonistic
agonistic interactions among individuals interaction networks
Predators Greater predation pressure often increases Higher mean degree in riskier habitats
group size
Physically shaping Facilitate Corridors Presence of preferred movement paths Increase network density and greater
movements (how roads (habitat corridors, animal tracks, roads) assortative mixing
and where to move) increase frequency of encounters among
individuals
Impede Barriers (e.g., Reduced frequency of social encounters by Increased network clustering and
fences, roads) individuals restricted to different patches community structure, resulting in higher
Habitat and greater rates of social encounters modularity
transformation among individuals in the same patch
Reprinted under CC BY 4.0 International, from He, P., Maldonado-Chaparro, A.A., Farine, D.R., 2019. The role of habitat configuration in shaping
social structure: a gap in studies of animal social complexity. Behavioral Ecology and Sociobiology 73. # 2019 The Authors
7.4.3.2 The Spotted Owl: Population Predictions habitat availability (old growth forests) using movement
and Conservation Planning rules determined from actual dispersal abilities of juvenile
The northern spotted owl (Strix occidentalis caurina) is an owls. After repeated simulation experiments, the researchers
endangered subspecies that breeds primarily in old-growth came to a disturbing conclusion. Sharp extinction thresholds
forests in the US Pacific Northwest. Conservation scientists appeared at the point where old-growth habitat was reduced
who have studied the biology of the owl, and in particular its to ~20% of the landscape. At that level, the ability of
reproductive ecology, were able to determine the dispersal juveniles to locate suitable territories (a social interaction)
distances of juvenile owls during periods when they left their was reduced, leading eventually to population extinction.
natal areas and traveled in search of new areas to establish These results led to recommendations for the following habi-
breeding territories. Based on such investigations, they tat design criteria: individual habitat areas should be large
constructed a landscape-level model that could evaluate the enough to support at least 20 breeding pairs
probability of population persistence at different levels of (a recommendation of habitat amount); habitat areas should
7.5 Conservation Through Protected Areas 289
be no farther than 19 km apart (a recommendation of habitat patches essential for the maintenance of landscape connec-
configuration); and the matrix between habitat areas should tivity. Assuming a maximum dispersal range of 45 km for
be at least 50% forested with tree diameters >28 cm and with juvenile owls, the connectivity of forested habitat in this
canopy closure >40% (a recommendation of habitat connec- region is tenuous, at best (Fig. 7.23a). At this dispersal
tivity to facilitate social interaction) (Noon and McKelvey distance, only one patch linked the southwestern and north-
1996). eastern parts of its range, old growth forests in the US Forest
Notice how similar the findings of this species-specific Service’s Mount Taylor Ranger District in the Cibola
model are to the findings of the model developed by Flather National Forest in New Mexico (Fig. 7.23b). Analysis
et al., how closely such findings match predictions of perco- revealed that the Mount Taylor Ranger District was, in fact,
lation theory, and how such findings recognize the role of a crucial stepping stone connecting owl populations in
social interaction in both individual and population response. Arizona and New Mexico to those in Colorado and Utah.
Also observe that the amount of habitat preserved in reserve This was not obvious prior to the analysis of landscape
design was close to the theoretical persistence threshold connectivity. In fact, the Mount Taylor District had previ-
defined by the model. What would happen to owl populations ously been considered unimportant habitat because it
if there was a catastrophic event that reduced habitat avail- supported so few resident Mexican spotted owls.
ability below 20%? This was a stochastic model, and its
outcomes reflected uncertainties specified within a defined
range of probabilities for demographic and environmental 7.5 Conservation Through Protected Areas
events. Suppose, within that uncertainty, that a given period
of time proved to be “unusually unlucky” for owls in terms of 7.5.1 Algorithms of Reserve Design
demographic and environmental stochasticity. Living on the
edge of their persistence threshold of 20% available habitat, a Where large areas of natural habitat still exist, the most direct
series of “bad luck” events in demographic or environmental strategy for habitat conservation is the establishment of
stochasticity could send the population “over the edge” into protected areas. All levels of government, from international
extinction. to local, as well as thousands of private conservation
In cases where habitats harbor resources of high value to organizations, pursue setting aside preserves as the best
humans, such as timber in an old growth forest, final conser- way to conserve habitat and landscape and the species they
vation plans may be determined as much by political pro- contain. And much land has been set aside, almost 15% of the
cesses as scientific modeling, in which case concerns about world’s total surface area by 2014 (World Bank 2016). Not
persistence thresholds and percolation theory are ignored. all “preserved areas” are equally well protected, but the
But important insights from such modeling must not be number and area of preserves continues to grow. What
lost. Some of the models we have already examined, and methods are used to select protected areas, and how does
the insights derived from them (Fahrig 1992, 2003; Flather the decision-making process work? The Nature Conservancy
et al. 2002) suggest that, where habitat loss is substantial, (TNC) provides one example.
effects of fragmentation become significant to population TNC is one of the world’s most influential
persistence, and such effects will occur suddenly when the non-governmental organizations in conservation,
“persistence threshold” is attained, although the exact level of administering the largest system of private nature reserves
that threshold is difficult to determine and varies by species. in the world with holdings on six continents. The problem
When that threshold is reached, fragmentation and isolation TNC and similar conservation organizations want to solve
do matter, and conservation efforts to achieve optimal can be stated simply: how can we best locate reserve units on
arrangement of remaining habitats become important. The a landscape in such a way that they contain the largest
northern spotted owl provides an example in which a possible number of biodiversity elements? To solve this
species-specific habitat suitability model, developed from problem, TNC follows a three-stage process of reserve sys-
actual field studies, made predictions and generated outcomes tem planning: 1) many sites are screened as potential
consistent with those of percolation theory. But can percola- reserves, (2) candidate sites are examined for their promise
tion theory be applied in the planning stages of a conservation as part of a functional system of reserves, (3) individual
effort, such as a species recovery plan? reserve sites are established, managed, and monitored – a
One such application was the inclusion of a percolation- process that continues for the life of the reserve system
based analysis of habitat connectivity in the recovery plan for (Urban 2002). How would a site merit consideration for
the northern spotted owl’s taxonomic cousin, the Mexican step one? Selection criteria used by TNC are (1) ecological
spotted owl (Strix occidentalis lucida) (Keitt et al. 1995). In uniqueness – what is site’s level of species richness, rarity,
addition to assessing habitat connectivity, this application of and endemism, or other ecological features of special con-
percolation theory led to identification of critical habitat cern; (2) viability – what is the likelihood that species would
Fig. 7.23 (a) Relationship of

maximum dispersal distance
(km) of the Mexican spotted owl
(Strix occidentalis lucida), and
connectivity distance (km), the
maximum distance between
“connected” habitats. Note that at
dispersal distances below 45 km,
connectivity distance drops
sharply, indicating that, to owls,
available habitats separated by
more than 150 km from one other
are effectively “disconnected.” (b)
Location and strategic importance
of the Mount Taylor Ranger
District in central New Mexico
(USA). Although having a low
number of Mexican spotted owls,
the Mount Taylor Ranger District
is the critical “stepping stone”
creating connectivity of habitats
for owls in the four state area of
Utah, Colorado, Arizona, and
New Mexico, without which the
northeastern and southwestern
populations lose connectivity with
one another. (Reprinted by
permission from Springer Nature,
from With, K.A., 2002. Using
Percolation Theory to Assess
Landscape Connectivity and
Effects of Habitat Fragmentation,
in: Gutzwiller, K.J. (Ed.),
Applying Landscape Ecology in
Biological Conservation.
Springer, New York, NY, pp.
105–130. # 2002 Springer New
York; Photo courtesy of US Fish
and Wildlife Service, public
domain
persist on this site if it were protected as a reserve; (3) threats – likelihood of dispersal among sites by minimizing distances
what agents might reduce the long-term viability or value of between sites or by providing stepping stone habitats or
the reserve; and (4) feasibility – what are relevant economic, dispersal corridors between sites. In this algorithm, note
sociological, and administrative factors related to likelihood how concepts of percolation theory are implicitly invoked
that reserve could actually be acquired and protected. This is to increase connectivity of the landscape and the contagion of
the “can we really do this” criterion. A site that is unafford- preserves to one another. Also note that these three
able, unavailable, or engenders hostility from local residents algorithms focus on two criteria, namely richness (the first
will fail the feasibility test. two algorithms) and viability (connectivity algorithm).
If a site meets all criteria, it becomes one member of a
portfolio of potential sites. For all its worldwide scope and
influence, TNC’s resources are limited. It cannot acquire
Consider two sites in your area that could have conser-
every site, and it must make acquisitions that form a network
vation value but are currently not protected. How
of reserves that complement one another in conservation
would these areas score using TNC’s criteria? Which
objectives. To aid its decision making process, TNC employs
area would rank higher, and why?
three algorithms for site selection. (1) The Greedy Richness
Algorithm – Get the most species with fewest sites. (2) The
Greedy Rarity Algorithm – Get the rarest species first, then
add increasingly common species after rarest ones are If we employ these algorithms to select the “right” sites for
accounted for. (3) The Connectivity Algorithm – Provide for conservation preserves, will our choices achieve conserva-
landscape scale population resilience by maximizing the tion goals? To answer with insight, we must consider further
criteria and technological innovation in reserve design. One GAP is designed to provide an information base enabling
of the most frequently employed is GAP analysis. managers and planners to make the best and most efficient
use of land in establishing reserves by showing where con-
servation efforts should be focused to achieve maximum
7.5.2 GAP Analysis and Reserve Design biodiversity or protection for endangered species. With the
aid of its GIS applications and technology, GAP can display
One of the most comprehensive efforts in reserve design and relationships of interest at varying cartographic scales,
conservation planning is the ongoing Gap Analysis Program, distributions of individual species or entire suites of species,
now often referred to simply as GAP or GAP analysis. and overlay maps of species distributions with different
Originally developed in the early 1990s by J. M. Scott and jurisdictions of land ownership and management objectives.
others at the University of Idaho (USA), GAP uses satellite A criticism of GAP analysis is that it often assumes that all
imagery and GIS technology to make computer-generated habitat of a specific type will support species that use such
regional maps of the distribution of dominant vegetation or habitat, but this is not always the case. Variables such as
geographic distributions of animals species and then relate habitat configuration (especially ratios of edge to interior
these distributions to existing conservation reserves (Scott habitat), habitat connections and connectivity, and interspe-
et al. 1993). More specifically, GAP analysis determines, cific interactions are important determinants of habitat use
through the use of such computer overlay maps, whether and population persistence. Traditional GAP analysis did not
populations of species targeted for conservation fall within consider these elements, but more recent GAP analysis
the boundaries of currently protected areas (Opdam 2002) efforts do (Santini et al. 2014; Fig. 7.24). Some critics have
and which elements of landscape biodiversity are underrep- argued that the inventory and monitoring of biotic resources
resented in reserve systems. GAP’s sequential tasks are to: on a national or international scale require a sampling uni-
(1) map existing vegetation to the level of dominant or verse of broad, landscape-scale assessments. GAP, in con-
co-dominant species (from satellite imagery); (2) map trast, has typically relied on small-scale, low-resolution
predicted distributions of vertebrate species (using museum assessments of biodiversity, so is not always ideal for
and agency collection records with existing general range identifying larger areas that should become candidates for
maps of each species); (3) map public land ownership and biological protection. But it is possible to combine techniques
private conservation lands; (4) show the current network of of GAP analysis with understanding of species-specific habi-
conservation lands (the combined distribution of public lands tat suitability models at larger scales.
and private conservation lands); (5) compare distributions of
native vertebrates and vegetation communities with the net-
work of conservation lands; and (6) from this comparison, 7.5.3 Reserve Design and Habitat Suitability
provide an objective basis of information for options in
managing biological resources. That is, GAP’s final outcome To use or not to use; that is the question every species asks
is to attempt to identify “gaps” in the conservation reserve when presented with an array of habitat choices, and conser-
network: where in the landscape are important biodiversity vation biologists try to see the world through the eyes of
resources not protected? non-human species to anticipate which habitats are worth
Fig. 7.24 Sequence of steps in a GAP analysis defining networks of Wiley, from Santini, L., Marco, M.D., Boitani, L., Maiorano, L.,
protected clusters of suitable habitat, an example of how recent efforts in Rondinini, C., 2014. Incorporating spatial population structure in gap
GAP analysis consider habitat configuration and connectivity to identify analysis reveals inequitable assessments of species protection. Diversity
species needing conservation protection. (Reprinted by permission from and Distributions 20, 698–707. # 2014 John Wiley & Sons Ltd)
preserving. Can previously discussed habitat suitability reserve system, 2.90 million km2 to conserve 10% of suitable
models be used at larger landscape scales and broader geo- areas, and 3.36 million km2 to conserve 10% of highly
graphic contexts to identify appropriate locations and suitable areas (Fig. 7.25). As the authors themselves noted
dimensions of proposed reserves, and can these models accu- “This last figure would mean nearly doubling the existing
rately evaluate the effectiveness of existing reserves in biodi- reserve network in Africa” (Rondinini et al. 2005).
versity conservation? Biologists Carlo Rondinini, Simon This analysis revealed that the reserve system in Africa
Stuart, and Luigi Boitani of IUCN addressed this problem was not sufficient to include even a minimal amount of the
in an evaluation of conservation planning for African range of all mammals and amphibians. It also suggested that
vertebrates (Rondinini et al. 2005). They were able to devise planning discussions should be shifted from arguments about
a plan for a systematic reserve selection for 1223 African the size of the reserve to analysis that more precisely
mammals and amphibians in which habitat suitability models determines the nature of the target. That is, managers should
were used as estimates of the area occupied by each species. look for the species-specific quality of what they are trying to
A standard conservation goal in reserve design is to con- protect (i.e., the right kind of habitat) rather than the quantity
serve 5–10% of present species ranges within the reserve of land area in the proposed reserve.
system. First Rondinini et al. wanted to find out if, in fact,
the present reserve system in Africa really did that for
mammals and amphibians. Using data from the IUCN’s 7.5.4 Determining Appropriate Reserve Size
Global Amphibian Assessment and Global Mammal Assess-
ment, they determined the geographic range (extent of occur- Although habitat quality is an increasing concern in reserve
rence) and habitat preferences for each of these 1223 species. design, the question of how large a reserve should be remains
They then used the habitat preference data to construct habi- a critical issue in conservation biology. If the purpose of the
tat suitability models inside each species geographic range, reserve is to ensure persistence of a particular species or
and for 181 species, verified the models by comparing suit- group of species, then the needs of such species become the
ability levels to presence-absence data collected in the field. operative criteria for reserve size. If time, expertise and
They used the suitable areas as estimators of the area of money permit, species within the proposed refuge, or at
occupancy and compared these to results of systematic least those of highest priority for conservation, should be
reserve selection based on geographic ranges. subject to a population viability analysis (Chap. 6) which
This was an ingenious and inventive use of habitat suit- can reveal the average minimum viable populations (MVPs)
ability models, previously discussed in our examples of for important refuge species. These MVPs, divided by
HSMs for the giant panda and corncrake, in conservation estimated population densities, should yield the quotient of
planning and assessment. Unfortunately, the results were minimal area for population persistence (Simberloff 1988).
discouraging. When Rondinini et al. compared their habitat- With minimal area estimates in mind, Soulé and
based approach to the traditional method of determining Simberloff (1986) devised what has become a classic three-
refuge placement by overlap with species’ geographic ranges, step approach to the problem of estimating the optimal size of
they found that geographic ranges overestimated the actual a conservation reserve or collection of reserves. The first step
area occupied by species and underestimated the amount of was to identify species whose disappearance would signifi-
area that needed to be conserved. Further, their analysis cantly decrease the value of the reserve or its diversity,
revealed that every protected area in Africa contained fewer including threatened and endangered species, species of
species than predicted by analysis of geographic ranges. high public visibility or aesthetic appeal, species whose
Because species are more specialized than estimates of geo- abundance provides an index of habitat quality (“indicator
graphic distribution suggest, underestimation of land areas species”) and species that create habitat or perform functions
needed for reserves is probably not unique to Africa, but a that enhance populations of other species (“keystone spe-
systemic problem in global conservation. cies”). Second, determine the minimum number of
Based on their habitat suitability analysis, Rondinini and individuals needed to guarantee a high probability of sur-
his colleagues determined that the reserve system in Africa vival. Third, using known densities, estimate the area needed
would need a 30–100% expansion to achieve minimal con- to sustain this minimum population.
servation targets. At the time of their analysis, existing The dilemmas of these choices emerge in practical ways
protected areas covered 3.44 million km2, or 10% of the when planners can, or must, choose between making the
African landmass surface. But to achieve the 5% target for reserve a single large area or several smaller reserves of
geographic ranges, the current reserve system would require approximately equal area (more broadly known as the single
an increase of 1.11 million km2 (32%). The same target large or several small (SLOSS) debate). Determining the best
would require an increase of 2.36 million km2 to conserve choice depends upon several factors. First, knowing that
5% of suitable ranges and 2.85 million km2 for highly suit- smaller reserves will support smaller populations, one must
able ranges. Achieving the 10% target for geographic range determine the difference between extinction probabilities
would take an addition of 1.73 million km2 to the African associated with large and small populations of the most
a b
c d
Protected areas
Existing
Additional selected
0 2,000 Km
Fig. 7.25 Location of sites selected under four scenarios of reserve meeting conservation goals, would require the current African reserve
design for African mammals and amphibians. (a) 5% of geographic system to be increased by nearly 100%. (Reprinted by permission from
ranges represented in reserves. (b) 5% of highly suitable habitat areas Wiley, from Rondinini, C., Stuart, S., Boitani, L., 2005. Habitat Suit-
represented in reserves. (c) 10% of geographic ranges represented in ability Models and the Shortfall in Conservation Planning for African
reserves. (d) 10% of highly suitable habitat areas represented in Vertebrates. Conservation Biology 19, 1488–1497. # 2005 Society for
reserves. The fourth scenario (d), although likely the most effective in Conservation Biology)
important species. Large differences favor a single large each small reserve will not contain every species that might
reserve, while small differences argue for several small be present in the single large reserve? Third, what is the
reserves. A second key question is: how many populations correlation in the year-to-year fluctuation of the environments
will a series of small refuges preserve, since, presumably, of the populations in the proposed small reserves? If
environmental variation is independent (uncorrelated), multi- patches, and they should contain separate minimum dynamic
ple reserves provide protection against chance environmental areas of each habitat type. Additionally, the reserve should
disturbances or catastrophes that could reduce or exterminate include internal sources for repopulating local extinctions
a single population. On the other hand, if the separate and should include different ages of disturbance-created
reserves have a high degree of environmental correlation, patches (Pickett and Thompson 1978).
they confer no such advantage. Even large reserves may not preserve diversity of habitat
Determining reserve size also must consider factors in a regional landscape, especially if the selection process is
associated with the reserve’s habitat heterogeneity and insensitive to the realities of patchy habitat distribution.
patch dynamics. To preserve both species and habitats, Using a GAP analysis approach, Wright et al. (1994) exam-
reserves must be larger than the size of the largest ined four areas proposed as future national parks in the US
disturbance-created patch or “minimum dynamic area”, state of Idaho (Fig. 7.26). Although large, averaging
even including the rarest kinds of disturbance-created 220,000 ha each, the four proposed areas added little to the
Fig. 7.26 Four areas in Idaho

(USA) proposed for protection as
future national parks. Although
large and inclusive of unique
geologic formations, the areas add
little to the number of vegetation
types under protection and none
protect even 10% of the
vegetation types in the ecoregion.
Wiley, from Wright, R. Gerald,
James G. MacCracken, and Joel
Hall. 1994. “An Ecological
Evaluation of Proposed New
Conservation Areas in Idaho:
Evaluating Proposed Idaho
National Parks.” Conservation
Biology 8 (1): 207–16. # 1994
Society for Conservation Biology)
7.6 Preserving Habitats in Human-Modified Landscapes 295
number of different vegetation types already protected, and modified by human use and activity, is often not completely
none met even the modest goal of protecting 10% of vegeta- hostile to other species. Many species can obtain resources
tion types in the ecoregion (Wright et al. 1994). Even if areas from the matrix which aid their persistence in their current
were expanded, they did little to increase the preservation of habitat fragment and many can move through the matrix for
habitat and vegetation types. Alternatively, current proposals the purpose of dispersal to and colonization of neighboring
for preserving habitat in the same area could have increased fragments. Managing matrix habitat may be the key to saving
the number of habitats preserved with relatively few ha added many of the species that remain in the landscape. What would
to their land area. happen if we shifted emphasis from managing habitat
Smaller reserves might better maintain the variety of fragments to managing the matrix?
habitats protected, but conservationists have traditionally Matrix habitat has usually been subjected to more modifi-
assumed that smaller reserves would have negative effects cation by humans than habitat in remaining fragments.
on population levels of some species. This is not always the Ecologists have growing evidence to support the so-called
case, however. For example, Tscharntke et al. determined “intermediate disturbance hypothesis” which asserts that the
that, in central European grasslands, 10 ha of protected area highest levels of biodiversity will be associated with sites,
comprised of 29 small, individual, and separated grassland habitats and landscapes subject to intermediate levels of
remnants contained more species than the same amount of disturbance in terms of frequency and intensity. Today
area contained in one or two large habitat preserves humans are the world’s most common and powerful agents
(Tscharntke et al. 2002), findings that support Fahrig’s of ecological disturbance, intermediate or otherwise (Chap. 3).
assertions associated with the habitat amount hypothesis Intensively disturbed landscapes, such as those subject to
(Fahrig 2013, 2017). high levels of human disturbance and modification, show
For all the abstract intricacies that theories of reserve lower levels of biodiversity than intermediately disturbed
design may generate, conservation biologists must never areas. No conservationist is surprised at that. What is
forget that, in the real world, the most common size for surprising is that intermediately disturbed areas also show
reserves is 10–30 km2 (Bolton 1997). Regardless of whether higher levels of biodiversity than undisturbed areas. This
or not such reserves are adequate for conserving biodiversity pattern can be observed more precisely in the relationships
on a theoretical basis, they still represent resources and between biodiversity and specific kinds of human-modified
opportunities for conservation. With intensive and intelligent landscape structures, especially landscapes dominated by
management, small reserves can make important agricultural use and commercial forestry.
contributions to conservation efforts. But whether a proposed
reserve is large or small, plans for refuge design must be
informed by ecological data or they may have little value in 7.6.2 Conservation in Agricultural
preserving biodiversity. and Commercially Forested Landscapes
Today almost 40% of the earth’s terrestrial land area is used

7.6 Preserving Habitats in Human-Modified in some way for agricultural production, making croplands
Landscapes and pastoral lands the most common and widespread habitat
on the planet. Faced with the daunting task of trying to
7.6.1 Intermediate Disturbance: Where Does preserve biodiversity in a highly modified agricultural land-
Conservation Matter Most? scape, conservationists often default to a mindset that the
only way to achieve conservation goals is to preserve natural
In the early years of conservation biology, conservation areas from future agricultural production, restore unneeded or
biologists often applied the theory of island biogeography unproductive agricultural areas to natural conditions and, to
to habitat fragments, perceiving the fragments as “islands” compensate for land lost to production, intensify agricultural
and the matrix as an inhospitable “sea.” Such logic predicted production on remaining areas. But an alternative vision,
that species richness would decrease with decreasing area of which could make conservation practices applicable to
fragments (“islands”) and increasing distances (isolation) much larger areas, would be to manage interactions between
between the fragments (MacArthur and Wilson 1967). This remaining natural or semi-natural areas in agricultural
view was easy and attractive to apply, but incorrect in its landscapes and the surrounding matrix of pastoral lands and
foundational assumptions and often wrong in the predictions croplands. Such a change in emphasis would shift conserva-
it generated. As we have already seen, the matrix, although tion effort from that of continuing to find and protect
Fig. 7.27 Visual depiction of the intermediate landscape complexity relation to landscape structure. (Reprinted by permission from Wiley,
hypothesis. In agricultural landscapes of Central Europe, (a) the differ- from Tscharntke, T., Tylianakis, J.M., Rand, T.A., Didham, R.K.,
ence in biodiversity and ecosystem services between managed and Fahrig, L., Batáry, P., Bengtsson, J., Clough, Y., Crist, T.O., Dormann,
unmanaged areas was greatest in “simple” (intermediate complexity) C.F., Ewers, R.M., Fründ, J., Holt, R.D., Holzschuh, A., Klein, A.M.,
landscapes consisting of 1–20% noncrop area compared to “cleared” Kleijn, D., Kremen, C., Landis, D.A., Laurance, W., Lindenmayer, D.,
(low complexity) landscapes of <1% noncrop area or “complex” (high Scherber, C., Sodhi, N., Steffan-Dewenter, I., Thies, C., van der Putten,
complexity) landscapes containing more than 20% noncrop area. The W.H., Westphal, C., 2012. Landscape moderation of biodiversity
S-shaped curve of this relationship shows that maximum biodiversity patterns and processes – eight hypotheses. Biological Reviews 87,
changes induced by management actions occur in simple landscapes. (b) 661–685. # 2012 The Authors)
Effectiveness (difference between managed and unmanaged sites) in
remaining natural areas (work that is, in many parts of the conservation should be preservation of habitat through estab-
world, finished), to one of creating an agricultural landscape lishment of protected areas. But, in simplified landscapes like
that permits movement between fragments for most species. those in agricultural areas, agri-environmental management
Might conservation efforts be better applied toward that makes the agricultural “matrix” more attractive for native
modifying agricultural practices to mitigate negative effects species by mitigating negative effects of agricultural
of intensive agriculture rather than creating a dichotomy of practices will have disproportionately larger effects on biodi-
preserved (non-agricultural) lands and non-preserved (full versity and environmental processes and can be applied to
commercial production) lands? more extensive areas than a strategy of preservation
In an analysis of landscapes in agricultural lands in central (Tscharntke et al. 2012).
Europe, Tscharntke et al. found that “management effi- Similarly, commercial forests, including both natural
ciency” (conservation efforts intended to increase biodiver- forests used for wood production and “plantation forests”
sity in the landscape) was highest (generated the greatest cultivated solely for wood products, are now estimated to
response) in “simple” landscapes (landscapes of intermediate cover over 264 million ha of the earth’s surface, about 14% of
structural complexity with 1–20% noncrop land area), but the planet’s land area, about the same percentage as devoted
had relatively little effect in both “cleared” landscapes to nature preserves. Plantation forests, in particular, are not
(monotypic agriculture with less than 1% noncrop land only planted for wood production, but to reforest deforested
area) and “complex” landscapes (more than 20% noncrop sites or abandoned agricultural areas that do not otherwise
land area dominated by natural vegetation) (Fig. 7.27). In succeed to natural vegetation. Such reforestation can initiate a
the Netherlands, for example, which has less than 1% of positive feedback loop of successional processes, as seed-
natural vegetation remaining (a “cleared” landscape), efforts dispersing species of wildlife, attracted to the plantation
to increase biodiversity through altered agricultural practices forest, begin to import seeds from surrounding natural forests
had no effect. In contrast, bird abundance and diversity were (Brockerhoff et al. 2008). Although not achieving conserva-
higher in simple agricultural landscapes (for example, organ- tion values as great as native forests managed solely for
ically cultivated fields) across Central Europe than in cleared conservation, plantation and commercial forests can add sig-
or complex landscapes in the same region (Tscharntke et al. nificant conservation value to landscapes in which they occur
2005, 2012). (Fig. 7.28) (Brockerhoff et al. 2008).
Powerful implications emerge from these and similar For example, commercial and plantation forests can
findings. Where natural, complex vegetation (and corre- “buffer” negative effects of habitat fragmentation by forming
spondingly complex habitat structures) exist, the goal of a matrix between fragments of natural forests that increases
connectivity between these fragments for plants and animals

(Fig. 7.29), or even as habitat that can be used by such species
(“interstitial habitat”). This connective effect of commercial
and plantation forests has been shown to have facilitated
movements between forest fragments in endangered species
such as the Iberian lynx (Lynx pardinus) (Ferreras 2001)
(Fig. 7.30) and in many species of Australian marsupials
(Lindenmayer et al. 1999). When planted so as to border
native forests, plantation forests also can lessen impacts of
edge effects, for example, reducing microclimate effects of EI
by 50% compared to the same effects on edges between
native forests and pastures (Denyer et al. 2006). As commer-
cial and plantation forests age, they become more diverse
biologically and more complex ecologically as they are
increasingly colonized by species from surrounding native
forests (Barlow et al. 2008). As more countries and private
wood production companies adopt standards of sustainability
and biodiversity set by the Forest Stewardship Council or
other international organizations, such increases in biodiver-
sity and ecological complexity have become more common
and more normative in commercial forest production. For
example, China and the United States (which possess, respec-
tively, the largest and second largest production plantation
forests in the world) have plantations consisting mostly of
native species.
Fig. 7.28 Conceptual model of the relative conservation value of

planted forests relative to conservation forests and intensive agriculture.
Some plantation forests serve multiple purposes on the same site,
including production, protection, and conservation. (Reprinted by per-
mission from Springer Nature, from Brockerhoff, E.G., Jactel, H.,
Parrotta, J.A., Quine, C.P., Sayer, J., 2008. Plantation forests and biodi-
versity: oxymoron or opportunity? Biodiversity and Conservation 17,
925–951. # 2008 Springer Science+Business Media B.V.)
Fig. 7.29 The role of a plantation

forest in a corridor-patch-matrix
landscape model in a fragmented
land with ~85% loss of natural
forest and ~20% plantation forest.
Springer Nature, from
Brockerhoff, E.G., Jactel, H.,
Parrotta, J.A., Quine, C.P., Sayer,
J., 2008. Plantation forests and
biodiversity: oxymoron or
opportunity? Biodiversity and
# 2008 Springer Science
+Business Media B.V.)
large clear-cuts of lodgepole pine and treated the residue of

CWD in three ways. Using three replicates sites on three
different study areas, they (1) dispersed CWD uniformly
over the site as their control treatment (the conventional
way CWD is distributed after a logging operation); (2) placed
CWD in piles at an average of 2–3 piles per ha in another
treatment and (3) distributed CWD in windrows in a third
treatment. Over time, they determined the species richness
and abundance of forest-floor small mammals in all three
treatments and compared their treatment results to the rich-
ness and abundance of the same species in surrounding uncut
old-growth forests. There were site-specific and species-
specific differences, but, overall, Sullivan et al. found that
community species richness of forest floor mammals and
Fig. 7.30 The Iberian lynx (Lynx pardinus), the world’s most abundance of most species was higher on sites where CWD
endangered species of feline, has been shown to use plantation forests
as corridors to fragments of its preferred habitats in native forests. (Photo had been arranged in piles and windrows than those where it
courtesy of the Iberian Lynx Ex-situ Conservation Program, reprinted had been dispersed. Not only were species richness and
under CC BY 3.0) abundance greater in piles and windrows than on uncut
sites, they were also higher than those in uncut forests
(Fig. 7.31). The greater the volume of CWD, the more pro-
7.6.3 Mitigation in Forest Environments nounced the difference between the treatments, and the
greater the difference between control and uncut sites
Few forests today, even “natural” ones, are undisturbed by (Fig. 7.32) (Sullivan et al. 2012). The response of one spe-
commercial logging and other activities that remove wood cies, the southern red-backed vole (Myodes gapperi) was
and other products from forest ecosystems. The methods of particularly noteworthy. A specialist of closed canopy
logging, as well as the management of a site after logging, forests, the southern red-backed vole is normally absent
affect habitat quality and site biodiversity. In North America, from forest openings, including clear-cuts, and does not
many stands of lodgepole pine (Pinus contorta) have been return to cut sites until decades later (Ransome et al. 2009;
decimated by sudden, site-specific population increases Sullivan et al. 2011). Yet, when slash was arranged in rows or
(“outbreaks”) of the mountain pine beetle (Dendroctonus piles, the vole persisted in cut areas (Sullivan et al. 2012).
ponderosae). Where such outbreaks occur, lodgepole pines Sullivan et al. provide an example of how intentional
are damaged by the beetle and often die. Forest managers management of the effects of human disturbance can be
contract with commercial loggers to remove lodgepole from manipulated to enhance species diversity and abundance of
such areas through “salvage sales,” taking the timber quickly the community of forest floor mammals. Can human activity
and while it is still standing to maximize its market value. For also be manipulated to mitigate loss of habitat by species,
lodgepole pine, one of the most common methods of removal even large mammals with extensive home ranges?
is clearcutting, in which most or all of the trees on relatively
large areas are moved in a single cutting period. Unmarket-
able trees, as well as limbs and branches of marketable ones, 7.6.4 Mitigating Human Effects to Avoid
are normally left on the ground after cutting has been Habitat Loss and Range Displacement:
completed. Such residue, commonly called slash, is referred The Case of the Line Creek Elk
to by forest managers and forest ecologists as coarse woody
debris (CWD). CWD can provide physical structures that In a remote region of south central Montana (USA), a popu-
provide habitat to small forest mammals, but benefits are lation of elk (Cervus elaphus) known as the Line Creek herd
affected by how the CWD is distributed. winter in sagebrush-covered foothills embedded in a mosaic
In the China Valley of British Columbia, Canada, Thomas of private, state, and federal ownership. The herd’s name
Sullivan and his colleagues selected three study areas with comes from a stream on their winter range that follows the
Fig. 7.31 (Left) Mean numbers of forest floor mammals/ha in three arranged in piles, or arranged in windrows, compared to forest floor
areas, each with three clear cut sites (replicates) monitored over 3 years mammal densities in surrounding uncut forests. Means with different
(2007–2009) in British Columbia (Canada) where slash on logged sites letters are significantly different (Duncan’s multiple range test).
was dispersed (no treatment), arranged in piles, or arranged in (Reprinted by permission from Oxford University Press, from Sullivan,
windrows, compared to forest floor mammal densities in surrounding T.P., Sullivan, D.S., Lindgren, P.M.F., Ransome, D.B., 2012. If we
uncut forests. (Right) Mean species richness (number of species) of build habitat, will they come? Woody debris structures and conservation
forest floor mammals in three areas, each with three clear cut sites of forest mammals. Journal of Mammalogy 93, 1456–1468. # 2012
(replicates) monitored over 3 years (2007–2009) in British Columbia American Society of Mammalogists)
(Canada) where slash on logged sites was dispersed (no treatment),
boundary line between the US states of Montana and conservation organizations, and local residents voiced con-
Wyoming for several miles. Here the US Forest Service cern that drilling would displace the elk and degrade their
granted a lease to Phillips Petroleum Company (now habitat. There was also concern that the elk might move
ConocoPhillips) permitting them to engage in exploratory permanently south across the state line in to Wyoming,
drilling for oil. Other state and federal agencies, private depriving Montana of revenue from license and hunting fees.
the site was limited to a single road used only by vehicles of

the company and the Forest Service, then closed after explo-
ration was completed. Workers confined activity to the drill
site. After each drilling season, the drilling rig was lowered
from a vertical to a horizontal position to make it impossible
to see from adjacent drainages, reducing its visual impact.
When drilling activity ended, the drilling site was re-seeded
to native grasses.
The Line Creek elk had been monitored through radio
telemetry prior to drilling, and such monitoring continued
during and after drilling. Of interest to all, and surprising
to many, the population did not move from its original
range. The size, shape and position of the herd’s home
range remained unchanged before, during and after drilling.
Elk avoided the drill site itself and increased their use of
forested habitats (“hiding cover”) near the well but did not
change their overall patterns of range use, even when
measured at fine spatial scales (Van Dyke and Klein 1996)
(Fig. 7.33).
The Line Creek study illustrates techniques and principles
that can be used to mitigate, or even altogether avoid, habitat
loss associated with displacement of animal populations by
human activities. The first is that of timing limitations. If
animals use habitats on a seasonal basis, human activities can
be scheduled to take place in those habitats in seasons when
animals are absent. A second mitigation technique is that of
limited access. If roads are built for human use, their detri-
mental effects can be reduced if their access is limited to only
those performing essential activities. A third technique is that
of visual minimization, exemplified here by lowering the
drilling rig to a horizontal position. Reducing the distance
at which an object associated with disturbance can be seen by
Fig. 7.32 Regression relationships of mean (a) total abundance
(individuals/ha) (b) species richness, and (c) species diversity of forest
animals reduces the animals’ response to the object and
floor small mammals to mean volume (m3) of woody debris in dis- makes more of the area available for the animals’ use. A
persed, piled, and windrowed treatments at study sites in three areas fourth technique is that of reclamation. By immediately
logged by clearcutting in British Columbia (Canada), 2007–2009. Dif- reseeding the disturbed site, the oil company began succes-
ferent symbols represent different study sites. (Reprinted by permission
from Oxford University Press, from Sullivan, T.P., Sullivan, D.S.,
sional activity on the actual drilling site quickly, and acted in
Lindgren, P.M.F., Ransome, D.B., 2012. If we build habitat, will they a way that could more rapidly make the site resemble its
come? Woody debris structures and conservation of forest mammals. previously undisturbed state. A fifth technique, although not
Journal of Mammalogy 93, 1456–1468. # 2012 American Society of used in this example, is that of no surface occupancy.
Mammalogists)
Through technology like directional drilling, extraction of
oil, natural gas, and other subsurface resources located
Phillips adopted a comprehensive series of rules to mini- beneath a site covered by high quality habitat can be accom-
mize adverse effects of drilling. The company agreed to use plished by surface drilling on a remote site of low quality
the drilling site only in summer and early fall, when elk habitat. This practice can reduce detrimental effects of these
would be using other areas at higher elevations. Access to activities on sensitive populations.
Fig. 7.33 Seasonal ranges of an elk population before (solid line), place (see text) did not significantly alter the size or boundaries of their
during (dotted line), and after (dashed line) oil drilling activity on the range. (Reprinted by permission from Oxford University Press, from
population’s winter range. Elk avoided the drilling site and altered Van Dyke, F., Klein, W.C., 1996. Response of Elk to Installation of Oil
patterns of habitat use, but, with a variety of mitigation measures in Wells. Journal of Mammalogy 77, 1028–1041)
Synthesis biologists to determine ways to permit human use in

Habitat conservation is the foundation of population designated areas with minimal disturbance. In a world
conservation. Yet conservation biologists still struggle where human presence continues to grow, conservation
to understand the separate and interactive effects of biologists must employ mitigation strategies to main-
habitat loss, fragmentation, isolation, and degradation tain viable habitats and healthy landscapes even in the
on plant and animal populations. Conservationists midst of human presence.
attempt to communicate these effects coherently to
the public and translate their understanding into mean-
ingful policies that manage habitat and processes that
shape it. The importance of habitat is recognized in
concepts like that of “critical habitat,” written into the
US Endangered Species Act, yet ignored in many other
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The Conservation of Aquatic Systems
8
By the law of nature, these things are common to mankind – the air, running water, the sea, and consequently the
shores of the sea.
Justinian, 535 A. D.
Keywords degradation, and invasions by non-native species. In fresh-

Aquatic environments · Freshwater ecosystems · Eutro- water systems, alterations of natural flow regimes by land-use
phication · Flow regime · Hydrologic connectivity · changes, river impoundments, and water removal also have
Wetlands · Marine protected areas · Benthic profound impacts on their biotic communities.
environments · Coral reef restoration · Marine fisheries One reason freshwater ecosystems receive relatively little
attention, and even less protection, is due to lack of a reliable,
global, synthesized database providing trustworthy informa-
tion on ranges of freshwater species, especially fish.
Overview Although freshwater systems contain only 0.01% of the
In this chapter, you will learn about: world’s water and cover only 0.8% of the earth’s surface,
they have been estimated to contain 126,000 described spe-
1. Basic properties of aquatic habitats. cies including fishes, mollusks, reptiles, insects, plants, and
2. Types of freshwater and marine ecosystems. mammals, and projections of the number of undescribed
3. Conservation problems, goals, and management species put this number over one million (IUCN 2019), an
strategies associated with freshwater and marine estimate which could include nearly 6% of all species on the
ecosystems. planet, (Dudgeon et al. 2006), distributed in complex conti-
nental, regional, and local patterns, often with disproportion-
ately high levels of endemism (Fig. 8.1). Of these, 7,614 were
listed as critically endangered, endangered, vulnerable, or
near threatened on IUCN’s Red List (Chap. 2).
Like freshwater ecosystems, marine systems are dispro-
8.1 Conservation Challenges of Aquatic portionate contributors to global biodiversity. Of
Habitats 29 non-symbiont animal phyla known on earth, all but one
have representatives in the ocean, and all of these have
8.1.1 Reservoirs of Global Biodiversity representatives in benthic (bottom-dwelling) communities.
In fact, most of the diversity found in marine ecosystems
Freshwater and marine environments are among the world’s consists of invertebrates that live in or on bottom sediments
most diverse ecosystems. Even if considered only on the (Snelgrove 1999). We are only now beginning to appreciate
basis of human self-interest, they merit care and nurture the biodiversity of such communities. For example, 64% of
because they provide goods and services vital to human polychaete (tubeworm) taxa identified in one classic deep-sea
societies. Yet they receive little protection, being among the study were previously unknown to science (Grassle and
most altered ecosystems on earth, threatened by overexploi- Maciolek 1992). Given such ignorance, it is not surprising
tation, water pollution, habitat fragmentation, habitat

308 8 The Conservation of Aquatic Systems
a b
Number of freshwater fish species Number of endemic freshwater fish species

1–19 67–101 214–322 1–11 28–40 74–118 No endemics
20–41 102–151 323–490 12–19 41–55 119–195 No data
42–66 152–213 491–880 Estimate 20–27 56–73 196–387
c d
Percentage freshwater fish species endemism Freshwater fish species per ecoregion area
> 0–5% > 15–21% > 39–51% 0–1 > 3–4 > 6–7 No data
No data > 5–10% > 21–29% > 51–71% > 1–2 > 4–5 > 7–8
0% Endemism > 10–15% > 29–39% > 71–100% > 2–3 > 5–6 >8
Number of species per 104 square kilometers
Fig. 8.1 Recent freshwater species data of freshwater ecoregions Bussing, W., Stiassny, M.L.J., Skelton, P., Allen, G.R., Unmack, P.,
displaying patterns of (a) species richness, (b) number of endemic Naseka, A., Ng, R., Sindorf, N., Robertson, J., Armijo, E., Higgins, J.V.,
species, (c) percentage endemism, and (d) species per ecoregion. Note Heibel, T.J., Wikramanayake, E., Olson, D., López, H.L., Reis, R.E.,
the consistent pattern of non-uniform, region-specific variation of the Lundberg, J.G., Sabaj Pérez, M.H., Petry, P., 2008. Freshwater
distributions of all four variables. (Reprinted by permission from Oxford Ecoregions of the World: A New Map of Biogeographic Units for
University Press, from Abell, R., Thieme, M.L., Revenga, C., Bryer, M., Freshwater Biodiversity Conservation. BioScience 58, 403–414. #
Kottelat, M., Bogutskaya, N., Coad, B., Mandrak, N., Balderas, S.C., 2008 American Institute of Biological Sciences)
that we do not know the exact number of marine benthic absorb and store large quantities of heat with relatively little
species, but past estimates have ranged from 500,000 (May change in temperature. This makes aquatic environments
1992) to more than 100 million (Lambshead 1993). Like their more stable in temperature than terrestrial environments,
freshwater counterparts, they are disproportionately and their biotic communities less well adapted to rapid tem-
threatened, with 1,683 species listed as critically endangered, perature changes. Water also differs from air in being more
endangered, vulnerable, or near threatened by IUCN. Both viscous – more resistant to internal flow, and therefore more
freshwater and marine systems merit attention in conserva- resistant to movement. Aquatic organisms must expend more
tion, so each will receive detailed focus in this chapter. We energy in moving about, possess a physique which minimizes
will begin with the waters that most directly intersect and resistance, be highly adapted to specific environments that
interact with the terrestrial environment – freshwater minimize need for movement, or all or some combination of
ecosystems. these.
The greater density of water, compared to air, affects
transmission of light. As light passes through water, it
8.1.2 Basic Properties of Aquatic Environments changes in wavelength and weakens in intensity. Such
changes create a photic zone in which light penetrates to
All living systems are constrained and driven by the func- some maximum depth, determined in part by the relative
tional properties of the medium in which their life exists. The clarity of the water. If the water contains high concentrations
properties of water are vastly different than those of air. of suspended material, light will be absorbed more rapidly
Compared to air, water has a higher heat capacity. It can and depth of light penetration will be less. Since light is
8.1 Conservation Challenges of Aquatic Habitats 309
essential for photosynthesis, this means that an aquatic envi- with pools often leads to conditions that are less productive
ronment has limits, in depth, in which plants and other and where decomposition is the more dominant process.
photosynthetic life can grow, a condition which creates, in As a stream continues its course, it interacts with the
aquatic environments like lakes, a trophogenic zone of high landscape in ways that affect both. To the landscape, the
light intensity (more shallow water) dominated by the process stream is an agent of erosion as it cuts and shapes its channel,
of photosynthesis. Below this is a tropholytic zone of lower carrying away with it sediments and other material from the
light intensity in which the dominant process is land around it. To the stream, the landscape is a source of
decomposition. material and energy. At any given point in most streams, 90%
Oxygen is available in much lower concentrations than in of its inputs come from upstream or from the surrounding
air and always a limiting factor. Aquatic organisms must area. As other streams join it, the stream increases in magni-
compensate either by having respiratory mechanisms that tude or stream order. At its origin, a stream is referred to as a
can more efficiently remove oxygen from water, metabolic first order stream. When that stream joins another first order
processes that can function at low levels of oxygen, or by stream, the new stream becomes a second order stream.
periodically coming to the surface to take in oxygen from air. When two second order streams join, they form a third
Broadly, freshwater ecosystems can be categorized as order stream. Whenever two streams of the same order join,
lentic systems, like lakes and ponds, in which there is less they form a stream of the next highest order. Increasing
internal movement of water, and therefore where zonation stream order reflects an increasing area of watershed, the
and stratification are more pronounced, and lotic systems like area of landscape drained by the stream.
rivers and streams where water is constantly flowing. Such Many ecologists refer to streams and rivers as derived
ongoing movement of water in streams makes them more ecosystems because they draw the majority of matter and
resistant to physical stratifications. However, because energy from outside their own systems. It is precisely their
streams flow over the landscape, their characteristics change derivative ecological nature that makes them vulnerable to
over distance and space. Near its source or headwaters, activities on surrounding lands. Therein lies one of the most
streams typically flow straighter and faster. As they travel, important dangers to aquatic habitat. What kinds of activities
streams slow in velocity, meander from side to side in their on land create threats to the water in streams, lakes, and
courses, and increase in volume of water in their channel. rivers?
The more uniform flow near the headwaters begins to be
replaced by an alternating pattern of faster, shallower, turbu-
lent riffles and slower moving, deep water pools (Fig. 8.2). 8.1.3 Threats to Freshwater Ecosystems
With light penetrating throughout the riffles, photosynthetic
organisms like diatoms, green and blue green algae, and Today freshwater ecosystems are among the most
aquatic moss that can persist in the stream or on the substrate endangered on Earth because of the historically unprece-
are more abundant, making riffle areas important zones of dented use and withdrawal of their most valuable resource –
primary production. The deeper, slower water associated water. Globally, freshwater withdrawals have almost doubled
Fig. 8.2 Classic, normal

alternation of riffle (foreground)
and pool (background) habitat in a
river. Riffles provide shallow
water habitat of increased oxygen
levels and higher rates of
photosynthesis. Pools provide
deeper water habitat of reduced
current and increased prevalence
of decomposition. (Photo courtesy
of Joel Betts)
since 1960, with more than half of all accessible freshwater of threat to different freshwater species groups. Exploitation
runoff now being appropriated for human use. Today primarily affects freshwater vertebrates, especially fishes,
increased water demands for irrigation, industrial, and reptiles and some amphibians. The other four are more sys-
domestic uses threaten freshwater rivers throughout the temic, threatening all forms of freshwater life. Of these,
world. The Nile in Egypt, the Ganges in South Asia, the introductions of exotic species may be especially harmful in
Amu Dar’ya and Syr Dar’ya in Central Asia, the Yellow freshwater systems already experiencing high levels of
River in China, and the Colorado River in North America human-induced stress, precisely because such species can
are among the world’s major rivers that are dammed, more easily invade systems where native species have been
diverted, and overused to the point that little or none of reduced or eliminated (Dudgeon et al. 2006).
their freshwater reaches the sea. With such excessive use These stressors are increasing extinction rates in freshwa-
comes degradation, with the health of many of the world’s ter systems far more than those in even the most human-
freshwater ecosystems showing a decline of 50% between impacted terrestrial ecosystems. At a regional scale, the
1970 and 1995 (Loh et al. 1998). projected mean future extinction rate for North American
Combined with massive withdrawals, threats to freshwater freshwater fauna has been estimated to be about five times
ecosystems arise from (1) overexploitation of species by greater than for terrestrial fauna and three times that of coastal
humans; (2) habitat degradation associated with destructive marine mammals (Ricciardi and Rasmussen 1999) – a rate
land use change, particularly land use change generating comparable to the range of estimates predicted for tropical
high levels of erosion leading to increases in sediment or rainforest communities. Among these are approximately
chemical input; (3) water pollution, especially nutrient 790 native freshwater fish species threatened with extinction,
enrichment in otherwise nutrient-limited aquatic systems; 627 as a direct result of dams, pollution and introduced
(4) direct human-mediated modification of flow volumes species (Stendara et al. 2012). Among freshwater mussels,
and rates that lead to destruction or modification of the published conservative estimates of extinction rates range
natural habitats within the aquatic environment; and (5) intro- from 2–7% per century, one of the highest current extinction
duction or invasion of non-native species which may reduce rates of all taxonomic groups. Less conservative estimates
or eliminate native species (Fig. 8.3). These five categories predict future mussel extinction rates in continental North
and their interactions with one another create different kinds America to exceed 6% per decade (Geist 2011). More than
60% of the total nominal freshwater snail fauna are listed by
IUCN as Critically Imperiled or Presumed or Possibly
Extinct and more than 40% of freshwater snail species are
negatively affected by anthropogenic factors, including over-
exploitation, habitat degradation, and climate change (Lysne
et al. 2008).
Despite their current peril, freshwater species often
receive little or no consideration at the level of conservation
awareness, management, or planning. Most species in aquatic
communities are rare, and their natural histories little
investigated and poorly understood (Dudgeon et al. 2006).
For example, an exhaustive global inventory of freshwater
taxa admitted serious survey gaps and assigned species
distributions only to the level of continent (Lévêque et al.
2005), demonstrating how unprepared humans are to under-
take the conservation of aquatic organisms. Similarly, con-
servation concerns regarding freshwater species have long
been under-represented in scientific literature.
Such high estimates and risks of extinction, combined
with scientific ignorance and professional inattention to
Fig. 8.3 Major global threats to aquatic species and habitats and their aquatic species, expose the vulnerability of freshwater
established or potential pathways for interactions and subsequent detri- systems. Despite their combination of richness, endemism,
ment to freshwater biota. (Reprinted by permission from Wiley, from vulnerability and threat, few comprehensive conservation
Dudgeon, D., Arthington, A.H., Gessner, M.O., Kawabata, Z.-I.,
Knowler, D.J., Lévêque, C., Naiman, R.J., Prieur-Richard, A.-H.,
planning efforts have targeted freshwater systems and their
Soto, D., Stiassny, M.L.J., Sullivan, C.A., 2006. Freshwater biodiver- dependent organisms. But to develop appropriate responses
sity: importance, threats, status and conservation challenges. Biological to the conservation of freshwater systems, we must have a
Reviews 81, 163. # 2005 Cambridge Philosophical Society) more detailed understanding of what threatens them.
8.1.4 Chemical and Biological Degradation pH is a change in chemical reactions occurring in the aquatic
system, especially in metallic ions such as aluminum, lead, or
Chemical alteration or pollution can take many forms in cadmium. Aluminum is deadly to fish because, under acidic
lakes and streams, but two of the most common are eutro- conditions, it binds to their gills and impedes respiration
phication and acidification. Eutrophication occurs when (Brönmark and Hansson 1998). When fish populations are
nutrients are released into aquatic systems from upstream or reduced in acidified lakes, many invertebrates are released
surrounding agricultural areas (in the form of fertilizer run- from predation pressure and their populations then increase
off) or from towns and cities (in the form of human waste) (Brönmark and Hansson 1998). In addition, once aluminum
(Brönmark and Hansson 1998). Higher levels of nitrogen and begins to precipitate out of solution, it binds with phosphorus
phosphorus, for example, trigger increases in primary and precipitates as aluminum phosphate. When this occurs,
producers (formerly limited by a scarcity of these nutrients). phosphorus becomes unavailable as a nutrient for organisms.
Periphytic (attached) algae and submersed macrophytes Introductions of exotic species pose a greater threat in
increase at the beginning of the process, but then decline as aquatic habitats than terrestrial ones. In Canada for example,
phytoplankton and cyanobacteria (“blue-green algae”) invasive species are a primary threat to 26 of 41 threatened
increase in abundance and reduce the amount of light that listed fish species, and 6 of 11 threatened mollusk species
penetrates the water. Dead organisms accumulate as sedi- (Dextrase and Mandrak 2006). The true number of invasive
ment, and bacteria that remove minerals from decaying species, especially among invertebrates, is probably
organic matter consume large amounts of oxygen they extract underestimated because of poor or irregular monitoring,
from the water. Loss of some species (“fish kills”) follows as inability to distinguish invasive from native species, and
oxygen is depleted, but cyprinid fishes (Family Cyprinidae, lack of historical knowledge and data concerning native
carps and minnows) increase because they can survive in species (Ricciardi 2015). Such species are being discovered
poorly oxygenated waters and are efficient predators of zoo- at increasing rates in freshwater systems worldwide
plankton, organisms that increase during eutrophication. As a (Fig. 8.4), globally dispersed by shipping where they can be
result, populations of grazing zooplankton decrease. Levels concealed in ballast water and sediments, as well as by active
of phytoplankton, the prey of zooplankton, then grow, further and intentional live trade (Ricciardi 2015). Invertebrate
increasing turbidity (Brönmark and Hansson 1998). As eutro- invaders most damaging to their new ecosystems tend to be
phication progresses, the system declines in value as a source species that use critical resources differently and more effi-
of drinking water, recreation, and food. ciently than native species (creating shortages of food), may
Acidification is a process through which the pH of surface be predators or grazers that are not regulated by other species,
freshwaters declines (becomes more acidic) because of inputs and may have higher rates of reproduction than native species
of acidic precipitation in the form of rain, snow, or fog. (Ricciardi 2015).
Emissions of hydrogen sulfide (H2S), produced by burning Disturbed aquatic systems, and particularly those
coal to generate electricity, and nitrous oxide (NO), an exhaust associated with urban areas or other regions of high human
waste from cars, can combine with atmospheric water vapor to population densities and use, are more vulnerable to
form sulfuric and nitric acid that fall as precipitation into a invasions than undisturbed systems, partly because they
stream or lake or their drainage area. Acidic inputs generally lack the resiliency of undisturbed systems and partly because
do not affect pH in areas where soil and rock substrates contain their exposure to increased levels of human use lead to
significant amounts of calcium carbonate (CaCO3) or other greater and more frequent introductions of non-indigenous
carbonate compounds. These chemicals react with water to species, accidentally or on purpose, by humans (Dudgeon
form carbonate and bicarbonate ions that buffer a system et al. 2006). Many invasive aquatic plant species, particularly
against acidic inputs. In areas without these buffering in wetlands, tend to be large or spreading invasive perennials,
capacities, such as those with granitic substrates or granitic- such as purple loosestrife (Lythrum salicaria), reed canary
derived soils, the same inputs can have disastrous effects on grass (Phalaris arundinacea), European frog-bit
aquatic communities, as has happened in lakes and streams in (Hydrocharis morsus-ranae), and others, which can, under
the Adirondack Mountains in the northeastern United States. the right (or should we say “wrong”?) conditions, become
When an aquatic system becomes acidified, problems dominant in freshwater lakes, streams, and wetlands.
begin with a lowering of pH due to acidic inputs, especially Pollution and invasive species can pose threats to aquatic
during periods of heavy rain or during spring snowmelt. The systems that can act independently, but more often the effects
most common and immediate effect of lower pH is a reduc- of pollution and invasive species are created by changes to
tion or cessation of reproductive effort in many species of the habitat of the freshwater system itself. One of the most
fish, amphibians, and aquatic invertebrates. Some species important and widespread causes of such changes in aquatic
suffer direct mortality. In others, sperm or eggs may become habitat occur when humans make changes to flow regimes of
inviable. An indirect but more devastating effect of the lower freshwater systems through the construction of dams.
Fig. 8.4 Accumulation of alien invertebrate species recorded as increasing rate of accumulation in the last 40–80 years. (Reprinted by
established over time in four major freshwater ecosystems worldwide: permission from Elsevier, from Ricciardi, A., 2015. Ecology of Invasive
(a) the Laurentian (US – Canada) Great Lakes, (b) the Rhine River in Alien Invertebrates, in: Thorp and Covich’s Freshwater Invertebrates
Europe, (c) the Thames River in the UK, and (d) the Ebro River on the (Fourth Edition). Elsevier, pp. 83–91. # 2015 Elsevier Inc)
Iberian Peninsula (Spain and Portugal). Note, in every system, the
8.1.5 Dams, Levees, and Flood Plains: Flow, California (USA), for example, rivers whose flow regimes
Impoundments, and Connectivity have been altered by dams have more predator-resistant
invertebrates and fewer fish than undammed rivers (Wotten
Because streamflow is the “master variable” limiting distri- et al. 1996). This change occurs because more uniform flows
bution of riverine species and regulating ecological integrity created by dams allow non-native species to replace locally
of flowing water systems, human-caused flow modification adapted native fauna. As barriers, dams affect movements of
or alteration of flow regimes is, as we have already noted animals and plants, altering metapopulation dynamics within
(Fig. 8.3) one of the most dangerous threats to freshwater the aquatic ecosystem (Saunders et al. 2002).
species worldwide. Such alteration most commonly occurs as Dams restrict naturally flowing rivers and streams to cre-
a result of building dams to create artificial impoundments of ate stable and readily available supplies of water for drinking,
water. The hydrological changes caused by dams and other irrigation, and industrial processes. Dams provide hydroelec-
human-created impoundments and diversions create systemic tric power, recreation; and flood control. But any dam, no
changes in aquatic systems. Among the most pervasive are matter what size, declines in usefulness overtime as
changes that occur in species composition. In northern accumulating sediment degrades the quality of water in the
reservoir, reduces the amount of energy that can be (3) flow events that permit or facilitate longitudinal dispersal,
generated, limits recreation, and eventually loses effective- such as flood plain development, affecting connectivity of
ness in flood control. For non-human species, effects of dams habitats with otherwise independent river systems, and
can be more negative and highly species-specific. In (4) flow events that follow long-term natural patterns of
New Zealand, construction of dams changed flow patterns flow variability in a particular river system and favor native
on rivers used by the endangered black stilt (Himantopus species adapted to such variability and discourage establish-
novaezealandiae) which nests on gravel bars in those rivers. ment of non-native species (Dudgeon 2006).
Dams changed flow patterns that raised water levels upstream
from the dams, flooding the stilt’s nests and reducing recruit-
ment to near zero in affected areas (Boyce and Payne 1997). 8.1.6 Consequences of Dams on Fish
In the United States, dams have had similar effects on reduc- Biodiversity and Community
ing nesting sites for two endangered species, the least tern Composition
(Sternula antillarum) and piping plover (Charadrius
melodus), both of which nest on sandbars created by seasonal One of the most thorough studies of the effects of dams on
variations in the river’s flow rate. fish populations was conducted by fisheries scientist Angelo
One of the most straight-forward and increasingly Agostinho and his colleagues in Brazil. Examining the effects
employed means of managing flow in freshwater ecosystems of hundreds of dams in multiple watersheds, they found that
is through Environmental Water Allocations, or EWAs, that “Soon after the reservoir is filled, patterns of thermal/chemi-
mimic natural variability and provide for a range of flows cal stratification intensify progressively in the water column,
over annual or seasonal cycles (Dudgeon et al. 2006). Well- along with the decay of the flooded organic matter (vegeta-
designed EWA’s mimic a river’s historic flows, and so pro- tion, litter, and soil). Consequently, environmental quality
vide habitat and sufficient water for all native species may deteriorate in some layers of the column (e.g., thermal
associated with the river that are adapted to such variable stress, low dissolved oxygen, acidification), especially close
flow rates and their effects on stream and riparian habitat, to the bottom. . .” (Agostinho et al. 2008:1122). These
physical and chemical characteristics of water, and volume of changes in abiotic variables affected fundamental ecological
water in the river channel. dynamics of the entire system. Non-migratory (sedentary)
Four factors associated with flow that vary naturally or species were able to colonize inner zones of reservoirs
would be managed by an EWA are primary drivers of species because they had simpler demands for completing their life-
habitat and biodiversity in a river system (Fig. 8.5) These are cycles. In contrast, migratory species of fish declined due to a
(1) large (and often relatively rare) flow events that shape shortage of lentic (flowing water) environments, spatial frag-
overall river channel form, habitat complexity, and degree of mentation imposed by the dam, and loss of critical habitats
habitat patch disturbance; (2) timing and seasonality of flow needed to complete their own, usually more complex life
events and variation, which affect aquatic biodiversity by cycles (Agostinho et al. 2008).
favoring or discriminating against species with particular Because of the impoundments, the amount and quality of
life history patterns, such as seasonality of spawning or original habitat in the river was modified and degraded,
seasonal recruitment of new individuals to the population; changing the distribution and extent of lacustrine (deep
Fig. 8.5 Primary drivers Principle 3

(“principles”) affecting a natural lateral connectivity Principle 1
or managed flow regime in a river longitudinal connectivity channel form
showing how the river’s aquatic
habitat complexity biotic diversity
biodiversity is affected via inter-
patch disturbance
related mechanisms (Principles access to spates
1–4) operating at different scales floodplains
of time (X axis) and volume of
water discharged during flow variability Principle 2
events (Y axis). “Spates” refer to dispersal Life history patterns
sudden flood events in the river. triggers spawning
Discharge
(Reprinted by permission from reproductive triggers recruitment

Springer Nature, from Bunn, S.E.,
seasonality
Arthington, A.H., 2002. Basic
predictability
Principles and Ecological stable baseflows
drought
Consequences of Altered Flow
Regimes for Aquatic Biodiversity.
Environmental Management 30, Time
492–507. # 2002 Springer-
Verlag New York Inc) Principle 4
natural regime discourages invasions
lake), transition, and riverine zones relative to the original original river channel. Open and deep water areas were
river (Fig. 8.6) and creating an “environmental filter” that characterized by low levels of productivity, light, and oxy-
altered richness and abundance of fish communities in the gen. As a result, Angostinho et al. noted, “few species colo-
reservoir over time (Fig. 8.7). The formation of the reservoir nize the vast pelagic areas in reservoirs. The deep layers
created extensive pelagic (open water) and deep water areas exhibit similar qualities, particularly in the lacustrine zone.
at the expense of other types of habitat more common in the Few species are adapted to persist in such an environment,
Fig. 8.6 Longitudinal zoning of the Itaipu Reservoir in the upper A.A., Pelicice, F.M., Gomes, L.C., 2008. Dams and the fish fauna of the
Paraná River basin, Brazil, showing how predominant ecological pro- Neotropical region: impacts and management related to diversity and
cesses changed and differed in each zone once the reservoir was fisheries. Brazilian Journal of Biology 68, 1119–1132)
established. (Reprinted under CC BY 4.0 International, from Agostinho,
Fig. 8.8 The pacu (Piaractus mesopotamicus), one of many migratory

species in South American rivers whose numbers decline after construc-
tion of reservoir-creating dams. (Photo courtesy of Leonard L. Lovshin)
Fig. 8.7 Changes in fish species richness and abundance in each of the
riverine (Riv), transition (Tra) and lacustrine (Lac) zones in the Itaipu
Reservoir, Brazil, after formation the reservoir, between 1987 and 1997.
(Reprinted under CC BY 4.0 International, from Agostinho, A.A.,
Pelicice, F.M., Gomes, L.C., 2008. Dams and the fish fauna of the
Neotropical region: impacts and management related to diversity and
fisheries. Brazilian Journal of Biology 68, 1119–1132)
which eventually becomes inhospitable. . .” (Agostinho et al.

2008:1124).
Analyzing fish species richness in 75 Brazilian reservoirs,
Agostinho et al. found that even those that covered large
areas averaged only 30 species of fish, a low number relative
to the high diversity found in most unaltered South American
river systems. As the reservoir aged, species numbers
declined further, falling to an average of 20 species per
reservoir in reservoirs more than 20 years old. Because the
dams blocked upstream movement of fish, the loss of migra-
tory species in reservoirs was the greatest contributor to this
decline in species richness, which included many of the
larger fish species (Figs. 8.8 and 8.9). Loss of these migratory
species had detrimental economic effects. Agostinho et al.
noted that “. . .the migratory guild, which includes large
species such as the dourado Salminus brasiliensis. . ., the
pintado Pseudoplatystoma corruscans. . . the pacu Piaractus
mesopotamicus. . . and the giant jaú Zungaro jahu. . ., is the
first group to decline or disappear from impounded areas”
(Agostinho et al. 2008:1124) (Fig. 8.9). Only 5% of the Fig. 8.9 Fish caught in the Itaipu River before (1977) and after (1987
reservoirs studied had more than three migratory species and 1997) construction of the Itaipu Reservoir. Migratory species in
among the most abundant fish, and more than 50% lacked bold. Asterisk indicates species that normally grow longer than 60 cm.
(Reprinted under CC BY 4.0 International, from Agostinho, A.A.,
any migratory species (Agostinho et al. 2008). Pelicice, F.M., Gomes, L.C., 2008. Dams and the fish fauna of the
Managers tried to remedy loss of species and populations Neotropical region: impacts and management related to diversity and
by stocking fish raised in hatcheries, requiring great expense fisheries. Brazilian Journal of Biology 68, 1119–1132)
but yielding little result. “According to data on yield from Similarly, structures like levees (raised embankments along a
artisanal fisheries and stocking effort (number of fry released) river channel) that protect against flooding by forcing rising
in six reservoirs operated by the Electricity Company of São water to remain within the channel can have the same nega-
Paulo State,” noted Agostinho et al., “. . .there is no relation- tive effect on a floodplain and make flood events worse
ship between these variables [i.e., yield and stocking effort]. downstream.
In spite of these results, stocking is still the main management Flow rate and volume are controlling variables that most
strategy used by hydropower companies and environmental affect a river’s ecological dynamics. These are the things
agencies as well as the main strategy requested by dams most impede. In watersheds where dams have not
stakeholders” (Agostinho et al. 2008:1128). been constructed or where they can be removed, a natural
flow approach should be applied, especially to aquatic
systems and habitats in protected areas where historical
8.1.7 Dams as Barriers to Population hydrological regimes are relatively intact or could be
Persistence and Reproduction restored. Because of increasing demand for freshwater
supplies and services, maintaining such natural regimes will
The work of Agostinho and his colleagues demonstrated the require active management. But even where dams or other
effects of dams on the dynamics of river systems and impediments exist, reservoir releases, interbasin transfers,
accompanying changes in resident fish species. But dams groundwater withdrawals and periodic drawdowns of
have even more devastating impacts on populations of diad- reservoirs can be used to maintain flows and mimic natural
romous fish species which move from fresh water to salt discharge patterns, whereas weirs, embankments, and sluices
water in their normal life cycle. can be used to maintain water levels.
Limburg and Waldman (2009) conducted a comprehen- In the past, dam removal has been viewed is a radical
sive review of studies examining changes in the status of approach to aquatic conservation and restoration, but today is
diadromous fish populations in recent years in the North increasingly employed, with success, throughout the world
Atlantic Basin, a region of pronounced declines in fish (Table 8.1) (Saunders et al. 2002). In the United States, more
populations. Examining 35 separate long-term studies of than 1100 dams have been removed since the 1970s (Foley
diadromous fish populations, they found, in 13 studies, that et al. 2017). An increasing number of studies on the effects of
relative abundance in studied species dropped 98% compared dam removal are demonstrating that it is a workable strategy
to historic levels and to 90% in 11 other studies. Every to improve watershed connectivity, reduce environmental
species examined had suffered losses of some local hazards associated with aging dams, and promote recoloniza-
populations, and many species were now classified as tion by fish and other organisms previously blocked from
threatened or endangered. Factors contributing to declines moving upstream by the dam (Grant 2017). For example,
included overfishing, pollution, climate change, nonnative when the Edwards Dam on Maine’s Kennebec River (USA)
species, and aquaculture. But the greatest detriment to these was removed in 1999, benefits to diadromous fishes were
migratory species was habitat loss caused by damming the immediately apparent as these species rapidly reoccupied
rivers of their historic migration routes. As of 2009, there their historical spawning grounds (Limburg and Waldman
were more than 80,000 dams 6 feet or higher in the United 2009). After the Fort Halifax Dam, also on the Kennebec,
States, and even dams this small block thousands of miles of was breached in July 2008, over one million river herring
stream habitat for diadromous species. The American shad (Alosa spp.) moved upstream just 11 months later in June
(Alosa sapidissima), for example, has lost approximately 2009. As a result, managed river herring commercial harvests
4000 of an original 11,200 km of spawning habitat to dams were reinstituted for the first time in decades (Natural
(Limburg and Waldman 2009). Resources Council of Maine 2018).
Dams are but one threat to the hydrologic connectivity of
riverine aquatic ecosystems, defined as “water-mediated
transport of matter, or organisms within or between elements
Think of a dam on a river near your home. What effects
of the hydrologic cycle” (Jackson and Pringle 2010:37). As a
would removal of this dam have on the: (1) local econ-
means to control downstream flooding through regulation of
omy; (2) flow regime of the river; and (3) movement of
flow, dams are usually successful, but the price of success is
fish and other aquatic organisms? Considering these
often the loss of intact floodplains whose species and sys-
factors, would it be a good idea to remove this dam?
temic productivity depend on seasonal pulses of flooding.
8.2 Management of Freshwater Habitats for Conservation 317
Table 8.1 Examples of successful application of flow-regime management for conservation

Location Flow component Ecological success References
St. Mary River, Alberta, Increased minimum flows and maintained more Cottonwood recruitment Rood et al.
Canada gradual flow reductions after high-flow periods (1995)
Pecos River, New Mexico, Maintained minimum flow and mimicked natural Increased reproductive success of Pecos Poff et al.
US flow variation by regulating withdrawals for bluntnose shiner (1997)
irrigation
Kissimmee River wetlands Increased water flow using notched weirs and Recolonized native aquatic plants and Toth (1991)
and floodplain, Florida, diverted water over former floodplains insects, increased fish and waterfowl
US biomass
Winous Point Marsh, Simulated natural drought conditions with periodic Reestablished semiaquatic vegetation Meeks (1969)
Ohio, US drawdowns
Roanoke River, Virginia, Dampened unnatural flow fluctuations Increased juvenile abundance of striped Rulifson and
US bass Manooch
(1993)
Glasson Moss and Dammed drainage ditches Regenerated Sphagnum sp. Mawby
Wedholme flow, Cumbria, (1995)
England
Headwater streams, Decreased current velocities Increased moss cover and invertebrate Laasonen
northeastern Finland abundance et al. (1998)
Panshet dam, Pune, India Impounded water below dam into ditches Colonized fish, frogs, and aquatic Middleton
vegetation (1999)
Groot River, South Africa Released water at irregular intervals from Beervlei Induced spawning in smallscale redfin Cambray
dam minnow (1991)
Reprinted by permission from Wiley, from Saunders, D.L., Meeuwig, J.J., Vincent, A.C.J., 2002. Freshwater Protected Areas: Strategies for
Conservation. Conservation Biology 16, 30–41. # 2002 Society for Conservation Biology
over entire regions and drainage basins. They must use

8.2 Management of Freshwater Habitats techniques that enable them to stop such inputs from entering
for Conservation the system as they reach it, or they must remove or neutralize
such inputs as they enter.
8.2.1 Managing Chemical and Physical Inputs The most direct methods to stop such inputs are to
to Aquatic Systems (1) install filters and other devices at the proximate source
of input, such as the inflow stream, that remove the sediment
Because problems like sedimentation, eutrophication and and fertilizer when it arrives and (2) surround shorelines and
acidification are input-driven, their solution lies in input banks with vegetation that can provide high levels of nutrient
regulation. Sources of sedimentation and eutrophication are uptake from runoff. However, installation of filters and other
soil and fertilizer inflow, respectively, from surrounding devices is expensive, and the planting and management of
areas, especially agricultural lands, and urban waste. Both appropriate vegetation can also be costly, as well as labor-
are usually non-point source pollution problems, aggravated intensive.
through high levels of erosion from adjoining farmlands and Such practices might still be the best approach if they can
the abundance of impervious (high runoff) surfaces in urban lower levels of sediment and fertilizer entering aquatic
environments. The best management to address both habitats, but reductions of fertilizer input will not necessarily
problems would be socio-political in nature, specifically restore damage done by previous nutrient loading. What does
through laws and policies that (1) reduce the use of fertilizers, one do with the phosphorus and other nutrients that have
particularly on highly erodible lands and on lands near already entered and now remain in the system? Remedies for
watersheds, (2) require removal of fertilizers, especially this problem are dredging, chemical manipulation, and bio-
phosphorus, nitrate, and nitrite from urban sources before manipulation.
allowing urban discharge to proceed downstream, (3) reduce Dredging directly removes sediment from a lake, pond, or
erosion on agricultural lands through increased vegetative wetland, usually by physically scraping it off the bottom with
cover bordering streams and (4) encourage cultivation large, earth-moving machines. The sediment can then be
methods less destructive of soil structure. Social and political placed in a different, artificially constructed basin where the
remedies will receive the attention they deserve in later phosphorus is removed by physical or chemical means. Once
chapters (Chaps. 11 and 12). But managers of local aquatic purified, the sediment may then be returned to the system.
systems lack authority to implement such sweeping changes Dredging can be effective, but it is also expensive, labor
intensive, and disruptive to existing populations and

communities, especially benthic organisms. Dredging may
even require temporarily draining the system, and the method
is seldom suitable or effective in large, deep lakes. During the
dredging operation aquatic habitat may not be suitable for
other uses by humans or as habitat for other species.
Some chemical manipulation methods can be effective in
removing phosphorus by converting it into other chemical
states or that prevent it from entering or interacting in the
system. One of these is the so-called Riplox method
(Brönmark and Hansson 1998) in which the sediment surface
is first oxidized, causing the phosphorus in it to precipitate as
metal complexes. Then calcium nitrate (Ca(NO3)2) and iron
chloride (FeCl3) are added, increasing levels of oxygen and
iron. The pH of the system, which would tend to decline at
this point, is stabilized through the addition of calcium
hydroxide (Ca(OH)2). At a suitable pH, denitrifying bacteria
in the sediment will convert nitrate in the added calcium
nitrate to nitrogen gas (N2), releasing it to the atmosphere.
If these reactions proceed, the iron chloride then acts as a
chemical “lid” ion the sediment that prevents release of
phosphorus from the sediment into the water.
Bio-manipulation attacks the eutrophication problem by
altering the biotic community. First, densities of zooplank-
tivorous fish (usually the cyprinids) are reduced, either by Fig. 8.10 The “marble in a cup” model of alternative stable states of
adding piscivorous (fish-eating) species or by extracting the lakes relative to different levels of phosphorus inputs. Stability of the
cyprinids directly by trawling with gill nets. If the number of system is achieved through a combination of biomanipulation and
control of phosphorus inputs, but not one or the other exclusively.
zooplanktivorous fish decline, zooplankton populations
Typically, turbidity increases as nutrient levels increase. At low nutrient
should increase and the grazing rate on algae and phytoplank- level increases, turbidity may not change unless a disturbance
ton will rise. As a result, algal blooms decrease and water (represented by arrows) occurs. However, at a certain point () nutrient
clarity improves. Biomanipulation has worked best where at levels are too high for the water to remain in a clear state. (Reprinted by
permission from Scheffer, M., 1990. Multiplicity of stable states in
least 80% of the zooplanktivorous fish are removed, but its
freshwater systems. Hydrobiologia 200/201: 475–286 # 1990 Kluwer
success is not always due to reasons originally believed. Academic Publishers)
Rather, removal of fish often leads to an increase in
submerged macrophytic plants and periphytic algae at the
sediment surface. These absorb large amounts of nutrients gradual but rapid. The theory can be illustrated visually by
that are then no longer available for phytoplankton, which the “marble in a cup model” (Scheffer 1990) (Fig. 8.10). At
can now create conditions that oxidize the surface of the high levels of nutrient enrichment, the lake can exist only in a
sediment, reducing release of phosphorus. Removal of turbid state. As phosphate levels decline, alternate stable
benthic-feeding fish also contributes to a reduction in bottom states are possible, depending on which way the system
disturbance, lowers excretion of nutrients by fish, and (marble) is pushed. If, for example, macrophytes and peri-
reduces phosphorus released from the bodies of these fish phytic algae can be well established at intermediate nutrient
when they die and decompose. levels, they can take in excess amounts of phosphorus (“lux-
ury uptake”) that limits availability of phosphorus for phyto-
plankton and prevents their populations from increasing (and
8.2.2 Lake Systems as Alternative Stable States prevents the clarity of the water from decreasing). The lower
the level of nutrients in the water, the more stable the clear
Lake systems can, with respect to phosphorus, exist in state becomes. In cases like these, the system’s present con-
so-called alternative stable states, in which, at similar nutri- dition is a function not only of nutrient inputs and fish
ent levels, they may be dominated by submerged populations, but of populations of macrophytes and peri-
macrophytes in clear water or by high densities of phyto- phytic algae. And the system’s future condition is dependent
plankton and associated turbid water (Genkai-Kato and Car- on its present state, especially on how well established
penter 2005). The transition from one state to the other is not populations of macrophytes and algae are and how much
8.3 Managing Freshwater Systems at Landscape Levels 319
phosphorus they can absorb. This model is theoretical but has dilution of phosphorus concentrations in their deepest layers
empirical support from studies of Swedish lakes that exhibit (hypolimnion). The model predicted that lakes most vulnera-
alternative stable states (Blindow et al. 1997). ble to eutrophication, and least restorable after eutrophica-
Motomi Genkai-Kato and Stephen Carpenter of the Uni- tion, would be of intermediate depth - too deep to receive
versity of Wisconsin’s (USA) Center for Limnology took the benefits of phosphorus uptake by macrophytes, too shallow
concept of alternative stable states a step further through the to sufficiently dilute phosphorus in deeper waters. Regardless
development of a mathematical model designed to predict of depth, eutrophication was more likely to occur, and harder
lake transformations and the relative possibilities of restoring to reverse, at warmer temperatures (Fig. 8.11b). In fact, the
eutrophic lakes if a transformation occurred. Using data from model predicted that, in warmer lakes, shifts to alternate
their own and others’ studies, Genkai-Kato and Carpenter states (for example, clear to turbid) would occur at lower
found that both biotic and abiotic variables, and their levels of phosphorus input (Genkai-Kato and Carpenter
interactions, strongly affected not only the stable state of a 2005). These findings are disturbing in light of the global
lake, but also its potential for restoration. Shallow lakes could climate warming the Earth is now experiencing (Chap. 4).
be protected from eutrophication by the presence of
macrophytes that removed phosphorus from the water
(Fig. 8.11a). Deep lakes could be protected through the 8.3 Managing Freshwater Systems
at Landscape Levels
8.3.1 Protected Areas for Freshwater Systems
The concept of “protected areas,” in which freshwater

systems containing high habitat quality can be bounded and
preserved, is a technique commonly used in the conservation
of terrestrial ecosystems. This strategy of “fortress conserva-
tion” does not usually work well with freshwater systems,
unless the boundary for the protected area includes the entire
catchment or drainage basin of the stream intended for pro-
tection. Most protected areas have been designed to protect
terrestrial systems, so freshwater habitats often have been
protected only incidentally when they are included as part
of their inclusion in terrestrial reserves. Such inclusion, how-
ever, does not guarantee their protection, as managers of
many terrestrial parks have harmed the aquatic ecosystems
they contained through the stocking non-native sport fishes
(Saunders et al. 2002).
Today aquatic habitat conservationists are increasingly
employing a strategy in the design and management of fresh-
water protected areas known as whole catchment manage-
ment or WCM. WCM assumes the validity of the “landscape
filter” model (Poff 1997, Stendara et al. 2012), a theory of
aquatic community structure that sees freshwater aquatic
ecosystems, particularly rivers and streams, as products of
terrestrial environmental constraints or “filters” which control
Fig. 8.11 (a) The presence of aquatic macrophytes, rooted in the inputs of abiotic and biotic inputs to the stream ecosystem.
littoral (shoreline) zone of a lake, can reduce concentrations of chloro- The landscape filter concept can be used to accurately predict
phyll, an index of eutrophication, in water depths of 2–10 m. At greater characteristics of aquatic biological communities (Stendara
depths, there is insufficient light to establish substantial populations of
macrophytes, with consequent minimal effect on chlorophyll et al. 2012). Given such success, as well as support from
concentrations in the water. (b) As water temperature warms, chloro- ongoing theoretical and empirical research, most comprehen-
phyll concentrations increase rapidly, making eutrophication in the lake sive strategies for conservation of stream and river systems
more likely to occur and less likely to be reversible by management today are landscape-level approaches that focus on
actions. (Reprinted by permission from Wiley, from Genkai-Kato, M.,
Carpenter, S.R., 2005. Eutrophication due to phosphorus recycling in controlling characteristics and uses of terrestrial landscape
relation to lake morphometry, temperature, and macrophytes. Ecology and habitat within the watershed of the targeted stream or
86, 210–219. # 2005 Ecological Society of America) river.
WCM is arguably the most effective approach to riverine

aquatic conservation, but often the most difficult to apply. In Information Box 8.1: A Four-tiered Approach
any catchment, land adjacent to streams and rivers is usually to Aquatic Conservation
highly desirable for commercial, industrial, and private land In the Nature Conservancy’s (TNC) hierarchy of
owners. The first because it offers a convenient and cheap aquatic conservation categories (Information Box
opportunity for transportation of goods; the second because it Fig. 8.1), the highest level in the classification is called
can provide a convenient source of water for industrial pro- the aquatic zoogeographic unit (AZU), which serves
cesses; and the third because of its aesthetic attributes and as the overall planning element in initial conservation
opportunities for recreation. These interests may be limited to assessment. AZUs conform to major freshwater drain-
lands immediately adjacent to the stream, but because the age boundaries, generally 10,000–100,000 km2, and
catchment will include all the land in which surface runoff are distinguished by differences in continental and
drains into the stream, the stream will be affected by all land regional zoogeography that result from differences in
uses on all areas from which runoff is received. WCM, then, initial zoogeographic sources, patterns of drainage
must be able to regulate land use activities within the entire connections, and biotic changes over time in response
watershed if its conservation effort is to be effective. Gaining to climatic and geologic events. AZU planning units
such comprehensive authority over multiple corporate and delineate the area to be classified for a particular proj-
private landowners is a daunting, if not impossible, task in ect, and are differentiated on the basis of large-scale
larger watersheds (and even in smaller ones!), although lim- ecological differences rather than geopolitical
ited forms of WCM are possible with government authoriza- boundaries.
tion and local support. The second level of analysis relies on identification
When it is not possible to protect an entire catchment, one of ecological drainage units (EDUs) within an AZU.
alternative is to apply the Multiple-Use Module (MUM) EDUs represent regional biodiversity distinctions
approach. A MUM consists of a central, well-protected core within AZUs and are generally 1000–10,000 km2 in
surrounded by a series of buffer zones in which varying size. EDUs are delineated and classified by identifying
amounts of human activity are permitted. Only low-impact areas with similar biotic patterns and represent a finer
activities are permitted near the core, and potentially detri- scale of physiographic and zoogeographic diversity,
mental practices are excluded or relegated to the most distant allowing the selection of rivers and lakes for conserva-
zone. This design can be applied to freshwater protected areas tion to be stratified by environmental and biological
if the central core consists of a target waterbody (most appro- differences within an AZU.
priately, a river or stream) and its associated riparian zone. In The third level of classification is that of aquatic
this case, the MUM approach can be relatively easy to inte- ecological systems (AES) residing within a particular
grate with community-based conservation initiatives that EDU. AESs are stream networks representing a range
may already be in operation because some human activities of areas with distinct geomorphological patterns tied
would continue to be permitted within some zones of protec- together by similar environmental processes such as
tion (Saunders et al. 2002). hydrologic, nutrient, and temperature regimes. Patterns
of environmental conditions that determine the
characteristics of freshwater ecosystems and influence
8.3.2 Coarse-Filter Approaches for Regional biotic patterns are used to classify the AESs. Freshwa-
Representation ter ecosystem attributes such as water-body size,
hydrologic and temperature regime, chemistry, drain-
The World Database on Protected Areas indicates that, of the age network position, local connectivity, elevation, and
Earth’s 426 freshwater ecoregions, only about 15% enjoy some gradient can result in distinct aquatic assemblages and
form of protection (WDPA 2004; IUCN 2016). Where species population dynamics between and within streams and
data are deficient, regional habitat representation, a “coarse- lakes. Thus, it is an individual AES that can become the
filter” approach, serves as the primary tool for representing conservation target if it has high levels of regional
biodiversity in regional conservation planning. The Nature biodiversity, and if resources and circumstances permit
Conservancy (TNC), administering the largest system of pri- the preservation of the entire unit.
vate terrestrial refuges in the world, is also involved in global The fourth level of TNC’s system is the
conservation of freshwater aquatic ecosystems. As a first step macrohabitat, representing finer scale classification
toward organizing global information, TNC has developed a units that can be used to create the AES. Some parts of
coarse-filter classification for streams, lakes and rivers using
four spatial levels (Information Box 8.1). (continued)
8.3 Managing Freshwater Systems at Landscape Levels 321
a. One Aquatic Zoogeographic Unit
b. Ecological Drainage Units within

one Aquatic Zoogeographic Unit
c. Aquatic Ecological Systems within

one Ecological Drainage Unit
d. Macrohabitats within one

Aquatic Ecological System
Information Box Fig. 8.1 A four-tiered, hierarchical framework nested within EDUs (approximate scale 1:4,000,000). (d)
for classification of freshwater systems for conservation developed Macrohabitats are nested within AESs (approximate scale
by The Nature Conservancy. The highest level is (a) that of aquatic 1:1,200,000). (Reprinted by permission from Island Press, from
zoogeographic units (AZUs) (approximate scale 1:26,000,000). (b) Groves, C., 2003. Drafting a conservation blueprint: a
Ecological drainage units (EDUs) are nested within AZUs (approxi- practitioner’s guide to planning for biodiversity. Island Press, DC.
mate scale 1:26,000,000). (c) Aquatic ecological systems (AESs) are # 2003 Island Press)
the world do not have sufficiently detailed information of macrohabitats within an AES, these can serve as
on aquatic systems to identify individual macrohabitats conservation targets at smaller scales. The last two
within AESs. When that is the case, this level of classi- levels of classification consider how the physical envi-
fication can be omitted and targets for conservation ronment shapes local distribution patterns of aquatic
reserves designated by selecting appropriate AES organisms and thus are described using abiotic
units. If enough information exists for identification variables (Higgins et al. 2005).
(continued)
TNC uses these four levels because there are spatial data more important criteria for conservation. If two systems, A
available, mainly via remotely sensed data integrated into and B, contain similar levels of aquatic biodiversity in the
GIS, to classify and map ecological patterns at scales that same region, but A is under multiple threats of impending
are known to shape freshwater biodiversity patterns. Such a development and B, perhaps located in a relatively uninhab-
classification scheme can operate using either “top down” or ited area, is not, the strategic use of limited conservation
“bottom up” approaches. In regions where fine-scale, high- funding would be to purchase land in the watershed
quality hydrogeographic data and digital versions of other surrounding A because its biodiversity is more in need of
relevant data layers are available, the classification is immediate protection than B.
implemented by a “bottom up” approach, accomplished by
mapping relevant classification attributes in the hydrographic
data of a GIS file with a set of GIS decision-criteria 8.4 Wetlands, Pools and Ponds
algorithms created for this purpose. In contrast, a top-down
classification is used in areas where GIS data are insufficient 8.4.1 What Are Wetlands?
for identifying stream characteristics for individual habitats.
In these situations, AESs are defined based on similar envi- Wetlands (Fig. 8.13) have been defined as lands transitional
ronmental attributes (Fig. 8.12) (Higgins et al. 2005). This between terrestrial and aquatic systems where the water
approach provides TNC with rapid means to organize infor- table is at or near the surface or the land is covered by
mation at scales appropriate for ecoregional assessment. shallow water (Cowardin et al. 1979). Wetlands are distinc-
Although this approach provides a reasonable method for tive systems and habitats in three respects. First, they are
identifying conservation targets that can meet goals of unique in their hydrology – their characteristics of presence,
regional biodiversity representation, it does not assess the abundance, and sources or water. Hydrology interacts with
urgency of threat to such systems, which may be an even their second and third distinctive characteristics – their
Fig. 8.12 Example of a

top-down classification from a
portion of the Alto Cuiabá
ecological drainage unit in the
upper Paraguay River Basin, an
approach used in areas where GIS
data are insufficient for
identifying stream characteristics
at the level of individual habitats.
In these situations, the
macrohabitat classification is
omitted, but aquatic ecological
systems (AESs), such as those
shown in this drainage unit, are
defined based on similar
environmental attributes. Here
examples of different AES types
are represented as stream
networks. (Reprinted by
permission from Island Press,
from Groves, C., 2003. Drafting a
conservation blueprint: a
practitioner’s guide to planning
for biodiversity. Island Press,
Washington, DC. # 2003 Island
Press)
8.4 Wetlands, Pools and Ponds 323
Fig. 8.13 A wetland complex in

the Camargue Region of southern
France. Wetlands are important,
shallow water aquatic habitats
transitional between terrestrial
upland areas and other types of
aquatic habitats. (Photo courtesy
of A. Waterkeyn)
physico-chemical environment (geography, geology, and soil characteristics – the formation of hydric (water-logged) soils.
characteristics) and their unique biota (distinctive species of Due in part to their water-logged soils, wetlands are distinc-
plants and animals). Together, these three factors interact to tive in their abundance of hydrophytic (literally, “water-
form characteristic wetland habitat. For example, a surface loving”) vegetation. Because wetland soils are water abun-
depression (geographic formation) underlain by water- dant, wetland-adapted plants are often characterized by shal-
impermeable clay soils (geology and soil characteristics) low root systems, or even adventitious root systems in which
can facilitate the development of a wetland more easily than roots grow and function above ground. Such root systems
level terrain underlain by water-porous, glacially-derived often contain large air spaces, the better to store and move
sandy soils. If emergent vegetation in the wetland like sedges oxygen internally given that oxygen in the soil may be low or
(Carex spp.) and rushes (Juncus spp.), which have relatively absent.
low water demands, is replaced by extensive stands of cattail Wetlands typically exist between areas of open water
(Typha spp.), which have higher water demands, the water (lakes or streams) and upland areas whose soils are not
level and amount of open water in the wetland may decline. regularly or periodically saturated. Moving from open water
Beaver (Castor canadensis), when present, may build dams toward the upland area, the wetland system begins with areas
that reduce movement of flowing water through the wetland that are permanently flooded but lack open water because
and create larger open water areas. American alligator (Alli- they are dominated by vegetation, either in the form of trees
gator mississippiensis) will excavate depressions (“gator or shrubs (swamps) or herbaceous vegetation (marshes). Per-
holes”) that will retain water even during drought, thus manently flooded areas are replaced by periodically flooded
providing benefit for other species that might otherwise per- areas closer to uplands. Here vegetation is characterized by
ish or leave the wetland during dry periods. Muskrat species which can tolerate seasonal but not permanent
(Ondatra zibethicus), at high densities, can devastate (even flooding, like willow (Salix spp.), as well as animal groups
temporarily eliminate) emergent herbaceous vegetation like amphibians that can live as adults on land but require
(an event sometimes referred to as an “eat-out”) that can water for breeding and larval development. Moving still
radically change the wetland’s vegetation community. closer to the upland areas, wetlands may transition to areas
Together, these direct, indirect, and interactive effects drive with no open water but permanently saturated soils that
the formation of the wetland environment (Fig. 8.14). create distinctive vegetation communities. Prior to entering
The year-round presence and abundance of all types of a true upland habitat, soils may be periodically saturated near
wetlands not only maintains their unique ecological structure the surface. This condition, like that of permanently saturated
and function but drives the second of their most distinctive soils, can also lead to the formation of other different and
Fig. 8.14 Relationships among hydrology, physicochemical environ- and alligators can also significantly affect hydrology, soils, and other
ment, and biota in wetlands. Vegetation provides important feedback to biota. (Reprinted by permission from National Academies Press,
hydrology through evapotranspiration and increase in flow resistance from National Research Council (US), 1995. Wetlands: Characteristics
and to the physiochemical environment by affecting soil properties and Boundaries. National Academies Press, Washington, DC. # 1995
(organic content, dissolved oxygen) and elevation (accumulating National Academy of Sciences)
organic matter, trapping sediment). Animals such as beaver, muskrat,
unique vegetation communities (National Research Council in bog wetlands like peat is a wetland product used as fuel in
1995). many northern European countries, while actively growing
Wetlands support high levels of primary productivity and plants like wild rice (Zizania spp.) remain important food
biomass. Because water is shallow throughout the wetland staples for many Native American and First Nation
environment, all parts of the system can be photosyntheti- communities in the US upper Great Lakes states and Canada.
cally active, unlike deep water environments where light
cannot penetrate below certain depths. Because water levels
vary spatially and temporally (seasonally) within a wetland, 8.4.2 Managing Wetlands for Conservation
the environment experiences strong moisture gradients that
support a diversity of plant life characterized by plants having Water levels in wetlands can fluctuate widely. Therefore, the
unique life- and growth-forms, including carnivorous plants hydroregime of a specific wetland is a key ecological driver
like the pitcher plant (multiple families and genera), sundew defining characteristics of the system. A hydroregime is the
(Drosera spp.), and Venus fly trap (Dionaea muscipula). total contribution of different hydrological variables to the
Such plant diversity creates physical heterogeneity and com- stability of a temporary aquatic habitat (Vanschoenwinkel
plexity greater than most terrestrial environments, and so et al. 2009). These variables include, but are not limited to,
often supports a correspondingly more diverse animal com- (1) hydroperiod (the length of time there is standing water on
munity, often harboring high numbers of endangered species, the site, usually expressed as days per year); (2) desiccation
game animals, and furbearers. frequency (how often the site dries up); and (3) predictability
Plant communities in wetlands are valuable in both or regularity of alternating flooding and desiccation events.
services and products. They can absorb large quantities of A second consideration is habitat size, because the
water, thereby moderating variance in flow entering the wet- species-area relationship, an important predictor of species
land. Thus intact wetlands can mitigate flood effects and richness on a particular site (Chap. 2), can be important in
damage for downstream communities because they help determining the species richness of small wetlands, ponds,
maintain relatively stable downstream discharge even when and pools. In modeling experiments, hydroregime and habitat
experiencing high and variable levels of upstream input. size contribute substantially to differences in aquatic inverte-
Wetland vegetation and associated wetland systems can brate communities, explaining 47% of variation in relative
absorb greater quantities of nutrients, especially phosphorus invertebrate abundance and 59% of variation in species rich-
and nitrates, entering a lake’s drainage basin than can terres- ness. Elements of the hydroregime that increase species
trial vegetation, removing up to 79% of total nitrogen, 82% of abundance and richness are related to stability. Large deep
nitrates, 81% of total phosphorus, and 92% of sediment pools with long hydroperiods and low inundation frequency
(Chescheir et al. 1992). Slowly decomposing plant material support higher richness, while small shallow pools with short
hydroperiods, high inundation frequency and high variation Human disturbance often creates inhospitable areas between
in inundation frequency are associated with lower richness amphibian breeding areas in wetlands such as vernal pools and
(Vanschoenwinkel et al. 2009). appropriate sub-adult or adult terrestrial habitats. When this
Coordinated management of lake-wetland complexes can juxtaposition occurs, amphibians are forced to move through
produce more effective results for conservation than manag- areas of non-habitat to complete their life cycle and may suffer
ing each system separately. Managers can reduce inputs of disproportionately high mortality when doing so. This is a
phosphorus and other nutrients into a lake by maintaining or problem called “habitat split,” which scientists now consider
creating wetlands around it. As in island flora and fauna, as among the more important reasons for worldwide amphibian
the size of a given type of wetland increases, so does its decline. Habitat split has relatively small effects on amphibians
species richness. Thus, the conservation value of many with terrestrial larvae, but can depress populations of species
types of wetlands often rises with increasing area (Findlay with aquatic larvae, especially salamanders (Becker et al. 2007)
and Houlahan 1997), although this is not the case for all (Fig. 8.15). By affecting survivorship of individual amphibian
wetlands, such as vernal pools, small bogs, and prairie species, habitat split also affects structure and richness of entire
potholes which may harbor specialist species not found in amphibian communities.
larger wetlands. Because wetlands are often radically differ- Forest pools and amphibians that depend on them are
ent from their surrounding landscape, successful manage- difficult to manage and preserve. Because they are small
ment of wetland species may require management of and under forest canopies, they are difficult to detect remotely
landscape level processes that extend far from the wetland’s using aerial or satellite imagery. This problem is aggravated
borders. For example, Findlay and Houlahan (1997) deter- for vernal pools because vernal pools are ephemeral,
mined that, in southeastern Ontario, Canada, wetland species appearing and disappearing in different seasons. Since
richness in plants, herptiles (amphibians and reptiles), and amphibians using these pools have complex life cycles
birds was negatively correlated with the density of paved requiring both aquatic and terrestrial habitats, both kinds of
roads within 2 km of the wetland edge, and species richness habitat must be protected. Forest pools are especially vulner-
in wetland plants, herptiles, and mammals was positively able to stressors associated with human activities that degrade
correlated with the proportion of forest cover within the or destroy breeding and nonbreeding habitat, such as urbani-
same distance. Thus, a manager may be able to enhance zation, forestry operations, changes in temperature and pre-
biodiversity in a wetland as much by managing land use cipitation patterns (i.e., climate change) and spread of
processes around it as by managing the wetland itself. diseases that can be particularly deadly to amphibians.
Finally, forest pools can be difficult to manage because they
are easily overlooked among larger landscape elements like
8.4.3 The Special Case of Forest Pools: Critical streams, lakes, marshes, and forests. As a result, they do not
Elements for Amphibian Biodiversity always have a “constituency” of people who advocate their
protection. But, in some places, this is changing.
Seasonal and permanent forest pools are critical habitats for
maintaining amphibian biodiversity at landscape levels in
temperate climates, but one of the most difficult of all aquatic
habitat categories to protect. Seasonal, so-called “vernal”
(spring) pools normally fill during the spring or fall and
may disappear during summer. In contrast, permanent forest
pools have similar characteristics, but are more stable, year
round aquatic environments. Most permanent and vernal
forest pools have no persistent inlet or outlet stream, and
therefore no predatory fish. This makes them ideal habitats
and breeding areas for amphibians, who otherwise suffer high
mortality from fish predators.
Amphibians that breed in forest pools have complex life
histories requiring both terrestrial and aquatic habitats.
Among these species, larval amphibians spend most or all
of their time in wetland habitats, but sub-adult or adult
amphibians live in terrestrial upland areas. Efforts to protect
these species must focus on distribution and connectivity of Fig. 8.15 A red eft, the sub-adult life history stage of the eastern newt
multiple habitats within a landscape, especially connectivity (Notophthalmus viridescens), an example of the phenomenon of “habitat
split” in many forest-dwelling amphibians. The sub-adult red eft will
of forest pools and surrounding forest areas that provide live a terrestrial life in upland forests, then develop into a fully aquatic
habitat for adult life stages. adult. (Photo courtesy of US Geological Survey, public domain)
8.4.4 Engagement of Legislators states, while BMPs for development around vernal pools
and Stakeholders in Forest Pool have been used as a standard by the Army Corps of Engineers
Conservation: A US Case History and US Fish and Wildlife Service in New England. . .” (Hart
and Calhoun 2010:265). This work not only shed light on the
In the state of Maine (northeast USA), permanent and vernal importance of preserving these kinds of wetlands, but
forest pools are abundant and common landscape features. demonstrated a way of engaging broad community support
Many research studies have highlighted the importance of that led to their conservation.
such pools to native amphibian species. Such studies have The VPWG in Maine provides an example of politically
provoked increasing public awareness of the importance of and socially effective conservation effort at a local sphere of
vernal pools, an awareness that has begun to have political influence, but conservation efforts to protect wetlands are
effects. David Hart and Aram Calhoun are two conservation also at work at national and international levels. Internation-
scientists who have done extensive work with vernal pools ally, wetlands were one of the first ecosystems in a global
and worked effectively to mobilize the public for their con- agreement, the Ramsar Convention on Wetlands of Interna-
servation. They report that “Maine legislators and regulators tional Importance (1971, ratified in a meeting in Ramsar,
were receiving feedback from frustrated constituents about Iran), focused on the protection of an ecosystem instead of
the lack of coordination between federal and state wetland a species. Ramsar obligates its signers to land-use planning
regulations. In response, the Maine legislature passed a Leg- and preservation for wetlands, requiring them to establish
islative Resolve in 1993 setting up a Wetlands Task Force to national wetland nature reserves and designate at least one
recommend changes to the state wetland programme.” This wetland in their country as a “wetland of international impor-
task force led to formation of the Vernal Pool Working Group tance” (Koester 1989). As of 2018, there were 2314
(VPWG), “a multi-stakeholder group including designated sites covering more than 245 million hectares,
representatives from various federal and state agencies, the largest being Ngiri-Tumba-Maindombe in the Demo-
biologists/researchers, consultants and non-profit cratic Republic of Congo and Queen Maud Gulf in Canada,
organizations, to address the vernal pool issues that were each covering 60,000 square kilometers. (Ramsar 2018).
not adequately addressed in earlier legislation. . .” (Hart and
Calhoun 2010:264).
The VPWG realized that one problem of protecting vernal 8.4.5 Big Impacts of Small Habitat: Pond
pools was a lack of common definition among different Biodiversity
interest groups about what exactly a vernal pool was. Hart
and Calhoun, who were active participants in the process, Conservationists are habitually conditioned to think of “big”
noted that “our first step in developing a conservation strat- habitats as the most valuable conservation targets, but recent
egy was to create a working definition of vernal pools which research shows that small aquatic habitats may be dispropor-
was acceptable to a diverse array of stakeholders [the]. . .the tionately important to aquatic biodiversity, particularly in
VPWG developed a multi-pronged approach to meet the areas burdened with strong anthropogenic influences. In
mandate to improve conservation of pools; identify and fill western Europe, for example, conservation scientists have
gaps in knowledge (the science), educate stakeholders about conducted extensive surveys of aquatic habitat biodiversity
the resource and engage them in problem solving and at alpha (site specific), beta (habitat and landscape specific),
develop mechanisms (regulatory or non-regulatory) to con- and gamma (region specific) levels (Chap. 2) in a variety of
serve vernal pools” (Hart and Calhoun 2010:264). aquatic habitats, evaluating each for its absolute and relative
The VPWG avoided the common mistake of speaking contributions to landscape diversity at all of these levels
only to scientists in scientific terms. Instead, VPWG (Davies et al. 2008). They found that, at the level of individ-
presented scientific findings in non-technical publications ual sites (alpha diversity), rivers were the most species-rich
accessible to non-science stakeholders. Emphasis was on waterbodies, but, at regional levels (gamma diversity), ponds
best management practices (BMPs) to preserve the pools were the most species-rich aquatic habitat for both wetland
rather than technical research results. The process was inclu- plants and macroinvertebrates. In terms of rare species, ponds
sive, reaching out not only to traditional conservation groups, also had a higher rarity value, as measured by a Species
but groups whose work might most affect vernal pools, such Rarity Index, than other habitat types. Ponds are common
as loggers and foresters. As Hart and Calhoun describe it, in agricultural landscapes but have historically been consid-
“The inclusion of loggers and foresters in designing ered insignificant in their biodiversity contributions because
guidelines increased the level of ‘buy-in’ by stakeholders, of their small size, and therefore unworthy of conservation
including large industrial paper companies. . .Since their pub- effort. This might not be the case.
lication, the guidelines for forestry practices around vernal In a typical agricultural landscape in southern England, for
pools have been used as a model in other northeastern US example, small bodies of water like ponds (Fig. 8.16),
Fig. 8.16 A typical pond

embedded in an agricultural
landscape of western Europe.
Although small in size, recent
studies have demonstrated that
small water bodies like ponds and
ditches may make
disproportionately important
contributions to aquatic
biodiversity, especially in
environments affected by
agriculture or urban development.
(Photo reprinted under Creative
Commons CC0)
distributed throughout the landscape, contribute more to Table 8.2 Unique species contributed by ditches, lakes, ponds, rivers,
regional aquatic biodiversity than larger ‘downstream’ rivers, and streams from five locations in three biogeographic zones in western
with the largest proportion of the regional species pool occur- Europe 1997–2002
ring in ponds, the smallest type of waterbody surveyed Ditches Lakes Ponds Rivers Streams
(Williams et al. 2004). Ditches and small streams also proved Macrophytes
important reservoirs for biodiversity in this landscape, Avignon 6 1 10 3 3
despite being surrounded by areas subject to intensive land Braunschweig 5 1 45 1 2
uses (Davies et al. 2008). In particular, the Species Rarity Funen 5 1 42 7 2
Coleshill 3 0 22 10 3
Index (SRI) calculated for UK sites showed that ponds usu-
Whitchurch 10 – 36 – 8
ally had the highest mean values for both macrophytes and
Total 29 3 155 20 18
macroinvertebrates, as well as the highest recorded SRI
Macroinvertebrates
values and greatest range of values (Davies et al. 2008). In
Coleshill 11 0 55 24 1
regional species richness (gamma diversity), ponds were Whitchurch 14 – 50 – 27
again major contributors, supporting the highest values for Total 22 0 96 25 32
both macrophytes (155 unique species) and macroinver- Reprinted by permission from Elsevier, from Davies, B., Biggs, J.,
tebrates (96 unique species) (Table 8.2). Likewise, in beta Williams, P., Whitfield, M., Nicolet, P., Sear, D., Bray, S., Maund, S.,
diversity, ponds had the most dissimilar (unique) macrophyte 2008. Comparative biodiversity of aquatic habitats in the European
communities from one pond to another (Davies et al. 2008). agricultural landscape. Agriculture, Ecosystems and Environment 125,
1–8. # 2007 Elsevier B.V.
After examining these patterns of biodiversity distribu-
tion, principal investigator Bella Davies and her colleagues
noted “. . .smaller waterbodies are not simply inferior landscapes,” noted Davies et al., “are not ‘deserts’ for aquatic
versions of their larger counterparts in terms of species rich- biota: a range of aquatic species were supported in each
ness as has often been assumed. . .the relatively large ranges landscape and all of the surveyed waterbody types
in their [ponds’] physiochemical conditions provide a variety contributed to this biodiversity, with each supporting unique
of habitats which cumulatively results in greater species species. . .Encouraging the more extensive protection and
richness across the region . . .The classic species-area rela- creation of ponds throughout agricultural landscapes, so
tionship has not always been observed in lentic that a variety of successional stages are maintained, is likely
waterbodies. . ., with higher levels of biodiversity often to enhance the aquatic biodiversity of agricultural areas at the
being found in smaller waterbodies. . .” (Davies et al. landscape scale” (Davies et al. 2008:7).
2008:7). Consistent with the conclusions of Davies et al., Régis
Such patterns of biodiversity have implications for Céréghino and his colleagues, after making a comprehensive
gaining a right perspective on conservation in landscapes review of numerous studies of European ponds, similarly
markedly altered by human land use. “Agricultural concluded, “. . .there is growing evidence that ponds are
functionally different from larger lakes. . .and that, despite nearly 23,000 km2. Discharge from these watersheds is
their small size, they are collectively exceptionally rich in received into the Pacific Ocean through the waters of
biodiversity terms. . .Thus, ponds often constitute biodiver- Moreton Bay and associated estuaries (Fig. 8.17), all of
sity “hot spots” within a region or landscape, challenging which are of high conservation value, support significant
conventional applications of species-area models (“big is fisheries, and are extensively used for recreation and tourism.
best”) in practical nature conservation” (Céréghino et al. But the value of these waters is also highly utilitarian. The
2008:2). western drainage basins provide the region’s primary water
supply, not only for household and industrial use but also for
agriculture. Human impacts in the area have been significant
8.5 Policies and Practices that Protect since European settlement began in the 1700s. Today less
Freshwater Habitats than a quarter of the area still holds areas of intact native
vegetation, and there is much less than that along stream and
8.5.1 Connecting Stakeholders, Scientists, river corridors in more heavily used watersheds (Bunn et al.
and Policy Makers 2010).
Stuart Bunn, Director of the Australian Rivers Institute,
Given the array of problems and threats confronting the led a multi-disciplinary team of conservation scientists,
conservation of freshwater habitats, what practices and managers, and policy experts to not only document the cur-
policies, especially those influenced by conservation rent conditions of aquatic habitat in this region, but to
biologists and private “citizen conservationists,” can effec- develop an effective plan for watershed conservation that
tively move from a condition of understanding the threat to could engage the large and diverse community of
reducing or eliminating the threat? In Australia, the region of stakeholders with vested interests in the use and quality of
South East Queensland (SEQ) is the fastest growing area in these waters.
the country, with a human population of over three million. Rather than beginning with research, Bunn and his
When viewed from the standpoint of aquatic habitats, it colleagues initiated their efforts with extensive consultation
contains 15 major watersheds draining a combined area of and engagement with various interest groups, while at the
Fig. 8.17 Moreton Bay in South

East Queensland, Australia, the
receiving area for multiple
waterways affected by rapid
human population growth and
urban development. (Photo
courtesy of NASA Earth
Observatory and US Geological
Survey)
8.5 Policies and Practices that Protect Freshwater Habitats 329
same time developing a monitoring program that would not of the South East Queensland Healthy Waterways Partner-
merely document the decline of the watersheds, but use ship (SEQHWP). First, members created a clear expression
metrics that could provide predictive capability about which of their vision. “By 2026, our waterways and catchments will
direction the system was headed so that management be healthy ecosystems supporting the livelihoods and
responses to trends found in the watersheds would be accu- lifestyles of people in South East Queensland, and will be
rately informed and rightly directed. As Bunn et al. describe managed through collaboration between community, gov-
the effort in their own words, their intentions were “to derive ernment and industry” (Bunn et al. 2010: 226).
a shared understanding of the range of important environ- SEQHWP developed an Environmental Management
mental assets and values that need to be considered. . .Such Support System (EMSS) to understand how point and diffuse
an approach also requires an understanding of the probable sources of sediment and nutrients were influencing water
causal factors that influence the condition or health of impor- quality, and then determine whether water quality objectives
tant environmental assets and values. . .Without ‘diagnostic’ could be met downstream. They then designed and
capability, monitoring programmes cannot be used to guide implemented a comprehensive Ecosystem Health Monitoring
management intervention with any confidence” (Bunn et al. Programme (EHMP), conducting monthly sampling of water
2010:224). quality parameters at over 250 sites across the region (Bunn
et al. 2010). In such a process-oriented approach (Fig. 8.18),
the monitoring program informed scientists who incorporated
8.5.2 Forming an Issue-Driven Coalition: The monitoring data into models showing (i) how healthy aquatic
Healthy Waterways Partnership ecosystems function; (ii) how they are likely to respond to
disturbance; (iii) what biological components and processes
Through their work with stakeholders, Bunn and his in the system are best indicators of system response, and
colleagues formed the South East Queensland Regional therefore the best variables to monitor; and (iv) informed by
Water Quality Management Strategy, which led to formation monitoring data, how to determine best management actions
Fig. 8.18 Flow chart illustrating

the process developed to create
the freshwater Ecosystem Health
Monitoring Programme (EHMP)
for South East Queensland,
Australia. (Reprinted by
Bunn, S.E., Abal, E.G., Smith, M.
J., Choy, S.C., Fellows, C.S.,
Harch, B.D., Kennard, M.J.,
Sheldon, F., 2010. Integration of
science and monitoring of river
ecosystem health to guide
investments in catchment
protection and rehabilitation.
Freshwater Biology 55, 223–240.
# 2010 Blackwell Publishing
Ltd)
Table 8.3 Indicator groups of ecosystem health used in wetland evaluation in South East Queensland, Australia
Direct measures of stream
Ecosystem processes Water chemistry Biological patterns disturbance
Benthic metabolism (including GPP, Nutrient concentrations (water Structure and function of fish Measure of riparian
R24) & sediment) communities canopy cover
Catchment disturbance of the nitrogen Salinity/conductivity/ionic Structure and function of Assessments of channel
cycle (using δ15N) composition invertebrate communities integrity
Use of δ13C as a potential surrogate of Alkalinity/pH Structure of microbial community-
GPP Dissolved oxygen – snapshot ECOplates™
& diel
Water temperature – snapshot
& diel
Nitrogen cycling – denitrification Turbidity
Chlorophyll a – as surrogate of GPP
Algal bioassays
Reprinted by permission from Wiley, from Bunn, S.E., Abal, E.G., Smith, M.J., Choy, S.C., Fellows, C.S., Harch, B.D., Kennard, M.J., Sheldon, F.,
2010. Integration of science and monitoring of river ecosystem health to guide investments in catchment protection and rehabilitation. Freshwater
Biology 55, 223–240. # 2010 Blackwell Publishing Ltd
Table 8.4 Indicators recommended for the ambient ecosystem health monitoring program of rivers and streams in South East Queensland,
Australia
Indicator type Recommended indices
Water quality Conductivity
pH
Diel changes in temp (includes max & min)
Diel changes in DO (includes max & min)
Ecosystem processes GPP
R24
δ13C (aquatic plants)
Algal bioassay (controls)
Nutrient processes δ15N (aquatic plants)
Algal bio-assay (N + P/control)
Macroinvertebrates PET richness (edge habitats)
Stream invertebrate grade number average level (SIGNAL) score (edge habitats)
Family richness (edge habitats)
Fish Percentage of native species expected (PONSE)
% alien individuals
Fish assemblage O/E50
Reprinted by permission from Wiley, from Bunn, S.E., Abal, E.G., Smith, M.J., Choy, S.C., Fellows, C.S., Harch, B.D., Kennard, M.J., Sheldon, F.,
2010. Integration of science and monitoring of river ecosystem health to guide investments in catchment protection and rehabilitation. Freshwater
Biology 55, 223–240. # 2010 Blackwell Publishing Ltd
to enhance water quality. Individual models were then devel- assigned a score of one and sensitive taxa a score of 10, cal-
oped to show how different system components (“indicator culated by averaging scores for all families at a site. Three
groups,” Table 8.3) would respond to land-use disturbance indices that evaluated status and changes in the fish commu-
gradients measuring loss of riparian vegetation, increased nity also proved to have high sensitivity to the disturbance
water turbidity, levels of stream nutrient enrichment, and gradient. These were (i) Percentage of Native Species
other variables. Expected (PONSE) – the proportion of native species
Three indices proved more reliable than others in observed divided by the number of native species predicted
predicting the status of the aquatic system and its direction from a referential model based on relationships between
of change. These were (i) total richness – the number of environmental conditions and fish assemblages from mini-
macroinvertebrate families identified in each sample; mally disturbed reference sites, (ii) percentage of alien spe-
(ii) PET family richness – the insect orders Plecoptera, cies, i.e. the proportion of individuals in the sample that were
Ephemeroptera and Trichoptera which are collectively con- alien (i.e., introduced) species, and (iii) Fish assemblage)/E50;
sidered most sensitive to water quality and (iii) the SIGNAL the comparison of the species composition of the community
index (Stream Invertebrate Grade Number Average Level) - a observed against that predicted by a referential model
measurement in which pollution-tolerant invertebrate taxa are (Table 8.4.)
8.5 Policies and Practices that Protect Freshwater Habitats 331
All of this work was outstanding science, but the key to recognize the actual state of the watersheds in their own
real results in conservation could only come through linking jurisdiction. To accomplish this, Bunn and his colleagues
the scientific findings to policy formation, and that meant adopted a dramatic, media-oriented approach, the Catchment
getting the attention of policy makers and managers to Report Card (Information Box 8.2).
Information Box 8.2: Creating Policy Response: Information Box 8.2 (continued)
The “Catchment Report Card”
To ensure that scientific recommendations generated recognized from the beginning that distributing and
from the monitoring program of Australia’s South East publicizing Report Cards would cause anxiety among
Queensland Healthy Waterways Partnership had the public officials and agencies, but they were able to
best possible chance of actually being adopted as man- overcome the understandable reluctance to publicly
agement policies, Stuart Bunn and his colleagues knew receive the report card (in a televised ceremony)
that they needed a way to generate media-intensive because there was “broad acceptance of the need for
public attention on those recommendations and show open and transparent reporting to address public
that they had been conveyed to public officials. To do concerns about the health of regional waterways”
this, they developed the “Catchment Report Card.” As (Bunn et al. 2010:236).
Bunn et al. describe it, a “key element of the monitor- This approach is working. As Bunn and his
ing programme is the development and public presen- colleagues noted, “Since 2001, the Partnership has
tation of annual ‘Report Cards’ on the health of placed considerable emphasis on the science and mon-
waterways in the region. These are presented to itoring required to underpin the implementation of the
politicians and senior policy makers each year in a Healthy Waterways Strategy. Many of the management
public (televised) ceremony. . . where [a grade of] ‘A’ actions agreed to . . .have been completed or are well
reflects a catchment in near reference condition and ‘F’ underway, including multi-million-dollar upgrades to
is where the condition fails to meet the ecosystem wastewater treatment plants. This has led to a 40%
health objectives that underpin agreed values for the reduction in nitrogen loads to Moreton Bay and other
region’s streams and rivers. . .” (Bunn et al. 2010: 236) estuaries, despite a considerable increase in population
(Information Box Fig. 8.2). Bunn and his colleagues at the same time.” (Bunn et al. 2010:236).
Information Box Fig. 8.2 Presentation of a watershed-specific watershed quality and the need for conservation of aquatic habitats.
“Report Card” to public officials in South East Queensland, Pictured left to right, Healthy Land and Water Chairman Stephen
Australia, providing a “grade” for the condition and quality of Robertson, Healthy Land and Water Chief Executive Officer Julie
waterways in their district. Through such annual and televised McLellan and Environment Minister Dr. Steven Miles. (Photo cour-
ceremonies, conservationists have raised public awareness of tesy of Healthy Land and Water)
(continued)
The South East Queensland Healthy Waterways Partner- human activities that can alter and degrade them when their
ship focuses on maintaining health, quality, and biodiversity resources are extracted in destructive ways.
of an aquatic system in which human inputs and activities One way that structures of benthic (bottom-dwelling)
affect mainly freshwater habitats. But water in these systems communities can be altered is by the use of bottom trawling
eventually enters estuarine wetlands, both receiving from and nets (Fig. 8.19). Auster (1998) provides a picture of a location
giving input to a marine system, the Pacific Ocean. Marine on the bottom of the Gulf of Maine off the East Coast of the
systems, harboring their own unique array of world biodiver- United States before and after bottom trawling. The top
sity, must be considered carefully and separately in photograph (before) reveals a complex and diverse assem-
evaluating the threats that can degrade them, and the blage of creatures, including tubeworms, sponges, and many
strategies that can preserve their abundant life and resources. other forms of life. The bottom photograph shows the same
spot after a trawl net was dragged across it. The complexity of
the habitat has been obliterated, along with all its residents.
8.6 Marine Habitats and Biodiversity This vivid, visual example of marine habitat destruction
can be understood more generally through a conceptual
8.6.1 Destruction of Benthic Environments model of the effects of fishing gear upon different marine
habitats, such as might be found on a continental shelf.
When evaluating threats to marine ecosystems, there is logic Consider eight different categories, ranging, at the simplest
in starting from the bottom up, literally. Marine benthic level, from flat sand or mud to the most complex, piled
communities are among the most unique and diverse on boulders (Table 8.5). Auster (1998) assigned a “numerical
earth, physically and biologically, but they are vulnerable to complexity score” to each habitat category. Note that, as
Fig. 8.19 A portion of the

Atlantic Ocean bottom before
(top) and after (bottom) being
swept by a trawler net. Note that,
prior to trawling, a complex and
diverse community is present in
and on the sediments. After
trawling, it has been obliterated.
Wiley, from Auster, P.J., 1998. A
Conceptual Model of the Impacts
of Fishing Gear on the Integrity of
Fish Habitats. Conservation
Biology 12, 1198–1203. # 1998
Blackwell Publishing)
8.6 Marine Habitats and Biodiversity 333
Table 8.5 A classification of fish habitat types on the outer continental shelf of the temperate northwest Atlantic
Complexity
Category Description Rationale score
1 Flat sand and mud Areas with no vertical structure such as depressions, ripples, or epifauna 1
2 Sand waves Troughs provide shelter from current; previous observations indicate that species such as 2
silver hake hold position on the downcurrent sides of sand waves and ambush drifting
demersal zooplankton and shrimp
3 Biogenic structures Burrows, depressions, cerianthid anemones, hydroid patches; features that are created or 3
used by mobile fauna for shelter
4 Shell aggregates Provide complex interstitial spaces for shelter; also provide a complex, high-contrast 4
background that may confuse visual predators
5 Pebble-cobble Provide small interstitial spaces and may be equivalent in shelter value to shell aggregate, 5
but less ephemeral than shell
6 Pebble-cobble with Attached fauna such as sponges provide additional spatial complexity for a wider range of 10
sponge cover size classes of mobile organisms
7 Partially buried or Partially buried boulders exhibit high vertical relief; dispersed boulders on cobble 12
dispersed boulders pavement provide simple crevices; the shelter value of this type of habitat may be less or
greater than previous types based on the size class and behavior of associated species
8 Piled boulders Provide deep interstitial spaces of variable sizes 15
Note: Classification is based on Auster et al. (1995), Langton et al. (2008), Auster et al. (1996), and unpublished observations; habitat complexity
scores do not increase at a constant rate but reflect cumulative effects of structural components added at each succeeding level
Reprinted by permission from Wiley, from Auster, P.J., 1998. A Conceptual Model of the Impacts of Fishing Gear on the Integrity of Fish Habitats.
Conservation Biology 12, 1198–1203. # 1998 Blackwell Publishing
Fig. 8.20 A conceptual model of

the effects of fishing gear on sea
floor habitat. Note that increases
in fishing effort produce
disproportionately greater
reductions in habitat complexity
in complex habitats compared to
simpler habitats. (Reprinted by
Auster, P.J., 1998. A Conceptual
Model of the Impacts of Fishing
Gear on the Integrity of Fish
Habitats. Conservation Biology
12, 1198–1203. # 1998
Blackwell Publishing)
habitats become more complex, scores do not increase line- predicts that effects of fishing activity on habitat complexity
arly. For example, category 6, pebble-cobble with sponge are nonlinear (Fig. 8.20) – the more complex the original
cover, receives five (not one) additional points because it habitat, the greater the loss of complexity from trawling
contains elements of all previous categories plus dense emer- activity (Auster 1998).
gent epifauna (animals living on the surface of the seabed or
objects on it). Category 7 receives 10 points for containing all
the elements of category 6 plus 2 points for shallow boulder 8.6.2 Pollution in the Water Column
crevices and current refuges. Finally, category 8 receives an
additional 3 points for its addition of deep crevices (Auster Marine habitats are degraded from land-based sources. This
1998). The effect of intensive fishing activity, primarily indirect, but extensive degradation has multiple causative
trawls and dredges, is to reduce habitat complexity by agents, most of which directly affect the quality and
smoothing bedforms (habitat categories 1 and 2), removing characteristics of marine water itself (Table 8.6). Many of
epifauna (categories 3, 4, and 6) and removing or dispersing these, such as eutrophication, sedimentation, and thermal
physical structures (categories 5, 7 and 8). Such a model pollution, are proximity-based relative to sources of
Table 8.6 Some types of land-based pollutants that degrade marine habitats and ecosystems
Pollutant Potential threat to system
Herbicides May interfere with basic food chain processes by destroying or damaging zooxanthellae in
coral, free living phytoplankton, algal, or seagrass communities.
Can have serious effects even at very low concentrations.
Pesticides May selectively destroy or damage elements of zooplankton or benthic communities;
planktonic larvae are particularly vulnerable.
May through accumulation in animal tissues have effects on physiological processes such as
growth, reproduction, and metabolism.
May cause immediate or delayed death of vulnerable species.
Antifouling paints and agents May selectively destroy or damage elements of zooplankton or benthic communities.
Likely to be a significant factor in harbors, near shipping lanes, and in enclosed, poorly mixed
areas with heavy recreational boat use.
Sediments and turbidity May smother substrate.
May smother and exceed the clearing capacity of benthic animals, particularly filter feeders.
Reduce light penetration, likely to alter vertical distribution of plants and animals in shallow
communities such as coral reefs.
May adsorb and transport other pollutants.
Petroleum hydrocarbons A wide range of damaging effects depending upon type of hydrocarbon, dilution, weathering,
dispersion, emulsification or interaction with seawater or other chemicals.
Direct contact with living tissue usually results in local necrosis and, with longer exposure,
death.
Exposure to water-soluble hydrocarbons results in mucus production, abnormal feeding,
changes to a wide range of physiological functions, and with longer exposure, death.
Detrimental effects on reproduction and dispersion; premature discharge of larvae, distorted
larvae, decreased larval viability.
Residual hydrocarbons in substrates may lead settling larvae to avoid affected areas, and thus
block recolonization and repair.
Sewage-detergent phosphates Inhibit a wide range of physiological processes and increase vulnerability of affected biota to a
range of natural and human induced impacts.
Inhibit calcification, e.g., in corals and coralline algae.
Can cause effects at very low levels.
Sewage and fertilizers—Nitrogen Increased primary production in phytoplankton and benthic algae distorts competitive and
predator/prey interactions in biological communities in areas such as coral reefs, which are
characterized by very low natural nitrogen levels.
Reduced light penetration through absorption and turbidity of increased planktonic
communities.
Increased sedimentation of detritus from planktonic communities.
Increased nutrient levels in benthos from sedimentary organic material.
Selectively favors growth of some filter or detritus feeders such as sponges and some
holothurians.
Some species such as corals are affected at very low levels.
High or low salinity water—Freshwater runoff, Low salinity water floats on top of water column, high salinity water sinks, prior to mixing and
effluents dispersion.
Tolerance of species highly variable so changed regime may alter biological communities,
particularly those in shallow, poorly mixed or enclosed waters.
Salinity is a key factor in settlement and physiological performance of many shallow benthic
and reef organisms.
May (e.g., for corals) cause physiological stress evidenced by elevated mucus production,
expulsion of zooxanthellae, or death.
High or low temperature water—From Tolerance of species highly variable so changed regime may alter biological communities,
industrial plant heating or cooling particularly those in shallow, poorly mixed or enclosed waters.
Temperature is a key factor in settlement and physiological performance of many shallow
benthic and reef organisms.
(continued)

Pollutant Potential threat to system
Heavy metals, e.g., mercury, cadmium May be accumulated by, and have severe effects upon, filter feeders and, by accumulation up
the food chain, pass these effects to higher predators.
Can interfere with physiological processes such as the deposition of calcium in skeletal tissue.
Surfactants and dispersants Most are toxic to marine biota.
Synergistic effects of dispersant/hydrocarbon mixes can be more toxic than either component
unmixed.
Can interfere with a wide range of physiological processes, e.g., photosynthesis.
Chlorine At low levels inhibits external fertilization of some invertebrates, e.g., sea urchins.
Can be lethal to many species.
Reprinted by permission from Taylor and Francis Group LLC Books, from Kenchington, R.A., Managing marine environments Taylor & Francis,
New York. # 1990 Taylor and Francis
pollution, and thus have their greatest effects on coastal and dollars by Short et al. 2011), greater in value than many other
estuarine environments. But others, such as radioactive terrestrial and marine areas. Seagrass habitats support arti-
wastes and persistent toxins such as PCBs, DDT, and similar sanal fisheries and provide livelihoods for millions of people
compounds, can travel long-distances in ocean currents or be in coastal communities, especially in tropical regions.
deposited far out to sea through atmospheric circulation Seagrass is an important food source for marine animals,
patterns. Likewise, some kinds of military wastes like radio- including many of conservation concern like the dugong
active material or chemical weapons may be transported long (Dugong dugon), green sea turtle (Chelonia mydas), and
distances from shore before being deposited. These pollutants Cape seahorse (Hippocampus capensis). Leaves of seagrass
can cause habitat destruction and devastate populations act as filters, clearing water of suspended sediments, and their
thousands of miles from their source of origin. More recent leaves, roots, and rhizomes take up and cycle nutrients. The
challenges also include rising temperatures and ocean acidi- complex root structure of seagrass beds secures and stabilizes
fication, both results of human-caused climate change sediments, providing essential shoreline protection and
(Chap. 4). reduction of coastal erosion from extreme storm events. For
Although problems of marine habitat and species preser- biological components of the marine community, seagrass
vation show expected local and regional variation, major leaves serve as surfaces for attachment for encrusting algae
threats to marine environments are consistent throughout and small animals, which in turn provide food for larger
the world. Some are similar to those in freshwater seagrass-associated animals. Seagrass beds are a nursery
environments. Others are unique to marine systems. We ground for juvenile and larval stages of many commercial,
now examine the most important of these in greater detail. recreational, and subsistence fish and shellfish. But for all
their value, seagrasses are not yet valuable enough to pre-
serve. The area they occupy declined at a rate of 110 km2 per
8.6.3 Habitat Destruction in Shallow Water year between 1980 and 2006. The rate of decline has been
Environments – The Plight of Seagrass accelerating (Short et al. 2011).
8.6.3.1 Seagrass as a Component of Marine 8.6.3.2 Seagrasses as Species of Conservation

Ecosystems Concern
Seagrasses (Fig. 8.21) are flowering plants that form vast Nearly one-quarter (15 species, 21%) of all seagrass species
underwater meadows throughout the world’s coastal oceans, are classified by the IUCN as threatened (Endangered or
supporting a myriad of marine food webs, providing habitat Vulnerable) or Near Extinction (Table 8.7). Three species
for many coastal species, and playing a critical role in the have been listed as Endangered, five as Near Threatened.
equilibrium of coastal ecosystems and human livelihood Twenty-two species (31%) have declining populations.
(Short et al. 2011). Although seagrasses themselves create Declining seagrass species are found worldwide, particularly
important marine habitats, they are also elements of more north of the equator in the most developed parts of the world,
complex ecosystems, contributing to the health of coral reefs but also in Australia and throughout the Indo-Pacific
and mangroves, salt marshes, and oyster reefs. The value of (Fig. 8.22) (Short et al. 2011).
their ecosystem services has been estimated at $34,000 USD Coastal seagrass beds are degraded, damaged, and
per hectare per year (Costanza et al. 1997, adjusted to 2010 destroyed as a result of human development and associated
Fig. 8.21 Seagrass (various

genera and species), a keystone
species that provides foundational
habitat for multiple species of
marine species, both vertebrate
and invertebrate, especially in
shallow water areas. (Photo
courtesy of P. Colarusso/US
Environmental Protection
Agency)
Table 8.7 Number and percent of all seagrass species listed in each IUCN Red List Category (n ¼ 72) and number and percent of population trends
(increasing, decreasing, stable, or unknown) in each category
Red List category No. of species Increasing Decreasing Stable Unknown
Endangered 3 (4%) 0 3 (100%) 0 0
Vulnerable 7 (9.5%) 0 7 (100%) 0 0
Near threatened 5 (7%) 0 5 (100%) 0 0
Least concern 48 (67%) 5 (10%) 6 (13%) 29 (60%) 8 (17%)
Data deficient 9 (12.5%) 0 1 (11%) 0 8 (89%)
Reprinted by permission from Elsevier, from Short, F.T., Polidoro, B., Livingstone, S.R., Carpenter, K.E., Bandeira, S., Bujang, J.S., Calumpong, H.
P., Carruthers, T.J.B., Coles, R.G., Dennison, W.C., Erftemeijer, P.L.A., Fortes, M.D., Freeman, A.S., Jagtap, T.G., Kamal, A.H.M., Kendrick, G.
A., Kenworthy, W.J., La Nafie, Y.A., Nasution, I.M., Orth, R.J., Prathep, A., Sanciangco, J.C., van Tussenbroek, B., Vergara, S.G., Waycott, M.,
Zieman, J.C., 2011. Extinction risk assessment of the world’s seagrass species. Biological Conservation 144, 1961–1971. # 2011 Elsevier Ltd
pollution and habitat destruction. The primary impact to including hydraulic dredging (a technique used to harvest
seagrasses from such activity is loss of water clarity and clams in large quantities) (Figs. 8.24a, c), scarring of plants
quality due to eutrophication accompanied by sediment load- and their substrates by boat propellers (Fig. 8.24b), and
ing. Damaging fisheries practices such as trawling and aqua- habitat loss from aquaculture operations (Fig. 8.24d). By
culture also harm seagrass habitats, as does mechanical the mid-1970s, eelgrass in Chesapeake Bay had been
damage from boats and associated installation of moorings eliminated from half of its historically habitable area. By
and docks. Diseases, such as wasting disease, threaten some this time, scientists were reporting a decline of all species
seagrasses, causing large scale declines (Short et al. 2011). of submersed aquatic vegetation in Chesapeake Bay,
One species of seagrass, commonly called eelgrass prompting resource managers, politicians, environmental
(Zostera marina), has become the subject of intensive con- groups, and the general public to mobilize a major effort to
servation and restoration efforts. In the Chesapeake Bay improve overall water quality. One part of the resulting
region of the Atlantic Ocean bordering the US states of conservation initiative was an effort to restore eelgrass to
Maryland and Virginia, eelgrass grows in a variety of sedi- where it was no longer present but had historically occurred
ment types in water 2–10 meters deep. Eelgrass has been (Orth et al. 2010).
declining in Chesapeake Bay since the late 1960s (Fig. 8.23), One portion of the Chesapeake Bay, the Virginia coastal
with losses attributed to deteriorating water quality and bays, suffered extensive losses of eelgrass during this period.
increased nutrient loading. The latter problem results in over- Large-scale eelgrass restoration in these bays had been tried
growth of macroalgae on eelgrass leaves, reducing their earlier, but these efforts had transplanted adult plants onto
ability to photosynthesize and respire. Eelgrass also has targeted areas. Most transplants died within 5 years of plant-
been damaged in direct, physical ways by human activities, ing, many within 2 years. Conservationists changed tactics
Fig. 8.22 Global distribution of seagrass species in stable and declin- Fortes, M.D., Freeman, A.S., Jagtap, T.G., Kamal, A.H.M., Kendrick,
ing populations in (1) Temperate North Atlantic, (2) Tropical Atlantic, G.A., Kenworthy, W.J., La Nafie, Y.A., Nasution, I.M., Orth, R.J.,
(3) Mediterranean, (4) Temperate North Pacific, (5) Tropical Indo- Prathep, A., Sanciangco, J.C., van Tussenbroek, B., Vergara, S.G.,
Pacific and (6) Temperate Southern Oceans ecosystems. (Reprinted by Waycott, M., Zieman, J.C., 2011. Extinction risk assessment of the
permission from Elsevier, from Short, F.T., Polidoro, B., Livingstone, S. world’s seagrass species. Biological Conservation 144, 1961–1971.
R., Carpenter, K.E., Bandeira, S., Bujang, J.S., Calumpong, H.P., # 2011 Elsevier Ltd)
Carruthers, T.J.B., Coles, R.G., Dennison, W.C., Erftemeijer, P.L.A.,
and began using eelgrass seeds instead of transplants, with cup-shaped structures that serve as their homes and that they
resulting success. On seeded sites, eelgrass had reoccupied attach to one another. Over many years and generations of
591 ha (over 1500 acres, more than two square miles) of coral, these calcium carbonate secretions build a coral reef,
Virginia coastal bays by 2008 (Orth et al. 2010) (Fig. 8.25). each new generation enlarging the reef by building on the
bodies of their departed ancestors.
Coral reef ecosystems are centers of biodiversity because
8.6.4 Habitat Destruction and Marine they combine elements of structure, nutrients, water quality,
Biodiversity: Threats to Coral Reefs and light to create a favorable and productive environment
for living things. Physically, the body of the reef provides a
Coral reefs (Fig. 8.26) have been called the tropical substrate and point of attachment for many species, espe-
rainforests of the oceans. Worldwide coral reefs cover < 1% cially more sedentary species groups such as crustaceans
of the world and less than 10% of the ocean surface area but and mollusks. Even among more active species, physical
contribute over 25% of the Earth’s biodiversity. Between characteristics of the reef provide cavities for shelter and
35,000 and 60,000 reef-dwelling species have been breeding. Upon this structure, high densities of prey species
described, but, at the current rate of species’ discovery, it attract proportionally high densities of predators.
has been estimated that the total number of reef-associated Reef-building corals take up dissolved calcium from sea-
species is somewhere between one million and nine million, water and accrete it to produce the reef substrate. Because the
all of this habitat constructed by fewer than 1000 species of reef forms in well-lit waters, light is available in combination
reef-building corals (Microdocs 2012). Although the bulk of with calcium and other nutrients. Interacting with nutrient
any coral reef is non-living matter, the surface layer of living and light availability is generally high water quality, pro-
creatures is composed mostly of coral polyps. Relatives of duced in part by abundant populations of sponges on the
jellyfish and anemones, coral polyps have column-shaped reef’s surface. Sponges, using the reef for support, circulate
bodies topped with stinging tentacles. These polyps secrete and cleanse surrounding water through their own bodies,
calcium carbonate as a metabolic product, and from it fashion enhancing water quality, lowering turbidity, and increasing
Fig. 8.23 Eelgrass (Zostera marina) distribution in the US Chesapeake a result of restoration efforts in the Virginia Coastal Bays. (Reprinted by
Bay region from 1984 to 2007. Shaded polygon shows extent of eelgrass permission from Springer Nature, from Orth, R.J., Marion, S.R., Moore,
distribution through 1960. Closed polygon shows distribution in 2007. K.A., Wilcox, D.J., 2010. Eelgrass (Zostera marina L.) in the
Red and green dots show eelgrass transplant sites from 1978 to 2006. Chesapeake Bay Region of Mid-Atlantic Coast of the USA: Challenges
Note from the bar graphs that eelgrass declined during this period in in Conservation and Restoration. Estuaries Coasts 33, 139–150. #
Chesapeake Bay and nearby Chincoteague Bay, but began to increase as 2009 Coastal and Estuarine Research Federation)
Fig. 8.24 Aerial photographs sowing impacts to eelgrass (Zostera habitat has been lost from aquaculture operations. (Reprinted by permis-
marina) from human activities including scaring caused by hydraulic sion from Springer Nature, from Orth, R.J., Marion, S.R., Moore, K.A.,
dredging for clams (a, c), boat propellers (b), and habitat loss and limits Wilcox, D.J., 2010. Eelgrass (Zostera marina L.) in the Chesapeake Bay
to bed expansion by hard clam aquaculture. In (c), arrows associated Region of Mid-Atlantic Coast of the USA: Challenges in Conservation
with numbers 1, 2, and 3 point to circular scars formed by a dredge being and Restoration. Estuaries Coasts 33, 139–150. # 2009 Coastal and
pulled by a small boat being driven in circles, leaving only a small Estuarine Research Federation)
central area still occupied by an intact eelgrass bed. In (d), eelgrass
transparency of the water to allow penetration of light to Ocean, 46% of living coral on reefs have been killed. Each
greater depths. year, as the world human population adds over 80 million
Coral reefs are, for all their beauty and diversity, fragile people to its current level of 7.7 billion, most of the new
systems. They can be degraded or destroyed by disease, humans will be added in coastal areas of developing
bleaching, sediment, pollution, overfishing, or direct destruc- countries, in immediate proximity to the world’s great reef
tion by natural causes like hurricanes or human activities like systems. But the developed world adds to reef degradation as
mining (for minerals in the reef) or blasting (an effective but well. As coral reefs become more popular designations for
destructive method of harvesting fish). They are today in a western “ecotourists,” abuses associated with divers touching
global state of accelerating decline, with 10% of the world’s or walking on reefs, or physically breaking off pieces of coral
reefs now considered degraded, and projections for another for souvenirs, becomes an increasingly serious problem,
30% expected to be lost in the next 20 years. Ten to 16% of leading to the degradation of some of the world’s most
coral reefs worldwide have been destroyed. In the Indian complex reef structures.
Fig. 8.25 Aerial photographs of an eelgrass (Zostera marina) restora- beyond the initial seed plots to form a contiguous distribution covering
tion site in South Bay, one of four Virginia (USA) coastal bays most of the polygon. (Reprinted by permission from Springer Nature,
associated with the Chesapeake Bay area where eelgrass restoration from Orth, R.J., Marion, S.R., Moore, K.A., Wilcox, D.J., 2010. Eel-
efforts have been undertaken to extend the distribution of eelgrass. grass (Zostera marina L.) in the Chesapeake Bay Region of Mid-Atlantic
The polygons in each photo (a – 2004 and b – 2008) enclose an area Coast of the USA: Challenges in Conservation and Restoration.
of approximately 300 ha. Dark squares in (a) are 0.4 ha eelgrass plots Estuaries Coasts 33, 139–150. # 2009 Coastal and Estuarine Research
seeded in 2001 and 2002. Note that, by 2008, eelgrass has spread Federation)
Two threats to coral reefs deserve elaboration. Bleaching Destruction of coral reefs also occurs through blast fish-
is a response that occurs in temperature-stressed corals. When ing, a form of direct destruction of coral reefs in which
bleaching occurs, there is loss of endosymbiotic explosives are used to harvest fish from the reef. A single
dinoflagellates (zooxanthellae) living within the coral polyp blast can devastate thousands of cubic meters of coral reef,
that occurs as part of the coral’s stress response to tempera- destroying not only individual fish but the structure upon
ture (Lesser 2011). It does not take much temperature which the community depends. Such blasting destroys what
increase to cause it. Studies of bleached corals reveal that may have taken hundreds or thousands of years for marine
water temperatures as little as 1–2 C above normal organisms to build. Blast fishing, even though now outlawed
maximums, if sustained for even a few weeks, will lead to by most governments, is still practiced illegally. But there are
bleaching (Spalding 2004). Although recovery is possible, methods of managing, conserving, and rehabilitating
“recovered” corals typically show fewer polyps, less surface “blasted” corals that can be successful on local scales.
area, slower growth, reduced competitive ability, and greater
susceptibility and mortality to other diseases. Bleaching was
unknown in corals prior to the 1970s but is now a worldwide 8.6.5 Rehabilitation Techniques for Coral Reefs
problem (Fig. 8.27). By May 2017, the third global coral
bleaching event had ended, but it was the longest, most 8.6.5.1 Biological Approaches – Reef Building
widespread, and possibly most damaging coral bleaching with Coral Transplants
episode to date (NOAA 2017). Heat stress during this event One method that has long been used to restore coral reefs is
also caused mass bleaching in reefs never bleached before, known as “coral gardening.” Living corals, are grown in an
such as the northernmost part of the Great Barrier Reef near aquatic laboratory (a coral “nursery”) and then placed on
Australia. Current models of worldwide ocean temperature artificially constructed structures near coral reefs (Fig. 8.28)
change (Chap. 4) estimate that all six of the world’s major or directly on damaged reefs that have lost their original
coral reef regions will exceed thresholds for bleaching corals. Using this approach, living corals are transplanted
between 2030–2050. Under these projected temperature from one location to another, and reef building is accelerated
regimes, coral bleaching will become an annual event by using electrolysis (passing an electric current through
(Hoegh-Guldberg 2004). water currents near the reef) to increase deposition of calcium
carbonate and enhance growth of existing or transplanted

corals. This approach has been most widely used in the
Caribbean Sea and adjoining western portions of the Atlantic
Ocean where it has enjoyed increasing success to the point
that today “coral gardeners” grow and “outplant” tens of
thousands of corals onto degraded reefs each year (Lirman
and Schopmeyer 2016).
A further advance in this approach has come through the
application of the technique of micro-colony fusion in which
fragments of different colonies can be fused together to
produce larger colonies with faster rates of growth. Like
many other organisms, size in corals is correlated with survi-
vorship, fecundity, and increased competitive ability in
interactions with other corals. In corals, smallest size classes
often have the highest rates of mortality. However, corals of
identical or related genomes can fuse with one another.
Fusion of multiple small coral colonies increases their rates
of growth, permitting them to obtain more resources from
their substrate and environment, occupy more physical space,
reach needed size for sexual maturity and reproductive capac-
ity, and become less vulnerable to damage or predation
(Forsman et al. 2015).
David Vaughn, a pioneer in the development of micro-
colony fusion in corals, and his colleagues have studied the
performance of coral fragments developed from colonies
placed on ceramic tiles in laboratory aquaria which had
opportunity to fuse as they increased in size. The initially
small and separate colonies grew, fused and expanded, with
larger fragments then growing at more rapid rates. Perhaps
most importantly, none of the initially placed fragments died
or detached from the tiles (Forsman et al. 2015). Experimen-
tal “in-ocean” tests and applications of micro-colony fusion
are ongoing, and it holds potential for making efforts in coral
gardening even more effective.
8.6.5.2 Structural Approaches – Restoration

with Artificial Reefs
Indonesia’s Komodo National Park (KNP), a group of small
tropical islands in south-central Indonesia, contains some of
the world’s most beautiful and diverse coral reef systems. It is
also an area where reefs have been devastated by blast fish-
ing, leaving behind “reefs” that are little more than piles of
rubble on the ocean floor. Coral reef restorationist Helen Fox
and her colleagues at the University of California (USA) –
Berkeley, working with members of The Nature
Conservancy’s Coastal and Marine Program in Indonesia,
have been engaged in an extended rehabilitation effort to
Fig. 8.26 A coral reef in the Indian Ocean, exemplary of the high restore such damaged systems. The previously described,
biodiversity of marine species associated with these systems. (Reprinted biologically-based approach of coral gardening has had suc-
by permission from Springer Nature, from Graham, N.A.J., Jennings, S.,
MacNeil, M.A., Mouillot, D., Wilson, S.K., 2015. Predicting climate-
cess, but gardening techniques can be expensive, labor inten-
driven regime shifts versus rebound potential in coral reefs. Nature 518, sive, and sometimes experience high mortality in coral
94–97. # 2015 Macmillan Publishers Limited) transplants. Costs for restoration efforts using these
Fig. 8.27 Patterns of bleaching projected for coral reefs along the in which coral is remnant (< 5% coverage of reef), based on assumption
southern coast of (a) Jamaica, (b) Phuket, and (c) Tahiti from 1860 that coral communities cannot survive three or more severe bleaching
through 2100. Sea temperature data from the ECHAM4/OPYC3 model events per decade, informed by observations in Okinawa, Palau,
(Roeckner et al. 1996). Left side shows accumulated Degree Heating Seychelles, and Scott Reef in 1998. (Reprinted by permission from
Months (DHM) values once they exceeded 0.5 DHM. Right side shows Springer Nature, from Hoegh-Guldberg, O., 2004. Coral Reefs and
frequency of bleaching events per decade over next century Projections of Future Change, in: Rosenberg, E., Loya, Y. (Eds.),
(DHM > 0.5, solid line) and severe events (DHM > 3.2, dotted line). Coral Health and Disease. Springer, Berlin, Heidelberg, pp. 463–484
In right side graphs, left column shows period in which corals are in # 2004 Springer-Verlag Berlin Heidelberg)
decline (coral cover beginning to decrease). Right column shows period
techniques can range from US$13,000/ha to more than US measured colonization success of new corals on wide-mesh
$100 million/ha (Fox et al. 2005). These are prices that, fishing net attached to rock rubble, cement slabs pinned to
today, adjusted for inflation, can be beyond the budgets of rubble, and piles of rocks on top of rubble (Fox et al. 2005).
most conservation organizations and even many government Rock stabilization plots were the most successful, followed
agencies, especially in Majority World countries. by cement slabs, then netting. Untreated rubble did worst
Rather than using a biologically-based approach to reef (Fig. 8.29).
restoration, Fox et al. took a structural approach, conducting Although this experiment was a useful first step, coral
an experiment on reef restoration using low-cost materials rehabilitation cannot be successful using one square meter
obtained locally in Indonesia. In 1 square meter plots, they plots because, in areas of strong currents, rock rubble piles
8.7 Overexploitation of Marine Populations 343
m2, clearly unreasonable for large-scale rehabilitation in

developing countries. . .” (Fox et al. 2005:105). In their
study, rehabilitation costs were approximately US$5 per
square meter, a figure that included materials, transportation,
boat rentals, and labor. Although such rehabilitation
techniques hold promise for saving coral reefs at some
locations, they are of little value if reserve management is
ineffective at stopping destructive practices. Additionally, for
all the threats and problems faced in the conservation and
restoration of coral reefs, one great advantage is that coral
polyps stay put. More mobile marine species, especially
marine vertebrates, are also in need of conservation, protec-
tion, and restoration, but require different strategies for suc-
cess. First, however, we must understand their problems.
8.7 Overexploitation of Marine Populations

Fig. 8.28 A “coral tree” in the Atlantic Ocean off the coast of the US
state of Florida. Coral trees and other artificial structures are used to 8.7.1 The Collapse of Marine Fisheries
propagate corals that can then be transplanted to damaged or degraded
coral reeds. (Reprinted under CC-BY 4.0, from Lirman, D., In 2014, Marine biologist Rebecca Lewison and her
Schopmeyer, S., 2016. Ecological solutions to reef degradation: colleagues, after an extensive global survey of the world’s
optimizing coral reef restoration in the Caribbean and Western Atlantic.
PeerJ 4, e2597) marine fisheries, reported to the US National Academy of
Sciences that, since 1950, total world fisheries production
had increased from 19.3 million tons to more than 154 million
this small will be buried or broken into pieces over time. tons (Lewison et al. 2014). This level of harvest is not
Moving to a larger scale, Fox et al. created 100 square meter sustainable, a fact that is increasingly evident in the reality
(10 m 10 m) rock rubble piles and monitored their coloni- of having 63% of global fish stocks being now classified as
zation by corals. Results were encouraging. “Recruitment of overfished or collapsed, and increasingly leading to the phe-
hard coral and cover increased significantly in the mid-scale nomenon known as “trophic downgrading,” characterized by
studies. The rock piles quickly developed a ‘biofilm’ and massive declines or complete loss of megafauna in marine
were colonized by coralline algae and other encrusting ecosystems (Lewison and others 2014).
organisms” (Fox et al. 2005:102). Given the success of rock In the 1940s and 1950s, the emerging science of fisheries
rubble piles this size, Fox et al. increased the size of their management perceived fish stocks as renewable resources
colonization sites again, transforming approximately 6,430 that could be managed for maximum sustainable yield
square meters of dead coral rubble into four highly structured (MSY), a value that could be calculated precisely by various
designs at four different locations. On sites this size, Fox et al. means, primarily using estimates based on catch per unit
noted that “Scleractinian [hard coral] recruits quickly settled effort (Ricker 1958). All that was thought to be required for
on the rock piles, with considerable recruitment of hard corals a sustainable fishery was reproductive surplus. Today the
after approximately 1 year” (Fox et al. 2005:104). Overall, concept of MSY has all but disappeared from fisheries sci-
Fox and her colleagues demonstrated that coral recruitment ence, along with many of the fish stocks mismanaged under
could be enhanced by creating stable, spatially complex its assumptions. Biologists have gradually learned that most
structures high enough above reef rubble to minimize burial fish populations (1) show wide fluctuations of high and low
and abrasion. Recruitment on structured rock piles was more abundance; (2) do not always show a strong correlation
than 20 times that of untreated rubble and showed substantial between recruitment and number of adults present, and
recruitment of hard corals in as little as 6 months. Although (3) do not necessarily show advance warning of impending
labor was required to create appropriate structure, materials population decline or crash from overexploitation (Hilborn
were ready at hand, available at little or no cost. “Economi- et al. 1995). Rather, declines may be sudden, and stocks may
cally,” Fox et al. noted, “substrate stabilization using locally not recover in the short-term even when given complete
available rock compares favorably with other methods. . ., protection.
rehabilitation treatments in the Maldives cost from US$40 Exploited as they are, marine ecosystems show resiliency
to US$160/m2, and rehabilitation projects in the Florida Keys when given opportunity to recover. Among the large marine
National Marine Sanctuary cost from US$550 to US$10,000/ ecosystems of the world, the US Northeast Shelf Ecosystem
Fig. 8.29 Recruitment and

growth of corals onto 1 m2
treatments of cement, netting,
rock piles, and untreated sites on a
formerly blasted coral reef in
Komodo National Park, Indonesia
from sp. (spring) 1998 through
sp. 2001. (a) Mean and SE of
coral recruits plot and (b) mean
and SE of total area (cm2) covered
by coral recruits per plot.
Fa ¼ fall. (Reprinted by
permission from Wiley, from Fox,
H.E., Mous, P.J., Pet, J.S.,
Muljadi, A.H., Caldwell, R.L.,
2005. Experimental Assessment
of Coral Reef Rehabilitation
Following Blast Fishing.
Conservation Biology 19, 98–107.
# 2005 Society for Conservation
Biology)
has historically been one of the most productive, and most became mandated. Total allowable catch levels (TACs) for
heavily exploited, of all ocean fisheries. Stocks in this fishery species identified as “depleted” were reduced. Going further,
had become so depleted by the 1990s that, by 1994, days-at- the US Congress approved a plan for buying 79 groundfishing
sea for fishing trawlers were reduced to 50% of pre-1994 vessels from their owners, further reducing fishing effort
levels (Sherman et al. 2003). The problem was especially (Sherman et al. 2003). With the passage of the US Sustain-
acute among bottom-dwelling or demersal species, so-called able Fisheries Act in 1996, even more restrictions were
“groundfish” such as flounder and haddock, and became so imposed. But could an exploited stock recover, or was such
serious that government and private industry began to coop- effort too little too late? Remarkably, stocks of demersal
erate more effectively than ever before to save the fishery. species showed rapid recovery in this system after such
Changes resulting from that cooperation were sweeping. measures were implemented (Fig. 8.30), and stocks of
Four areas containing over 5000 nautical miles were closed pelagic (free-swimming, upper layer) species such as herring
to vessels with fishing gear capable of catching groundfish. and mackerel also increased after reductions in fishing effort
New regulations increased minimum net mesh size (allowing and TAC were imposed (Sherman et al. 2003).
younger individuals to escape). A moratorium was placed on The effects of over-exploitation on targeted commercial
new vessel entrants (i.e., no new fishing vessels were allowed species are not surprising, but effects on non-target species
to enter the industry). Vessel and dealer reporting of catches can be equally devastating. Removal of prey species can
Fig. 8.30 Recovery of yellowtail

flounder (a) and haddock (b) from
the Georges Banks/US Northeast
Shelf Ecosystem following
reductions in total allowable catch
and gear restrictions. Note that
stock biomass and recruitment in
both species begin to recover
almost immediately after
reduction in exploitation rate.
Elsevier, from Hempel, G., and
Sherman, K., 2003. Large Marine
Ecosystems of the World: Trends
in Exploitation, Protection, and
Research. Large Marine
Ecosystems, V. 12. Amsterdam:
Elsevier. # 2003 Elsevier B.V.)
reduce populations of predator species, and not of fish only, fishing in the sea lion’s critical foraging habitat violated the
but also of birds and mammals. Examples have been seen in ESA’s directive that “reasonable and prudent alternative
the decline of Peruvian seabirds following decimation of the (RPA) measures” be taken to avoid inflicting “adverse modi-
anchovy fishery and in the reduction in numbers of sea otters fication” on critical habitat of an endangered species. A US
off the California (USA) coast following over-fishing of district court upheld the decision and ordered the National
abalones (Agardy 1997). Marine Fisheries Service to revise its RPA (Stump 2000).
Some cases have had legal as well as biological As the removal of a prey species can cause declines in the
ramifications. In 1998, a coalition of US environmental predator, so the removal of a predator can cause changes in
organizations sued the North Pacific Fishery Management prey populations, and those changes do not always lead to
Council under the US Endangered Species Act for failing to uniform or long-term increases. Over-exploitation disrupts
protect foraging habitat for the Steller sea lion (Eumetopias equilibria of many populations (Agardy 1997) and can
jubatus) by allowing unregulated pollock fishing in the sea make them more susceptible to declines associated with
lion’s foraging areas (Stump 2000). Lack of food had previ- environmental and demographic stochasticity (Chap. 6),
ously been identified as a cause of decline in sea lion such that stocks may continue to decline even after take is
populations, and pollock is an important prey of sea lions. restricted or stopped altogether (Lauck et al. 1998). The
The plaintiffs argued that to allow unregulated commercial exploitation of great whales provides a singular example.
8.7.2 The Surplus-Yield Theory: Great Whales harvest. Large baleen whales feed primarily on krill (small,
and Ecological Function shrimp-like invertebrates), rather than commercial species of
fish. In commercial fishing, some krill is harvested directly,
The “great whales” (thirteen larger species of toothed and especially in the production of krill oil, but the primary value
baleen whales) have long been exploited by humans. of krill lies in its role in the food base for larger fish. If, as the
Although whale populations had been reduced by North surplus yield reasoning goes, whales decimate krill and lower
American and European whaling throughout the eighteenth primary production of the marine system, commercial
and nineteenth centuries, the so-called “industrial whaling” whaling should resume on an international scale in order to
period, beginning in the early twentieth century, brought prevent such losses and make more of this production (more
about even more precipitous declines and near extinction of krill) available to commercially targeted fish species.
many species. Australian marine scientists Trish Lavery and her
Today, as a result of a moratorium on harvesting of most colleagues evaluated assumptions of the surplus-yield
species of great whales under most circumstances, whale model from a comprehensive ecological standpoint, and
stocks are showing signs of recovery, a recovery coincident determined that, in its simplistic form, the surplus-yield
with the collapse of most of the world’s major commercial model ignores the role that large baleen whales play in the
marine fisheries. The increase in baleen whales has been primary production processes of marine systems, most
particularly noticeable. For example, the blue whale directly through their excretions of iron-rich feces into iron-
(Balaenoptera musculus) (Fig. 8.31), the largest creature deficient marine waters. Whale feces contain iron
ever to have lived on earth, saw its populations decline concentrations seven orders of magnitude greater than
from an estimated 239,000 individuals in the Southern surrounding marine seawater, thus representing a potentially
Ocean (Antarctic, Indian, South Atlantic, and South Pacific large source of iron input. Lavery et al. determined that “Our
Oceans) in 1905 to around 1700 individuals by 1996. modified surplus-yield model indicates that an equivalent
Increasing from that point, as result of protection, at a rate amount of carbon (1.3 1011 kg C yr1) [to that lost from
of 7.3% annually, the same population was estimated to krill consumed by whales] would be stimulated via the action
contain over 5000 individuals by 2012 (Lavery et al. 2014). of blue whales defecating iron into surface waters, suggesting
As blue whale and other whale species have recovered, some that the recovery of blue whale stocks would increase the
voices in the commercial marine fishing community have marine productivity of the Southern Ocean by 1010 kg C
advanced a variation of the now discredited Maximum Sus- yr1” (Lavery et al. 2014:888–889) (Fig. 8.32).
tainable Yield theory in the form of the “the surplus-yield The findings of the Lavery et al. model help to explain
model,” asserting that, if recovery of the great whales records of exceptionally high biomass of both krill and
continues, they will consume vast quantities of marine whales which characterized the Southern Ocean system in
fisheries production that will then be lost to commercial the 1920s and 1930s when whale populations were much
larger, particularly in former whaling grounds in the South-
west Atlantic Sector where extensive records exist. This area,
the so-called Antarctic Peninsula Plume (APP), is today
classified as only moderately productive. But, as Lavery
et al. point out, “If whales stimulate primary productivity
with their iron-rich feces, and if no other predator has
expanded to fill the trophic void left by depleted whale
stocks, one can expect that primary production will have
decreased in proportion to whale populations. Indeed,
declines in satellite-measured surface chlorophyll
concentrations since the 1970s indicate declining levels of
marine production in the APP. . .and there have been reports
of an 80% decline in APP krill stocks since the Discovery
surveys of the 1920s. . .” (Lavery et al. 2014: 900). Informed
by the results of their model and careful study of historical
records, Lavery et al. concluded “The recovery of Southern
Fig. 8.31 Populations of blue whale (Balaenoptera musculus), the Ocean blue whales to their historical abundances and popu-
largest creatures ever to have lived on Earth, were decimated during lation structures dominated by large adult animals is thus
industrial whaling throughout most of the twentieth century. As num-
bers recover, there is growing evidence that blue whale and other whale
unlikely to reduce fishery yields and may in fact enhance
species provide vital supplies of iron to marine ecosystems through their ecosystem productivity in the Southern Ocean” (Lavery et al.
iron-rich feces. (Photo courtesy of NOAA, public domain) 2014: 902).
Fig. 8.32 The influence of the projected recovery of blue whales to defecation of iron-rich feces into surface waters by blue whales.
their historical abundances on the net carbon availability in the Southern (Reprinted by permission from Wiley, from Lavery, T.J., Roudnew,
Ocean. The traditional surplus-yield model assumed that all primary B., Seymour, J., Mitchell, J.G., Smetacek, V., Nicol, S., 2014. Whales
production required to support krill consumption by whales was ren- sustain fisheries: Blue whales stimulate primary production in the South-
dered unavailable to fisheries. The modified surplus-yield model of ern Ocean. Marine Mammal Science 30, 888–904. # 2014 Society for
Lavery et al. (2014) integrates both primary production required to Marine Mammalogy)
support krill consumption and primary production increased by
8.7.3 Bycatch: The Preeminent Threat to Large

Marine Vertebrates
Threats posed to marine ecosystems do not arise merely from

overexploitation of targeted commercial species, but also
from mortality of non-target species which suffer as a result
of commercial fishing operations. In some fisheries, such as
shrimp, worldwide discarded biomass of non-targeted spe-
cies exceeds the targeted catch (Agardy 1997)! By far the
most dangerous and pervasive threat of all to larger marine
vertebrates occurs “accidentally,” an insidious form of “unin-
tended” mortality known as bycatch.
“Bycatch” can be defined as the inadvertent capture of
animals that have become hooked, trapped, or entangled in
fishing gear deployed with the intention of catching some-
thing else (Reeves et al. 2013) (Fig. 8.33). Many species are Fig. 8.33 Sea turtles killed by becoming entangled in fishing gear
affected, but large marine vertebrates such as sea turtles, intended to catch marine fish species. Bycatch mortality may be imme-
dolphins, sharks, rays, as well as both vertebrate and inverte- diate, as when a non-target species is “landed” onto a ship along with
brate benthic organisms, continue to be killed in large num- target species, or delayed because the turtle or other non-target species
escapes still carrying portions of fishing gear such nets, hooks, lines, or
bers as bycatch species. The problem is magnified because floatation devices. Delayed bycatch often inflicts prolonged periods of
mortality is multi-dimensional. Some bycatch species are pain, stress, and suffering before death occurs. Bycatch, in all its forms,
killed by fishers when they are retrieved with the targeted is considered the single largest threat to populations of large marine
catch. But there is also the problem of “cryptic” bycatch, vertebrates. (Photo courtesy of Joanna Alfaro-Shigueto)
animals that become entangled in fishing gear, escape, then

swim away injured, sometimes with gear still attached, only
to die later, even though they are not ‘caught’ or accounted
for in harvest statistics.
Bycatch mortality can be inflicted not only during active
fishing periods, but also when animals are “caught” in fishing
gear that has been lost or abandoned (“ghost” bycatch), a
form of mortality known to be significant but almost entirely
unquantified. Bycatch of marine mammals in gill nets alone
is estimated to kill between 300,000 and 600,000 individuals
annually (Simmonds et al. 2016, Taylor et al. 2016).
Although bycatch is most commonly associated with gill
netting, it also occurs when fishers use other techniques
including purse seine netting, longlines, trawling, traps and
drift netting. The last technique, drift netting, has been illegal Fig. 8.34 The vaquita, (Phocoena sinus), a species of dolphin endemic
since 1993, but is still practiced worldwide, and considered to the Gulf of California (Mexico), is currently considered the world’s
most endangered marine mammal. It is not threatened by commercial
by many marine conservationists to be the deadliest tech- exploitation, but solely through unintended captures in gillnets.
nique of all. Bycatch not only occurs during commercial (Photo courtesy of Alejandro Robles)
fishing operations, but individual fishers, as well as family
groups or communities practicing artisanal fishing also inflict species of dolphin, including the Mãui dolphin
heavy mortality. (Cephalorhychus hectori maui) of the Hawaiian Islands in
Lewison et al. have provided one of the most comprehen- the Pacific, the Irrawaddy dolphin (Orcaella brevirostris) of
sive recent assessments of bycatch and its impacts on a the Mekong River in southeast Asia, and the harbor porpoise
worldwide scale. They noted that “declines in marine mega- (Phocoena phocoena) in the Proper Baltic Sea (Simmonds
fauna lead to major changes in ecosystem function and et al. 2016).
process. . .This loss of megafauna, referred to as trophic Many of the worst effects of bycatch mortality are not
downgrading, has reverberating effects on biotic interactions, immediate. As Simmonds et al. reported to the International
disturbance regimes, species invasions, and nutrient Whaling Commission, “Recent investigations documenting
cycling. . . (Lewison et al. 2014: 5271–5272). Examining these scars [from encounters with fishing gear] on
data for marine megafauna from 1990 to 2011, Randall endangered North Atlantic right whales (Eubalaena
Reeves and his colleagues determined that 75% of toothed glacialis) and humpback whales (Megaptera novaeangliae)
whale species (odontocetes), 64% of baleen whale species along the East Coast of the US indicate that they regularly
(mysticetes), 66% of pinniped (“flippered”) mammals, all encounter and shed gear and that many more have been
species of sirenians (manatees and dugongs) and all species affected than have been reported entangled [and] between
of marine mustelids (sea otters) had been captured in gillnets 65–85% of the individuals within these populations bear
during this period (Reeves et al. 2013). scars from having encountered gear and. . . 12–16% of these
Bycatch can lead to the extermination of a species even populations exhibit new scars each year” (Simmonds et al.
when the species has been completely protected from com- 2016). Larger whales may be strong enough to break free of
mercial harvesting. The recent extinction of the beiji or the gear, but, even if the whale escapes, it often remains
Yangtze dolphin (Lipotes vexillifer) in China occurred solely partially entangled in the gear it has torn away. Remaining
as a result of bycatch mortality. The critically endangered rope and gear may gradually constrict one of more body
vaquita porpoise (Phocoena sinus) (Fig. 8.34), endemic to parts, or may wind through and around the whale’s mouth,
the Gulf of California (Mexico), has experienced population hindering its ability to feed. Such events are especially com-
declines of 34% per year even though it was never an mon in North Atlantic right whales, where time to death after
intended commercial target. The Mexican government placed entanglement averages 6 months (Moore and van der Hoop
a two-year ban on all gillnetting in areas used by vaquitas in 2012, Simmonds et al. 2016). In such cases, the individual
2015, but illegal gill netting has continued (Taylor et al. may experience impaired foraging, increased drag that
2016). Now considered the world’s most endangered ceta- impedes its ability to swim, infection associated with gear-
cean (Reeves et al. 2013), the vaquita is almost certain to inflicted wounds, internal hemorrhaging and tissue damage.
become extinct unless the current ban on gillnetting is made The description may be graphic, but not inaccurate, and
permanent and stringently enforced. But the vaquita, though underscores the fact that bycatch is not only the number one
in the most imminent danger of loss, is not an exceptional threat to marine megafauna conservation, but also to the
case. Bycatch is a serious threat to many other endemic health and physical welfare of large marine mammals.
8.8 Preserving Marine Habitats and Biodiversity through Protected Areas: The Marine Reserve 349
8.7.4 Reducing and Mitigating Effects Union’s (EU) Common Fishery Policy (CFP), which
of Bycatch includes provision for estimating and reducing the total inci-
dence of non-target species bycatch. Since 1992, the EU’s
There are many techniques and management strategies Habitats Directive has required bycatch monitoring by EU
which, employed and enforced, would reduce bycatch- Member States (Peltier et al. 2016). But for all the good
related mortality. Time and area closures during periods of intended by these directives, they are limited by observer
marine vertebrate use and activity, particularly of marine bias. One aspect of this bias is the “deployment effect,”
mammals, reduce the probability of an encounter occurring because accepting an observer on board a fishing vessel is a
between nontarget species and fishing equipment. Reducing matter of the vessel master’s discretion. A second part of
the amount of fishing effort, especially the allowable number observer bias is the “behavior effect,” because fishers may
of fishing vessels in an area or the amount of gear that is change their fishing practices to reduce bycatch when an
allowed to be set by each vessel would also reduce bycatch observer is present (Peltier et al. 2016).
mortality. Gear modification and use can reduce encounters
between marine vertebrates and fishing gear, or the harm that
occurs when there is an encounter. Such modification can
Recently the World Wildlife Fund, in cooperation with
include changing the configuration or amount of ropes used
the international fishing industry, has sponsored The
in the fishing effort, including reducing the length of vertical
International Smart Gear Competition to provide incen-
lines of rope in the water. When ropes or other netting
tive to commercial fishers to reduce bycatch. Fishers
material is in the water, acoustic deterrent devices (“pingers”)
submit ideas on new technologies to reduce incidental
can be attached to the rope or netting, sending an acoustic
capture of non-target species for relatively modest cash
signal that many marine mammals, especially cetaceans, can
prizes. The 2011 winner, for example, was a Japanese
recognize and interpret as something to avoid. Turtle
fisher whose invention reduced seabird bycatch by
excluder devices in trawl fisheries, especially in shrimping
nearly 90% (Aslan et al. 2013). For the fishing indus-
operations, have been implemented in Australian fisheries
try, paying the prize money is cheaper than paying the
and reduced bycatch of sea turtles by 90% (Lewison et al.
penalties associated with bycatch regulations, and gear-
2014). Similarly, a variety of seabird-specific mitigation
driven reduced bycatch has a much greater and more
devices has reduced bycatch of albatross and petrels in long-
lasting effect. How could a government regulatory
line vessels in Hawaii, Alaska, and the Southern Ocean
agency incorporate ideas from the Smart Gear Compe-
(Lewison et al. 2014). Fishing gear itself can be modified to
tition to increase enforcement success to reduce
reduce harmful consequences if contact occurs, such as using
bycatch while at the same time reducing its expenses?
ropes and nets with built in “weak links” or that have lower
breaking strengths (sufficient to hold the target fish species,
but easily broken by a large marine mammal) (Simmonds
et al. 2016).
For all the good that can be done with improved technol-
ogy and better fishing practices, bycatch will remain a threat 8.8 Preserving Marine Habitats
to marine megafauna as long as catch reduction measures are and Biodiversity through Protected
not required or enforced by individual nations and regional Areas: The Marine Reserve
fisheries management agencies, most of which still do not do
so. And because commercial marine fishing usually occurs in 8.8.1 Management Context, Goals
international waters, the problem of bycatch will require and Strategies in Marine Reserves
international cooperation and regulation if there is to be an
effective solution. At the present time, even the reporting of Historically, marine conservation efforts have used three
bycatch and bycatch mortality is sporadic. As Simmonds basic approaches, alone or in combination: (1) creating
et al. propose “. . .one of the areas most needed to address area-specific management plans that direct conservation
this issue globally is a system through which successful efforts for targeted areas or species; (2) establishing a marine
mitigation measures can be made available to a global audi- reserve in which “taking” activities are excluded, or
ence” (Simmonds et al. 2016). (3) establishing marine protected areas (MPAs). MPAs usu-
To solve the problem of bycatch mortality, what is needed, ally have less restrictive regulations than marine reserves,
at international levels, is the development and adoption of although they are sometimes nested within them. MPAs
participatory agreements, adaptive approaches, and serious usually restrict allowable catch, season of catch, catch effort,
enforcement procedures to reduce the bycatch of marine catchable species, or type of fishing effort and gear permitted
megafauna. One example of such an effort is the European for use.
Three goals that marine reserves and MPAs are designed least twice the median dispersal distance for an isolated
to meet simultaneously include: (1) protecting marine and marine protection area to sustain viable populations. As
coastal biodiversity, (2) improving productivity of local more marine protected areas are included in a network
fisheries, or at least ensuring that marine productivity is not design, the minimum size decreases because other protected
undermined by uncontrolled exploitation, and (3) focusing locations are within dispersal distances to provide propagules
efforts for restoration of vital areas that may be presently to sustain the local population.
degraded but have potential to support healthy marine Efforts to establish marine reserves have varied in effec-
ecosystems in the future (Agardy 1997). To these ends, tiveness according to region and country. Within a variety of
marine reserves have been established worldwide with a management goals, strategies and national efforts, we can
variety of names, jurisdictions, and purposes. Within examine specific cases of individual marine reserves and
reserves, areas closed to all types of marine fishing and their management approaches to better understand their role
harvesting are often designated as harvest refugia or no-take in conservation.
zones. These are designed to protect a particular commercial
stock or group of stocks from over-exploitation. At large
scales, some “biosphere reserves,” administered by the 8.8.2 Protection at Ecosystem Levels:
United Nations’ Educational, Scientific, and Cultural Orga- Australia’s Great Barrier Reef
nization (UNESCO), include marine reserves, like the Marine Park
Palawan Biosphere Reserve in the Philippines, which
includes not only the island of Palawan and smaller islands One of the best examples of a large marine reserve managed
near it, but also surrounding coral reefs, seagrass meadows, comprehensively as a functional ecosystem is Australia’s
and coastal mangrove forests (UNESCO 2013). Marine Great Barrier Reef Marine Park (GBRMP) (Fig. 8.35), one
portions of many such reserves may be divided into three
zones, much like the Multiple-Use Modules (MUMS)
discussed earlier in this chapter relative to protected areas
for freshwater systems (Section 8.3.1). Like a MUM, there
are usually “core” reserves with little or no harvesting
surrounded by “buffer” areas where limited harvest and
other activities are permitted. Around the buffers are “transi-
tion” areas that are minimally protected and administered
with regulations most like those outside the reserve (Sobel
1993). At small scales, more limited reserves may be
established to achieve a more restricted set of conservation
objectives.
As in terrestrial conservation reserves, size is a key con-
sideration for marine reserves. A protected area must be large
enough to retain a large proportion of marine organisms
within its boundaries, but migratory marine mammals, fishes,
and invertebrates require large marine protected areas. Their
high rates of offshore, seasonal, and ocean-wide migrations
mean that substantial portions of their lives might be spent
outside protected areas that are too small to contain their
movements. Many marine organisms that are sedentary as
adults, such as sponges and tunicates, have larval stages with
enormous dispersal capacities, often passively drifting long
distances in ocean currents. Therefore, patterns of per area
larval accumulation and retention should be compared
between different potential areas being considered for reserve
status because both behavior and oceanographic processes
affect dispersal distance. As a consequence, even relatively
small spatial errors in the placement of a marine reserve or Fig. 8.35 Australia’s Great Barrier Reef is a colorful and beautiful
network of marine reserves can mean the difference between example of a marine area protected for conservation of marine biodiver-
sity but managed under multiple-use and multiple-zonation approaches
successful dispersal and gene flow for many species of plank- that permit different activities in different portions of the reef system.
tonic invertebrates and a state of isolation and eventual (Photo courtesy of C. Jones # Commonwealth of Australia
extinction. Theoretical models suggest a minimum size of at (GBRMPA))
of the world’s premier marine protected areas, and part of the co-management. Co-management is a conservation strategy
Biosphere Reserve and World Heritage Site programs. The that attempts to simultaneously address biological, cultural,
Great Barrier Reef itself is a vast complex of some 2,900 economic, and political concerns through collaboration and
individual reefs and 250 cays (low islands or reefs made of integration in conservation efforts between local
sand or coral) stretching along the continental shelf of north- communities and government authorities. In a typical
east Australia from just south of the Tropic of Capricorn to co-management system, government agency personnel,
the Torres Strait. The system possesses 71 genera of coral such as park administrators, share responsibility and
alone. The Great Barrier Reef was relatively inaccessible to decision-making power with local residents, usually through
humans until the 1960s. The GBRMP that attempts to pre- the mechanism of a “council” or equivalent structure which
serve it is in many ways exceptional among marine preserves. includes representatives of the agency and local stakeholders
The preserve was not established to stop or solve an existing who represent various interest groups. In addition to partici-
problem or degradation of the reef but formed in anticipation pation in the council and its decision-making functions, local
of future problems that were just beginning to emerge. In residents also perform services in education, research, eco-
1967, a private Australian firm filed an application for per- logical monitoring, or law enforcement within the park. In so
mission to take coral limestone from a part of the reef for use doing, managerial expertise that is typically the domain of
in the production of agricultural lime. The Wildlife Preserva- government employees is complemented by traditional
tion Society (WPS) of Australia perceived this application as knowledge of the area and its natural community possessed
the first step in setting a precedent for dangerous and destruc- by local citizens. But could such a system work in a real
tive processes that could eventually destroy the reef. With marine reserve, and what would it look like if it did?
other conservation groups joining the lead of the WPS, public The ecosystems of the Comoros Islands in the West Indian
outcry led to the refusal of the permit application by the Ocean, a biodiversity hotspot with high endemism and
provincial government (Queensland). Further controversies diverse tropical marine environments, are adversely affected
over offshore oil drilling in the reef area and outbreaks of the by existing ecological, socioeconomic, and political
crown-of-thorns starfish (which destroys reef corals) led to conditions. Most local residents are poor. Many make a
legislation that established the GBRMP (Kenchington 1990). living in whole or in part by harvesting marine resources
Today the GBMRP is a vast multiple-use area managed by from the system, including the system “protected” within
establishing different zones within the park for different uses, the park. The government of the Comoros, like its citizens,
through which it has successfully accommodated a variety of is not wealthy, and its resources to enforce park protection are
user groups (Fig. 8.36). Much of its success is attributable to limited. So are its educational and professional expertise,
careful attention to the criteria earlier described for successful providing it with only limited scientific knowledge of the
marine reserves. Rigorous scientific study has helped define park’s marine systems and species. Despite these constraints,
park boundaries and management policies, but the key to its the Comoros Government, with help from IUCN and funding
success has been widespread stakeholder involvement from the World Bank’s Global Environmental Facility (GEF)
through an extensive participatory network. Participants in and the United Nations Development Programme (UNDP),
this network have incorporated not only scientific informa- established Mohéli Marine Park in the Federal Islamic
tion, but socioeconomic, political, and cultural objectives into Republic of the Comoros. The goals of the park were to
the park’s management, building a broad base of support for stop loss of biodiversity in the Comoros while at the same
its conservation objectives. However, even in this exemplary time developing increased local capacity for and participation
park there are problems. For all its size and jurisdictional in natural resource management. The Mohéli Marine Park
power, the GBRMP Authority that manages the park has no contains both core reserve (no take) areas as well as protected
control over land-based inputs that pose threats to its coral areas within the park which permit marine harvests according
reefs, commercial fish stocks, and endangered species. Its to prescribed methods and limits. The park contains a diverse
jurisdiction stops at the shoreline. It is powerless to stop environment of coastal mangroves, seagrass beds, inshore
influxes of sediments and chemical pollutants that pour into and offshore coral reefs, and its beach areas include some
system it is committed to protect (Agardy 1997). of the world’s best nesting beaches for green sea turtles
(Chelonia mydas) (Fig. 8.37), where up to 5,000 females
come ashore each year to build nests and lay their eggs
8.8.3 Co-management – Can Shared Authority (Granek and Brown 2005).
Provide Better Conservation? Faced with the daunting task of protecting and managing
the park with limited resources and capacities, the Comoros
The GBRMP is managed at a national level by the Australian government, with the help of IUCN, GEF, and UNDP, devel-
government. One alternative to the traditional government oped a co-management strategy requiring participation from
model of establishing and protecting marine reserves is both government and local community organizations. At the
Fig. 8.36 Management zones in the Capricornia section of the Great Barrier Reef Marine Park, Australia. (Photo courtesy of the Great Barrier Reef
Marine Park Authority # Commonwealth of Australia (GBRMPA))
political structures in the government that would have

excluded them. It inspired local interest in tracking the
park’s success, leading to greater participation in monitoring
park resources. In recognition of these achievements, Mohéli
Park received the UNDP Equator Initiative Award, presented
at the Word Summit on Sustainable Development in 2002,
with a prize of US$30,000 to further develop the park’s
programs and infrastructure (Granek and Brown 2005).
The Mohéli program was, by many standards, a success,
but it also had shortcomings. Park design was based on
limited scientific data which impeded ability to accomplish
conservation goals. Despite admirable performance by the
ecoguards, inadequate governmental resources for policing
the park worked against preserving its biodiversity. Many
violators were not arrested, and many who were arrested were
Fig. 8.37 The green sea turtle (Chelonia mydas), a species that has released by government officials with no more than a warn-
benefitted from conservation management in the Mohéli Marine Park. ing. And despite commendable local initiative, regional and
(Photo courtesy of D. McLeish) global issues such as overpopulation and climate change
could destroy achievements produced by local effort. As
local level, this required participation from village environ- conservation policy experts Elise Granek and Mark Brown
mental organizations as well as local village fishers’ note, “Co-management is not a panacea for incomplete sci-
associations, which included individuals with high levels of entific data and broader economic and political problems, but
concern for the state of biotic resources in the park, resources it offers a potential alternative in situations faced with limited
which had traditionally supported their livelihood. Overall resources for addressing biodiversity conservation and natu-
program goals were to protect Comorian biodiversity and ral resource loss.” (Granek and Brown 2005:1731).
improve local and regional fisheries. The plan required rep-
resentation of the local community in the process of conser-
vation management, scientific monitoring, and 8.8.4 Marine Protected Areas and Commercial
environmental education. To accomplish this, village Fisheries
residents nominated representatives to serve as “ecoguards”
who would be responsible for monitoring local marine Some fisheries scientists and conservationists have argued
resources, enforcing park regulations, and representing the that the answer to the problem of collapsing commercial
interests of the community to the government. Ecoguards fisheries may be the establishment of marine reserves
were selected based on their perceived commitment to con- intended to sustain them. They assert that marine reserves
servation, respect and trust of village members, ability to help should become the foundation of a new form of fisheries
foster community participation in park programs, and confi- management based on a radical change in perspective.
dence of fellow villagers in their ability to patrol their zone of Namely, that we should abandon the concept that every
the park, enforce park regulations, and educate park visitors available commercial fish stock should be exploited opti-
(Granek and Brown 2005). mally and replace it with the strategy of “bet hedging.” That
Ecoguards improved communication and cooperation is, one should assume that high levels of uncertainty are a
among local villages that had a long history of conflict with permanent and persistent dimension of estimating the size of
one another over resource use. They developed novel and fish populations and their future trends. If high uncertainty is
widely embraced educational programs for local school chil- taken as a given, the best strategy is not to attempt to harvest a
dren, including a special environmental magazine, Mwana population optimally wherever it occurs, but to harvest some
wa Nyamba (The Baby Turtle) and a special “field day,” of the populations at the predicted (but uncertain) optimal
Journée de la Tortue (“Day of the Turtle”), that permitted level and leave a large portion unharvested as a protection
school children to observe female turtles making nests and against unforeseen (and uncontrollable) declines in the
laying eggs on local beaches. Over several years, Mohéli harvested stock.
integrated education, local knowledge, and community com- Lauck et al. manifested their ideas in a model whose goal
mitment into a viable conservation program that overcame was to retain fish populations at more than 60% of carrying
limited financial resources, weak government enforcement capacity for at least 40 years. Through a series of equations
capacities, and limited scientific data. The co-management that permitted estimation of the proportion of the population
approach empowered community leaders and circumvented available for harvest outside a closed area, Lauck et al.
estimated the probability that the population could persist for 1998). Thus, Lauck et al. reach the radical but rational con-
specified periods and levels. They assumed that half of the clusion that marine protected areas provide the “best
available population outside the reserve was captured annu- approach to implementing the precautionary principle and
ally, but with coefficients of variation (CV, the measure of achieving sustainability in marine fisheries” (Lauck et al.
uncertainty about the mean) assigned at six different levels 1998).
from 18–61%, and they varied the fraction of total area Empirical data from marine reserves complement the the-
available for harvesting (Lauck et al. 1998). oretical work of Lauck et al. and support their value in
The effect of catch variation and variation in exploitation restoring fish populations. For example, Russ and Alcala
rates had dramatic effects in this model. Even with a moder- (1996) compared density and biomass of large predatory
ate amount of variation in the catch (CV < 50%), the proba- fish at two small marine reserves in the Philippines with
bility of the population persisting for 40 years dropped two similar control sites. They found that the longer the
drastically when the amount of exploitable area became reserve was protected from fishing, the greater the increase
greater than 30% of the reserve (Fig. 8.38). If catch percent- in density and biomass of large predatory fish (Fig. 8.39). But
age was more variable, probability of the population’s persis-
tence was less than one (not certain) even if only 5% of the 20
area was harvested. The probability of successfully
y = 6.87 + 1.15x; r 2 = 0.97
protecting the fish stock increased if the harvest was reduced
to lower levels, and, at lower levels, if more area was made
available to fishing. Two conclusions emerged from the 16
Large Predators (no./1,000 m2)

model. First, “a reserve can simultaneously lead to stock
protection and a higher level of catch,” and “it is possible to
20
Mean Density of
maximize catch while protecting the stock” (Lauck et al. 12
Large Predators (kg/1,000 m2)

16
Mean Biomass of
8 12
y = 2.58e0.18x; r 2 = 0.89
8
4 Sumilon
Reserve 4
0 0
–3 –2 –1 0 1 2 3 4 5 6 7 8 9 10 11
Years of Protection
12
Large Predators (no./1,000 m2)
10
Large Predators (kg/1,000 m2)

8 y = 0.56 + 0.72x; r 2 = 0.95 Mean Biomass of
Mean Density of
Apo
4 Reserve 4
0.16x 2
y = 1.61e ; r = 0.83
2
Fig. 8.38 The probability that a fish stock remains at a size greater than
0 0
60% of its carrying capacity for 20 years depends on the fraction of area –3 –2 –1 0 1 2 3 4 5 6 7 8 9 10 11 12
available for harvesting. When the total area for harvesting increases
beyond 30%, the probability for maintaining a population size that is Years of Protection
>0.6 K (K ¼ carrying capacity) drops rapidly. Each line represents a
different value for coefficient of variation associated with the average Fig. 8.39 Changes in density and biomass of large predatory fish at two
harvest (CV is defined as the standard deviation of the harvest fraction/ marine reserves in the Philippines with two similar control sites. The
mean of the harvest fraction). (Reprinted by permission from Wiley, longer the reserve was protected from fishing, the greater the increase in
from Lauck, T., Clark, C.W., Mangel, M., Munro, G.R., 1998. density and biomass of large predatory species (Reprinted by permission
Implementing the Precautionary Principle in Fisheries Management from McGraw-Hill Publishers, based on data from Russ, G.R., Alcala,
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Conservation Through Ecosystem Management
9
Only if we can comprehend and envision the entity we are trying to shape as a dynamic whole can we have any
hope of dealing with it creatively.
J. T. Lyle (1985)
Keywords ecosystem management came in June 1992. At that time,

Ecosystem-based management · Integrated ecosystem Dale Robertson, Chief of the US Forest Service, announced
assessment · Northern spotted owl · Adaptive manage- that his agency would be moving to an “ecosystem approach”
ment · Performance-based evaluation · Protected area- in their management of national forests; making the Forest
centered ecosystems · Ecosystem monitoring · Ecosystem Service the first environmental agency in the US to adopt
modeling · Ecosystem-based governance · Ecosystem ecosystem management as its official agency perspective
processes (Butler and Koontz 2005). Within 4 years, seventeen other
US federal agencies had adopted or were considering adop-
tion of ecosystem management programs (Christensen et al.
1996), producing a wealth of agency-specific definitions of
Overview what ecosystem management was (Table 9.1).
In this chapter, you will learn: The Forest Service, like other federal US environmental
agencies, had not developed in a culture of ecosystem man-
1. What is ecosystem management. agement, but of resource management (Chap 1). Federal land
2. How and why the concept of ecosystem manage- agencies have always had jurisdiction over ecosystems but
ment developed. did not historically manage their jurisdictions as ecosystems.
3. The scientific basis of ecosystem management. Rather, they managed ecosystems for the production of
4. Methods of implementing ecosystem management resources. The distinction between resource management
and what they can accomplish. and ecosystem management can be most clearly understood
when parallel elements in the two approaches are directly
compared (Table 9.2). Governed by a paradigm of resource
management, the entity of value is a particular “resource,”
either an individual species or an abiotic component of the
system such as water, soil, or minerals. The resource is seen
as a commodity and its value is “use.” Units of management
9.1 The Concept of Ecosystem Management are species or abiotic factors and sites on which they occur. In
this approach, single species are often managed on a site-
9.1.1 Resource Management and Ecosystem specific basis, usually through direct intervention. Time
Management scales are short and management decisions occur in individ-
ual agencies. The management goal is production of multiple
Ecosystem management is still a relatively new concept. natural resources as commodities for multiple uses.
Birthdays of concepts are hard to identify, but one of the Objectives are determined by demand for commodities that
earliest efforts to not only define but begin the practice of the system can supply.

360 9 Conservation Through Ecosystem Management
Table 9.1 Some definitions of ecosystem management in various US federal agencies

Department Specific agency Definition
Department of The integration of ecological principles and social factors to manage ecosystems to
Agriculture safeguard ecological sustainability, biodiversity, and productivity.
Department of National Oceanic and Activities that seek to restore and maintain the health, integrity, and function of natural
Commerce Atmospheric Administration ecosystems that are the cornerstone of productive, sustainable economies.
Department of The identification of target areas, including Department of Defense lands, and the
Defense implementation of a “holistic approach” instead of a “species-by-species approach” to
enhance biodiversity.
Department of A consensual process based on the best available science that specifically includes human
Energy interactions and management and uses natural instead of political boundaries to restore and
enhance environmental quality.
Department of Bureau of Land Management The integration of ecological, economic, and social principles to manage biological and
the Interior physical systems in a manner safeguarding the long-term ecological sustainability, natural
diversity, and productivity of the landscape.
Fish and Wildlife Service Protection or restoration of the function, structure, and species composition of an
ecosystem, recognizing that all components are interrelated.
National Park Service A philosophical approach that respects all living things and seeks to sustain natural
processes and the dignity of all species and to ensure that common interests flourish.
US Geological Society Ecosystem management to emphasize natural boundaries, such as watersheds, biological
communities, and physiographic provinces, and bases management decisions on an
integrated scientific understanding of the entire ecosystem.
N/A, Environmental Protection To maintain overall ecological integrity of the environment while ensuring that ecosystem
Independent Agency outputs meet human needs on a sustainable level.
N/A, National Science Foundation An integrative approach to the maintenance of land and water resources as functional
Independent habitat for an array of organisms and the provision of goods and services to society.
Design by M. J. Bigelow based on data from USCRS (1994)
Table 9.2 Fundamental differences between resource management and ecosystem management paradigms in entity of value, value application,
management units, time scales, jurisdiction, decision making, and management goals. Design by M. J. Bigelow
Resource management Ecosystem management
Entity of value Resource Ecosystem
Application of value Beneficial use Continuing function
Management unit Species or abiotic factor Landscape elements
Time scale Relatively short Relatively long
Management jurisdiction Single government agency Multiple government agencies and private landowners
Management decision Single government agency Multiple government agencies and private stakeholders
making
Management goals Production and use of resource Productivity and sustainability of ecosystem functions and
commodities processes
This view of resource management was compatible with transmitted to successively lower organizational levels
traditional bureaucratic organization of most resource man- (Meidinger 1997).
agement agencies (Fig. 9.1a). Organizations like the Forest Beginning in the 1970s, the resource management para-
Service were arranged in a hierarchical system of supervisor- digm and its attendant bureaucratic structures began to have
subordinate control and characterized by departmental orga- increasing difficulty in dealing with conservation problems
nization in which each department had defined jurisdictions created by new environmental legislation, such as the US
and detailed rules governing roles of agency officials in each National Environmental Policy Act (NEPA) and the US
department, their operating procedures, and their boundaries Endangered Species Act (ESA), producing anomalies that
of decision making. Policies were set by a “head” account- helped prepare a climate of acceptance for an alternative
able to politically appointed or elected officials. Departments approach to management. For example, the ESA created an
were staffed by salaried officials progressing through fixed entity, threatened and endangered species, that could not be
career lines, which limited communication, understanding, managed in traditional ways. As the number of these species
and common interests between employees following differ- increased, species preservation on a case-by-case basis
ent career paths. Goals were determined at the top and then became too expensive to implement and too ineffective in
9.1 The Concept of Ecosystem Management 361
Fig. 9.1 (a) Schematic organizational representation of a classical areas are shared and fluid according to needs associated with particular
bureaucracy in a traditional resource management agency, in which a project tasks, an organization conducive to ecosystem management
Head (H) determines policies transmitted to lower level employees approaches. (Reprinted by permission from Island Press, from Kohm,
organized in departments with separate functions and jurisdictions. (b) K.A., Franklin, J.F. (Eds.), 1997. Creating a forestry for the 21st
Schematic organizational representation of a “project organization” century: the science of ecosystem management. Island Press,
model in an agency in which persons and knowledge in different agency Washington, DC)
its outcomes to meet goals of population viability. Manage- and impacts exceeded agency jurisdictional powers and
ment of endangered species increasingly forced agencies to boundaries. Further, the resource management paradigm
recognize their deficiencies in expertise needed for manage- was not able to deal with what could be called the “nationali-
ment decisions and compelled them to seek greater coopera- zation” of environmental values. With the emergence of
tion with non-agency scientists and citizens groups. national environmental legislation in countries throughout
Traditional bureaucratic organizations of agencies also the world also has come the rise of strong citizen-based,
were poorly equipped to deal with the horizontal geographically diverse constituencies pursuing ecosystem
information-sharing and citizen input that new environmental preservation for aesthetic and recreational values. Although
laws stimulated or with the policy adaptability needed for groups such as The Nature Conservancy, BirdLife Interna-
such interaction. Thus, government resource management tional, and The World Wildlife Fund had international
agencies eventually had to change their organizational organizations, their members were concerned with local
structures as well as their management models to become conditions. Agency bureaucracies could no longer justify
functional under these new conditions because their scientists policies that degraded local environments by invoking
and managers had to begin answering questions put to them national policy directives received from the top down
by citizens, lawyers, and legislators. To do this, environmen- or shrug, “We work for Congress and the President,” espe-
tal agencies had to begin moving from a bureaucratic organi- cially when new laws gave citizens ways to pursue their
zational model to what could be called a “project complaints and grievances with an agency, even to the extent
organization” model (Fig. 9.1b) that was more responsive to of lawsuit.
local and social concerns. In the project organization model, Changes that fostered development of ecosystem manage-
departments still existed, but individuals and expertise within ment were not primarily driven by new scientific information,
departments became interchangeable, driven by demands of but by social, legal, and political changes that altered public
individual management “projects” demanding interdisciplin- perception of the environment, as well as changes occurring
ary perspective. Further organizational changes occurred as in how government agencies were organized. As these
conservation dilemmas required expertise and cooperation changes continued, management actions and results, tradi-
beyond the agency itself, leading the agency to form more tionally viewed as cause and effect relationships, increased in
or less permanent relationships with other agencies and complexity and uncertainty, creating a greater need for rigor-
non-government entities, a so-called “project organization ous science in decision-making processes. These changes did
with external linkages” model (Meidinger 1997). These orga- more than redirect management goals. They altered the way
nizational adjustments facilitated integrative approaches fos- management decisions were made. To understand how these
tered by ecosystem management. developments affected actual practices, we return to the
The resource management paradigm also withered US Forest Service, and examine its interaction with a single
because its bureaucratic organization within individual species, the endangered northern spotted owl (Strix
agencies, agency departments, and localized districts could occidentalis caurina) and how this interaction drove this
not cope with modern environmental problems, such as air agency to adopt an ecosystem management approach.
and water pollution, toxic waste disposal, atmospheric depo-
sition, soil erosion, and stream sedimentation, whose sources
9.1.2 How the Spotted Owl Started Ecosystem recommended that management agencies retain 300 acres of
Management old-growth forest around every spotted owl nest site
(Caldwell et al. 1994). This recommendation was rejected
Until the late 1960s, little was known about the uncommon by the Forest Service and Bureau of Land Management
and rarely seen northern spotted owl (Fig. 9.2), which (BLM) because both wanted a statewide population manage-
inhabits the US Pacific Northwest. In the summer of 1967 ment goal established for spotted owls before implementing
Eric Forsman, an undergraduate student at Oregon State site-specific management practices (Meslow 1993). How-
University (OSU), learned that he could elicit the owl’s ever, when the ESA became law later in 1973, the spotted
response if it was present by imitating its call (Meslow owl was not listed as an endangered species, and its omission
1993). When Forsman and fellow undergraduate, Richard seemed to resolve the controversy. But conflict was just
Reynolds began to use this technique to search for spotted beginning.
owls in Oregon, they discovered that the owls could consis- In 1976, the newly-enacted National Forest Management
tently be found in old-growth forests, but rarely in other Act (NFMA) directed the Forest Service to “maintain viable
habitats. Forsman and Reynolds brought their data to OSU populations of existing native and desired non-native verte-
professor Howard Wight. Wight was interested, and by 1972, brate species” on national forests (Wilcove 1993). In other
Forsman had begun graduate research on the spotted owl words, the NFMA told the Forest Service it could not create
under Wight’s direction (Meslow 1993). any more endangered species, nor destroy portions of a
Forsman’s studies revealed a pattern of habitat use and species’ habitat (Meslow 1993). Under the guidelines of
population distribution in conflict with management policies NFMA, it was also clear that the initial recommendation of
of the Forest Service (Forsman et al. 1984). Managers viewed 300 acres for each pair of owls was inadequate because the
the owls’ preferred habitat of old-growth forests as areas of protection of such small areas would not maintain enough
low productivity with trees near the end of their life. Rational individuals to sustain the population (Wilcove 1993). The
management policy, which emphasized timber production, OESTF recommended a goal of maintaining 400 pairs of
was to cut old-growth timber. Timber management policy on spotted owls on public lands in Oregon, protecting sufficient
US public lands, however, operates within a context of envi- habitat that would provide for clusters of three to six pairs of
ronmental law, and two laws enacted during studies of the owls. Core areas for clustered pairs were to be no more than
spotted owl began to create conflicts with this traditional 1.6 km apart and each pair was to have a core area of
management approach. 300 acres. But subsequent studies of radio-tracked owls
In 1973, the US Fish and Wildlife Service (USFWS) (Forsman 1980, Forsman and Meslow 1985) demonstrated
included the spotted owl in its “Red Book,” an early version that individual owls needed at least 1000 acres of old-growth
of the US list of endangered species. At the same time, forest for permanent territories. In light of these findings, the
interest in conservation of old-growth forests was growing. plan was revised and recommendations changed to
After the owl’s Red Book listing, an appointed interagency 1000 acres of old-growth forest for each pair of owls within
group, the Oregon Endangered Species Task Force (OESTF), 1.5 miles of their nest site. However, the recommendations
were rejected by the BLM and only partially followed by the
Forest Service (Meslow 1993).
A population viability analysis by Russell Lande
concluded that the spotted owl population was declining
and could not be conserved unless significant portions of
the landscape remained in old-growth forests (Lande 1988;
Wilcove 1993). Based on this analysis, Lande determined
that the population could not persist with less than 20% of the
landscape in old-growth forests (Lande 1988), but the Forest
Service had proposed management guidelines that would
conserve only 6% (Wilcove 1993). Consequently, the Seattle
Audubon Society sued the Forest Service for failing to adopt
a conservation strategy in compliance with NFMA, eventu-
ally gaining an injunction against 135 timber sales in spotted
owl habitat (Caldwell et al. 1994). The USFWS was peti-
tioned again in 1987 to list the spotted owl as an endangered
Fig. 9.2 The northern spotted owl (Strix occidentalis caurina), a spe-
cies that can only be preserved with an ecosystem management species. The agency claimed the listing was unwarranted, but,
approach to its obligate habitat, old growth conifer forests. (Photo in 1988, a coalition of conservation groups filed an appeal
courtesy of T. Ellis and the US National Park Service, public domain) against the agency’s decision in Federal Court. In his
judgment in Northern Spotted Owl v. Hodel, Judge Thomas relative to survival of the owl. Some alternatives were con-
Zilly wrote that “The [Forest] Service disregarded all the sidered in proposed legislation, but none made it to a vote or
expert opinion on population viability, including that of its were included in enacted legislation (Gordon and Lyons
own expert, that the owl is facing extinction, and instead 1997).
merely asserted its expertise in support of its conclusions.” Although an apparent failure at the time, The Panel made a
Zilly ordered the USFWS to reconsider its decision (Gordon valuable contribution to the development of ecosystem man-
and Lyons 1997). agement by demonstrating that a biologically sound solution
By 1989, such successful litigation by the Seattle did not simply require a different management plan, but a
Audubon Society led Forest Service Chief Dale Robertson fundamental change in traditional management practices. All
to appoint an Interagency Scientific Committee (ISC) to 14 management alternatives are now recognized as early
“develop a scientifically credible conservation strategy for models of ecosystem management approaches, each offering
the northern spotted owl” (Meslow 1993). The ISC strategies that broke from the traditional resource manage-
recommended a strategy of habitat conservation areas ment strategy of limiting or eliminating management from
(HCAs) on public forest land in Washington, Oregon, and specific areas of the forest (“preserves”) and permitting inten-
California (Thomas et al. 1990). No timber harvesting would sive management in others (“timber sales”). Consideration of
be allowed in HCAs (Caldwell et al. 1994). In HCAs, the these alternatives also changed political debate from being
so-called “50-11-40” rule provided for dispersal of juvenile discussions about forest acres reserved versus board feet
owls from one HCA to another by requiring 50% of land- harvested to discussions of how to prevent multiple species
scape timber coverage of trees at least 11 inches diameter management problems by integrating management across
with canopy closure of 40% in each quarter township in areas forest landscapes at regional levels (Gordon and Lyons
lying between adjacent HCAs (Harrison et al. 1992; Meslow 1997). It was not surprising, then, given his agency’s experi-
1993; Wilcove 1993). The new plan protected 7.7 million ence with the northern spotted owl, that Forest Service Chief
acres of forests, with 30% of the landscape preserved in Dale Robertson announced, in 1992, that his agency would
old-growth forests (Harrison et al. 1992). At the same time, begin taking an “ecosystem approach” to all subsequent
the USFWS re-examined the status of the northern spotted forest management plans.
owl in 1989 and proposed listing it as a threatened subspecies Had the Forest Service been permitted to follow its tradi-
(Wilcove 1993). By 1990, the northern spotted owl was tional bureaucratic methods, the spotted owl might have been
officially listed. exterminated. The driving force for the shift to an ecosystem
The ISC strategy was the subject of more lawsuits against approach was outside input from citizens, the scientific com-
the Forest Service in 1991 that prevented it from selling munity, and other agencies, all essential elements in the
timber under the new plan. Eventually, US District Judge implementation of ecosystem management. Despite missteps
William Dwyer ruled that the plan carried significant risks to and imperfect solutions, the development of the forest man-
the owl and that the Forest Service had not only failed to agement plan for the northern spotted owl remains a water-
consider the owl’s needs, but also needs of other species in shed event in the emergence of ecosystem management. This
old-growth forests (Harrison et al. 1992). In the meantime, case irrevocably shifted management emphasis from individ-
Congress commissioned its own investigation and assess- ual sites to functional ecosystems, and its resulting
ment committee, the Scientific Panel on Late-Successional recommendations have been incorporated into many
Forest Ecosystems, which came to be known simply as “The subsequent studies of old-growth forests and their manage-
Panel.” The Panel developed 14 management alternatives for ment (Wilcove 1993). Note the progression in management
congressional consideration, each of which was regional in focus and strategies as depicted in Fig. 9.3. Even as popula-
scope, ecosystem-based in management, and risk-driven tion management strategies for the owl changed over time, so
Fig. 9.3 Progression of

population management strategies
developed by the US Forest
Service in their approach to
conservation of the northern
spotted owl (Strix occidentalis
caurina) in the US. Pacific
Northwest. Note how progression
from emphasis on individual pairs
to that of large clusters of pairs
begins to require larger land areas
with appropriate connectivity.
(Figure by K. DeVoss)
did management of its surrounding environment (Fig. 9.4).

The story of the spotted owl is in fact the story of the Point of Engagement Question
emergence of scientific, social, legal, and political elements What elements of the northern spotted owl’s
necessary for ecosystem management to succeed (Fig. 9.5). demographics and habitat preferences created
problems that a resource management approach was
unable to solve? What features of an ecosystem man-
agement approach were better suited to address these
same problems?
9.1.3 The Modern Context: EBM

in Contemporary Conservation
From such initially varied expressions of ecosystem manage-

Fig. 9.4 Progression of habitat management strategies developed by ment, the idea, as well as the practice, is today more com-
the US Forest Service in their approach to conservation of the northern
monly called ecosystem-based management or EBM. EBM
spotted owl (Strix occidentalis caurina) in the US Pacific Northwest.
Note how progression from emphasis on small habitat units eventually has matured into a more widely accepted understanding of an
expands to protection of an ecosystem – the old growth forest. approach that “defines management strategies for entire
(Figure by K. DeVoss) systems, not simply individual components of the
ecosystem. . .As a consequence, EBM takes into account
interactions among ecosystem components and management
sectors, as well as cumulative impacts of a wide-spectrum of
ocean-use sectors. . . EBM considers humans as an integral
part of the ecosystem, since humans derive a portfolio of
services from the ecosystem and also act as a driver
influencing ecosystem processes” (Levin et al. 2009:23).
Before we can understand EBM, we must have a clear
understanding of what we mean by an ecosystem. One classic
definition is all the organisms in a given area interacting
with the physical environment so that a flow of energy leads
to trophic structure, biotic diversity, and material cycles
(Odum 1971). Put simply, ecosystems are energy- and
nutrient-processing systems with physical structures and
functions that circulate matter and distribute energy.
Ecosystems are valued by humans, and so become targets
of human management because of the array of ecosystem
services they provide. Ecosystem services can be defined as
direct and indirect benefits people obtain from ecosystems.
The most widely recognized framework for describing types
of ecosystem services is that of the Millennium Ecosystem
Assessment (MEA), which groups ecosystem services in four
categories: (1) provisioning services (services or goods
needed to produce basic material needs, such as food and
Fig. 9.5 Progressive development of scientific, social, legal, and polit-
ical elements in the case of the northern spotted owl (Strix occidentalis water), (2) regulating services (services which regulate
caurina) in the US. Pacific Northwest. Legislation (passage of characteristics of the ecosystem, such as climate or water
US Endangered Species Act of 1973), social perception and initiative quality, and thus provide stability for human activities and
(lawsuits by private conservation organizations), and judicial review and environments in which provisioning services can be pro-
enforcement (court injunction against employment of traditional natural
resource management approaches for spotted owl conservation) lead to duced), (3) cultural services (materials or experience humans
greater inter-agency cooperation and inclusiveness of public input, value for non-instrumental purposes, such as education, rec-
creating necessary elements for ecosystem management to succeed as reation, or spiritual appreciation), and (4) supporting services
a functionally effective and necessary approach to environmental con- (services which provide a medium (air, water, land) for
servation. (Figure by K. DeVoss)

representation of supporting,
provisioning, cultural and
regulating services with selected
category-specific examples
illustrating goods and services of
benefit to humans provided by
ter
So
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functioning ecosystems. (Graphic
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supporting or permitting the operation of other activities) appreciate that effort invested in identifying fundamental
(MEA 2005) (Fig. 9.6). services, final services, and societal benefits not only aids a
Consider, for example, how an ecosystem might contrib- manager in determining what kinds of benefits to manage for,
ute to regulation of the hydrologic cycle. A basic but also what not to consider in developing a management
eco-hydrologic process such as water uptake and use by plan. For example, although the system has high value in
plants affects an ecological attribute, the quantity of surface food provision, it has essentially no value in energy produc-
water and groundwater available in the system. This attribute tion. This kind of analysis also identifies aspects of the
directly affects a hydrologic service (an ecosystem service), ecosystem that need further investigation.
the amount of water that can be diverted from the system for Let’s take this thinking one step further and evaluate how
human use for municipal, agricultural, commercial, industrial different human activities affect ecosystem components and
or other uses. the ecosystem services a manager wants to derive from them.
Such consideration matters because, before determining a Irit Altman of the University of New Hampshire (USA) and
plan for ecosystem management, it is necessary to describe an her colleagues addressed this problem by devising an
ecosystem in terms of its (1) class (category); (2) fundamental ecosystem-based management system for the Gulf of Maine
services (underlying physico-chemical processes which set in the Atlantic Ocean. Altman et al. (2011) first developed a
up the ecological niches of species inhabiting the ecosystem); matrix (the “human impact matrix”), to evaluate the cumula-
(3) final services (biotic processes through which organisms tive relative impacts of human actions on ecosystem services
in the ecosystem interact with one another and internally (A) which takes the form
regulate the system); and (4) societal benefits (goods and
services taken or used from the ecosystem that add to a11 ... a1j
human well-being). Using a marine ecosystem as one class : : :
of ecosystem, Fig. 9.7 shows examples of each of three other
A¼ : : :
categories (Atkins et al. 2011). By analyzing these
: : :
categorizations in a given system, an ecosystem manager
can gain a better idea of what to manage for, as well as why ai1 ... aij
one should manage at all.
We can see an example of this approach to management of Here, rows are lists of human activities (1 through i, with
the North Atlantic marine system surrounding Flamborough i representing the last numbered row (i.e., the number
Head, UK (Table 9.3). Read this table carefully, and you will corresponding to i is the total number of human activities
Fig. 9.7 Separation of processes and functions of a marine ecosystem environment: integrating ecosystem services and societal benefits with
into fundamental services, final services, and societal benefits. the DPSIR framework in a systems approach. Marine Pollution Bulletin
(Reprinted by permission from Elsevier, from Atkins, J.P., Burdon, 62, 215–226. # 2010 Elsevier Ltd)
D., Elliott, M., Gregory, A.J., 2011. Management of the marine
identified in this system)), and columns are lists of ecosystem In this matrix, each cell represents the strength of interaction
services numbered 1 through j, where j is the number of the between two ecosystem components that intersect in that row
last column, the total number of services identified in the and column.
system. Each cell represents the effect of a human activity on Altman’s team then developed a method of quantitatively
a particular ecosystem service. scoring these interactions so that the relative strength of each
To determine impacts of interactions of natural ecosystem interaction could be quantified and shown in each cell
components on ecosystem services, Altman et al. (2011) (Fig. 9.8). The greater the effect of the activity, the higher
developed a similar matrix to assess the strength of its score. Through additional and more complex calculations,
interactions of the ecosystem components (B) Altman et al. (2011) were then able to score and categorize
human activities relative to their effects on ecosystem
b11 ... b1j components and ecosystem services (Fig. 9.9). Using this
: : : approach, they were able to rank effects of 21 human
activities and 15 ecosystem services. Human activities with
B¼ : : :
greatest effects on ecosystem components and services were
: : :
coastal and watershed development, fishing, extent of marine
bi1 ... bij protected areas, climate change, and toxic contamination.
9.2 Shaping Decision-Making Processes in EBM 367
Table 9.3 Examples of ecosystem services and societal benefits identified in and provided by the marine environment at Flamborough Head,
UK. Reprinted by permission from Elsevier, from Atkins, J.P., Burdon, D., Elliott, M., Gregory, A.J., 2011. Management of the marine environment:
integrating ecosystem services and societal benefits with the DPSIR framework in a systems approach. Marine Pollution Bulletin 62, 215–226.
# 2010 Elsevier Ltd
Category Ecosystem services Description
Fundamental Gas and climate regulation Kelp forests act as a source and/or sink for CO2
services Physical habitat The extensive chalk sea cliffs provide habitat for many nationally and internationally important
breeding populations of seabirds, whilst the waters contain other important features including
littoral and sublittoral reefs, submerged/slightly submerged sea caves, rocky shores, kelp forests
and subtidal faunal turf communities
Nutrient cycling The communities found at the north and south cliff, differ noticeably due to the Flamborough
Front, a summer oceanographic boundary between the northern and southern North Sea creating
a very productive, nutrient-rich environment
Final services Bioremediation of waste There has been a history of anthropogenic activity in the area e.g., distilling, brewing and food
malt production and sewage treatment discharges
Biologically mediated Significant kelp forests (Laminaria hyperborea) and forests of Laminaria saccharina with red
habitat algal undergrowth in nearshore. Physical habitats include chalk reefs, sea caves and maritime
cliff vegetation
Resilience and resistance Despite the anthropogenic activity in the area, current site designations suggest a relatively high
level of resilience and resistance although further evidence is needed
Societal Food provisions The site supports a high level of commercial and recreational fisheries including trawling, netting,
benefits potting and lines in order to exploit the mixed fishery. There is also a developing sea bass fishery
in the area under pair trawling techniques
Raw materials Intertidal areas around the headland have historically been subject to the collection of bait and
fossils
Transport and navigation Due to the presence of chalk reefs, obscured outcrops and strong tidal currents Flamborough
Head has always been a dangerous place for shipping
Energy There is no marine energy generation although offshore wind farm sites are proposed and gas
storage facilities are located in the south of the EMS
Residential and industrial There is no evidence of water being abstracted for residential and industrial uses
water supply
Disturbance prevention Not understood at present
Cultural heritage and The Heritage Coasts covers 19 km and includes archaeological and historical features
identity
Cognitive values Flamborough Head is an educational and research resourced used by schools and universities as
well as statutory monitoring undertaken by relevant authorities
Leisure and recreation Flamborough is a popular tourist destination, with over 56,000 visitors per year. Recreational
activities including angling, bathing, canoeing, walking, bird watching (from both land and sea),
rock-pooling, boating and diving
Feel good or warm glow Existence values are considered likely to be positive because of the site’s outstanding natural
features
Future unknown or Widespread current user values suggest that option use values will be positive
speculative benefits
Altman et al. (2011) were then able to determine that the five
ecosystem services most strongly affected by cumulative 9.2 Shaping Decision-Making Processes
human impacts were biodiversity, marine harvested species, in EBM
aesthetic values, habitat, and recreational values.
The approach used by Altman et al. (2011) requires care- 9.2.1 Criteria that Define Ecosystem-Based
ful analysis of ecosystem components, ecosystem services, Management
and human activities, and its outcomes are only as good as the
information used to score the interactions evaluated. Never- EBM requires inclusion of criteria that distinguish it from
theless, this kind of analytical thinking offers a path to a first more specialized management approaches limited to a partic-
approximation of how to identify the most important factors ular resource, habitat or species (Table 9.4, Arkema et al.
affecting an ecosystem, and then creating strategies likely to 2006). These criteria reflect an emphasis on management that
affect the most important drivers of ecosystem change and recognizes the complexity of the system, incorporates spe-
ecosystem outputs. Such an approach permits us to take our cific and longer-term temporal scales, and encompasses the
first steps toward practical applications of EBM. landscape over which the ecosystem’s processes occur.
Fig. 9.8 Flow chart of the process of scoring effective strength of compromised, or degraded by the interaction. (Reprinted by permission
interaction (“effect strength”) between any two ecosystem components from Wiley, from Altman, I., Blakeslee, A.M., Osio, G.C., Rillahan, C.
or between an ecosystem component and a human activity in a given B., Teck, S.J., Meyer, J.J., Byers, J.E., Rosenberg, A.A., 2011. A
system, ranging from 0 (component A has no effect on component B) to practical approach to implementation of ecosystem-based management:
5 (component B has no resistance to the effect of component A, spatial a case study using the Gulf of Maine marine ecosystem. Frontiers in
scales of the effect are large and, at the temporal scale, the effect is Ecology and the Environment 9, 183–189 # Ecological Society of
chronic (long-lasting or permanent)). The higher the final score, the America)
more strongly the affected ecosystem component is changed,
Consideration of human dimensions requires determination relevant to management issues; (2) an issue identification and
of specific ecosystem goods and services that humans derive scoping process that identifies specific ecosystem manage-
from the managed ecosystem, what economic factors drive ment drivers and pressures; (3) an identification of indicators,
that use, and which human interest groups (stakeholders) most sensitive to such drivers and pressures and thus provide
have the most to gain or lose from the way the system is the basis for assessment of the status and trends of the
managed. ecosystem; (4) a risk analysis that evaluates the probability
EBM approaches often use integrated ecosystem assess- of harm to the ecosystem indicators posed by human
ment (IEA) as a framework for organizing information activities and natural processes; and (5) ongoing monitoring
needed to guide EBM. An IEA is created in a five-step and assessment of ecosystem indicators (Fig. 9.10). For
process: (1) an abstraction of the ecosystem into subsystems example, in a marine ecosystem valued for fish production,
Fig. 9.9 Ranks of human

activities in the Gulf of Maine as
drivers of ecosystem change.
Activities with high relative
scores have disproportionate
effects on ecosystem components
and the ecosystem services they
provide. (Reprinted by permission
from Wiley, from Altman, I.,
Blakeslee, A.M., Osio, G.C.,
Rillahan, C.B., Teck, S.J., Meyer,
J.J., Byers, J.E., Rosenberg, A.A.,
2011. A practical approach to
implementation of ecosystem-
based management: a case study
using the Gulf of Maine marine
ecosystem. Frontiers in Ecology
and the Environment 9, 183–189
# Ecological Society of America)
Table 9.4 Requirements for criteria that can be used to define ecosystem-based management
Criteria category Criteria name Criteria requirement
General criteria Sustainability Emphasizes maintenance of one or more aspects of the ecosystem
Ecological health Includes non-specific goals for ecosystem health or integrity
Inclusion of humans Recognizes that humans are elements in an ecosystem and their education and well-being are
in ecosystem important components of management decisions
Specific ecological Complexity Acknowledges that links between ecosystem components, such as food web structure,
criteria predator-prey relationships, habitat associations, and other biotic and abiotic interactions,
should be incorporated into management decisions
Temporal Incorporates temporal scale and the dynamic character of ecosystems
Spatial Recognizes that ecosystem processes operate over a wide range of spatial scales
Specific human Ecosystem goods and Recognizes that humans use and value natural resources, such as water quality, harvested
dimension criteria services products, tourism, and public recreation
Economic Integrates economic factors into the vision for the ecosystem
Stakeholder Engages interested parties in the management planning processes to find common solutions
Specific management Science-based Incorporates management decisions based on tested hypotheses
criteria Boundaries Recognizes that management plans must be spatially defined
Technological Uses scientific and industrial technology as tools needed to monitor the ecosystem and evaluate
management actions
Adaptive Continue to improve management actions through systematic evaluation
Co-management Promotes shared responsibility for management between multiple levels of government and
stakeholders
Precautionary Manages conservatively when threats to the ecosystem are uncertain
approach
Interdisciplinary Bases management on scientific understanding from several disciplines (ecology, economics,
sociology)
Monitoring Tracks changes in biotic, abiotic, and human ecosystem components for management purposes
Reprinted by permission from Wiley, from Arkema, K.K., Abramson, S.C., Dewsbury, B.M., 2006. Marine ecosystem-based management: from
characterization to implementation. Frontiers in Ecology and the Environment 4, 525–532. # The Ecological Society of America
Fig. 9.10 The five-step process of integrated ecosystem assessment – or management targets and thus evaluate effectiveness of the overall
(1) abstraction of the ecosystem into subsystems relevant for manage- management strategy. (Reprinted under CC BY 4.0 International, from
ment permits initial conceptual understanding of the system; (2) issue Holsman, K., Samhouri, J., Cook, G., Hazen, E., Olsen, E., Dillard, M.,
identification and scoping process identifies key management objectives Kasperski, S., Gaichas, S., Kelble, C.R., Fogarty, M., Andrews, K.,
and constraints; (3) identification of indicators names variables that 2017. An ecosystem-based approach to marine risk assessment. Ecosys-
serve as proxies for the state of the ecosystem and are most sensitive tem Health and Sustainability 3, e01256 (right graphic), and from Levin,
to major drivers and pressures that affect it; 4) risk analysis evaluates the P.S., Fogarty, M.J., Murawski, S.A., Fluharty, D., 2009. Integrated
probability that indicators will fall below management targets, a sign of Ecosystem Assessments: Developing the Scientific Basis for Ecosys-
harm to the indicators posed by human activities and natural processes; tem-Based Management of the Ocean. PLoS Biology 7(1): e1000014
and (5) ongoing monitoring and assessment of ecosystem indicators will (left graphic)
be used to assess or “score” their state relative to historical benchmarks
the abstraction stage would focus on identifying subsystem Adaptive management is critical to EBM because ecosys-
components relevant and responsive to management actions, tem management is management of uncertainty. Managers
such as components for individual harvested species. The must predict responses and interactions in ever-changing
scoping stage would identify focal resources and primary environments but do so without a full understanding of
drivers of the system, and might include elements of fishery processes and components that determine ecological states.
description, detailed objectives, or fishing activities (and their They must draw inferences and make predictions from data
potential hazards). Indicators might be represented by indi- fraught with human unreliability, biases and limits in inter-
vidual species, habitats or communities (Table 9.5). Risks pretation (Christensen et al. 1996). Uncertainties cannot be
(actual probability of experiencing harm) would be calculated eliminated, but the effects of some kinds of uncertainty can
for specific hazards (things that actually cause harm) be mitigated. Understanding of ecological systems can be
associated with specific fishing activities, as well as external enhanced through research targeted to the most important
activities that could affect the ecological system. A plan for questions of ecosystem behavior. Biases and human
and practice of monitoring and assessment would track num- limitations can be constrained with dedication to high levels
bers, abundance, or status of previously identified species, of professionalism in analysis and interpretation of data, peer-
habitats, or communities designated to inform managers review of research efforts, and humility about one’s own
about the health of the system. hypotheses (Christensen et al. 1996).
The most important tool in this process is the practice of
adaptive management, “the structuring of policy or manage-
9.2.2 The Role of Adaptive Management ment actions as a set of testable hypotheses to promote
learning from policy implementation, and to allow for greater
The choice of method used to inform decisions is arguably the adaptability when change does inevitably occur within the
most critical step in creating effective systems of EBM. A system” (Lamont 2006:7). In adaptive management, the dual
good EBM plan does more than track values of indicator goals of system performance and acquisition of reliable
variables. It integrates management actions into experimental knowledge are accomplished simultaneously so that
analysis of change in indicators, an approach known as managers learn about the system through implementation of
adaptive management. management actions. An adaptive management approach
Table 9.5 Some measurable ecosystem variables that can be used as performance indicators in ecosystem management
Properties of fish and benthic communities (Performance measures) Metrics (Reference points)
Biodiversity of Species
Biomass Sum of weight across species from survey
Size-structure Slope of size-spectrum
Length-frequency distributions of species
Multi-dimensional ordination
Species identities Species presence/abundance
Index of declining or increasing species
Presence of indicator, charismatic, sensitive species
Nonindigenous species
Species diversity Theoretical distribution metrics
Taxonomic Diversity indices
Ecological Functionality
Resilience Return time of properties of food webs
Invasibility
Productivity P/B ratio
Carbon per unit area/time/volume
Partitioning of production between somatic and gonad material
Trophic structure Connectance
Path length
Throughput Internal consumption to yield
Ulanowiez index
Body well-being Condition factor
Incidence of disease, pathogens, parasites, contaminants
Reprinted by permission from Wiley, from Hall, S.J., Mainprize, B., 2004. Towards ecosystem-based fisheries management. Fish and Fisheries 5,
1-20. # 2004 Blackwell Publishing Ltd
requires designing management actions as experiments

through direct manipulation of the system and willingness Information Box 9.1 (continued)
to change management strategies based on experimental careful monitoring over time. McClanahan and Obura
results, as well as a willingness to change research priorities (1996) exemplified such an approach in a classic
according to management needs. Adaptive management ecosystem-scale “experiment” on African coral reef
also requires ongoing interaction with stakeholders to com- systems, using Kenyan marine reserves as controls
municate results in meaningful ways, and, from such interac- and comparing them to adjacent marine areas open to
tion, learn what stakeholders consider meaningful research. commercial fishing. The species richness of coral and
Study the application of these concepts in an example fishes was higher in the protected areas, and differences
of a marine reserve in Information Box 9.1. In current in species richness between protected and unprotected
management research, these kinds of efforts are more for- areas increased with size of sampled area (Information
mally developed as theories of “performance-based Box Fig. 9.1). This study was not fully “experimental”
management.” because it lacked baseline information (species rich-
ness of reserve areas was unknown before they became
reserves) as well as knowledge of specific causes of
Information Box 9.1: Adaptive Management:
decline for particular species. However, past efforts
Applications in a Marine Reserve
like this one are good examples of early attempts to
In adaptive management, management goals and
document differences that ecosystem protection can
strategies are hypotheses to be tested by experiments
make, and then begin, in this case, to identify fishing
characterized by careful design, inclusion of environ-
and collection methods most harmful to the diversity of
mental controls (unmanipulated sites or subjects), and
the system.
(continued)
(continued)
Information Box 9.1 (continued)
Information Box Fig. 9.1 Species richness in Kenyan marine reserves and adjacent areas open to collecting and commercial fishing. (Reprinted by
permission from McGraw-Hill Publishers)
9.2.3 Evaluating Ecosystem-Based violated, could be taken as prima facie evidence for conser-
Management as a Performance-Based vation concern. When the value of the indicator reaches the
System threshold level, the manager is facing an unacceptable risk of
harm to the resource or the system (Hall and Mainprize
9.2.3.1 Theoretical Constructs 2004).
for Performance-Based Evaluation Target and limit reference points have been compared to a
Like any management system, EBM must measure perfor- “green light – red light” mentality of management, with
mance against targets. So, what are the targets? Specifically, “threshold” reference points being the equivalent of a “yel-
what variables should be chosen as performance indicators? low light” (Fig. 9.11). In some cases, the same variable, at
During the process of ecosystem assessment described previ- different values, can serve as both target indicator and limit
ously, managers must identify appropriate indicators of eco- indicator. Although this kind of monitoring was originally
system states, then validate their choices by monitoring and developed for and most widely used in fisheries management,
manipulating the indicators to determine if they really do it can be applied to any ecosystem management effort in
track trends in the ecosystem (Levin et al. 2009). An indicator which performance measures and reference points meet key
may track abundance of a single species, or, more often, serve criteria. The performance indicator must be: (1) quantifiable,
as a proxy for more general ecosystem attributes. This means because a “best estimate” of the indicator is required, along
that, when indicators reach critical values (“thresholds”), this with a measure of uncertainty about the estimate; (2) simple,
should trigger pre-determined management actions. Perfor- because the indicator must be easy to understand and inter-
mance indicators are reference points intended to tell us pret, not only for managers but for stakeholders who take part
something about the state of the system and how to respond in the decision-making process; (3) relevant, because the
to it. indicator must be related to management objectives, such
Target indicators are appropriate in production-oriented that there is a clear connection between changes in the refer-
ecosystem management in systems managed to produce cer- ence point and changes in the system, not merely a mysteri-
tain levels of production, like a specified level of stock or ous correlation; (4) tractable, so that the indicator can be
biomass of some resource, such as timber or game that a changed by management action; (5) faithful, because the
manager wants to remove from the system. Limit indicators indicator must convey accurate information in every instance,
are used in risk-averse ecosystem management. Here the not just in some instances or under ideal conditions; (6) com-
concern is avoidance of unacceptable risk, such as endanger- parable, so that the value of the reference point at one time
ment of key species. The limit indicator serves as part of an can be compared to its value at other times, even over long
“early warning system” designed to detect or predict the point periods of monitoring; and (7) cost-effective, so that its value
where the system begins to come under stress. A good limit can be obtained and actions directed by the value applied
indicator would be a measure of a property of a resource, without excessive expense (Hall and Mainprize 2004).
such as recruitment rate, harvest per effort, density, dissolved A more recent effort to build these concepts into specific
oxygen level, pH, or other biotic or abiotic metric that, if management evaluation and assessment systems is the
Fig. 9.11 An illustration of

target (“green light”), threshold
(“yellow light”), and limit (“red
light”) reference points that can
serve as performance indicators of
ecosystem production, function,
or risk. In this example, level of
“sustainable system biomass” of
an unspecified system component
serves as the indicator variable.
Wiley, from Hall, S.J., Mainprize,
B., 2004. Towards ecosystem-
based fisheries management. Fish
and Fisheries 5, 1-20. # 2004
Blackwell Publishing Ltd)
Drivers – Pressures – State Change – Impact – Response terrestrial ones (Table 9.5). Here are two examples that illus-
(DPSIR) approach. DPSIR is a systems-based approach trate how these concepts can be employed in decision
designed to capture key relationships between an ecosystem making.
and the human community surrounding it, especially the
actions of human society that do or can change the ecosys- 9.2.3.2 The Black-Legged Kittiwake
tem. In DPSIR, Drivers are key demands made on the eco- and the Swamp Wallaby
system by the human society. Drivers create Pressures on the
system which are actions, stimulated by human demands, that Kittiwakes, Sand Eels and Performance-Driven
result in State Changes in the ecosystem, leading to Impacts Management
(actual effects of State Changes on the ecosystem) that The black-legged kittiwake (Rissa tridactyla) (Fig. 9.13a), a
require a Response by management if the human society is gull-like bird with widespread global distribution, is currently
to continue to receive needed ecosystem services (Atkins listed by the IUCN as “Vulnerable,” as the regional popula-
et al. 2011) (Fig. 9.12). The DPSIR system has contributed tion of the North Sea has declined more than 50% since 1990,
to better definition and understanding of key concepts in making it a species of increasing conservation concern.
EBM, better formulation of management assumptions, and Although kittiwakes can be killed by being caught on fish
improved development of decision-making frameworks for hooks of commercial fishing vessels (because they try to take
incorporating concepts into management strategies and bait from hooks) or entangled in nets or lines, the most
actions. significant threat is reduction of their primary food, the lesser
Fisheries managers have taken the lead in initiating ways sand eel (Ammodytes spp. and Gymnammodytes spp.)
to assess these ecosystem characteristics and developed (Fig. 9.13b). Historically abundant, sand eels, which are
indicators that can be used to assess system biodiversity or actually eel-like fish, represent the largest single industrial
ecological functionality, or that can be used effectively in fishery in the North Sea and are often a primary food for sea
combination, not only in aquatic ecosystems but also in birds, including kittiwakes. Breeding success of kittiwake
Fig. 9.12 Schematic representation of the Drivers – Pressures – State Burdon, D., Elliott, M., Gregory, A.J., 2011. Management of the marine
Change – Impact – Response System (DPSIR) as a framework for environment: integrating ecosystem services and societal benefits with
developing decision-making strategies for ecosystem manage- the DPSIR framework in a systems approach. Marine Pollution Bulletin
ment. (Reprinted by permission from Elsevier, from Atkins, J.P., 62, 215–226. # 2010 Elsevier Ltd)
Fig. 9.13 (a) The black-legged kittiwake (Rissa tridactyla) and (b) the responses which have led to closures of sand eel fishing in some areas.
sand eel (Ammodytes spp.), its most important food. Kittiwake produc- (Kittiwake photo courtesy of S. Schoen, US Geological Survey; Sand
tivity in the North Sea provides ecosystem managers with important eel photo courtesy of M. Arimitsu, US Geological Survey)
performance indicators that generate predetermined management
populations, indexed as the number of chicks per nest, is to extract high quality information from management actions
closely correlated with abundance and availability of sand by implementing them as experiments. As Australian ecolo-
eels. Certain regions of the North Sea have been declared gist Julian Di Stefano observed, “. . .the existence of ecologi-
critical habitat for kittiwakes, and stakeholders involved in cal data and expertise will not result in improved
managing the system, including representatives of the scien- management outcomes unless positive relationships between
tific community, fishing industry, and government environ- ecologists and managers exist” (Di Stefano 2004:62).
mental agencies, agreed that such regions would be closed to In the southeastern Australian province of Victoria, state-
sand eel fishing if breeding success of kittiwakes fell below administered forests are managed for optimal sustainable
0.5 chicks per nest for three successive seasons. This trigger timber harvesting, but a common species of herbivore, the
point has been reached in some kittiwake populations, and swamp wallaby (Wallabia bicolor) (Fig. 9.14), causes exten-
one area of the North Sea, the Ford of Firth, was then closed sive damage to potentially harvestable timber and timber
to sand eel fishing (Hall and Mainprize 2004). regeneration in recently logged stands through browsing.
Here we see a system in which ecosystem management is Swamp wallabies are generalist feeders that spend much of
both production-oriented (harvesting sand eels) and risk their time in densely vegetated environments, usually feeding
averse (avoiding endangerment of kittiwakes). In this case, close to shelter vegetation (escape cover). Thus, an optimal
risk aversion trumps production, and concerns for the viabil- combination of food and cover defines their preferred habitat.
ity of regional kittiwake populations override concerns to Past harvesting practices used by the Victorian Department of
sustain regional sand eel fishing, leading to fishing season Sustainability and Environment (DSE) often produced such
closure on sand eels. Notice that the performance indicator, preferred habitats for swamp wallabies on logged sites where
chicks per nest, meets all seven criteria needed to make it an the wallabies’ favorite food (seedlings and other regenerating
appropriate index for management actions. plants) grew close to dense cover vegetation. Changing
harvesting procedures and silviculture manipulations might
Linking Research Insights and Policy Decisions – The reduce the browsing problem. Di Stefano, an ecologist with
Swamp Wallaby in Australian Forests the DSE, designed an adaptive management study focused on
If EBM is to have a basis in science and a foundation for answering these questions: (1) How widespread was the
professional credibility, it must first understand the specific problem of excessive browsing of regenerating timber by
information needs of ecosystem managers and then be able to swamp wallabies? (2) Were there ecological and habitat
translate research findings into informed, “on-the-ground” factors that could be used to predict browsing damage before
management and policy decisions determinations at local, it occurred? (3) Were current browsing reduction strategies
site-specific levels of decision making. This requires meeting management objectives? (4) If not, what new brows-
established and ongoing channels of communication and ing reduction strategies might work?
high levels of trust between researchers and managers. As with kittiwakes and sand eels, we see here a manage-
Managers must know how to place knowledge of ecosystem ment system simultaneously concerned with two criteria. It is
management processes in the context of existing information production oriented (for sustainable timber harvest), and
and work closely with researchers to design and conduct new therefore concerned with performance indicators that mea-
studies to fill knowledge gaps and determine in advance how sure production levels, such as the rate of forest regeneration
Fig. 9.14 The swamp wallaby (Wallabia bicolor) provides ecosystem

managers with performance indicators based on its level of browsing
damage and effect on regeneration of tree species in harvested areas in
Australian forests. (Photo courtesy of J. O’Neill, reprinted under CC-
BY-3.0)
Fig. 9.15 Three theoretical threshold relationships between browsing

in logged stands. But it is also risk averse (desirous to prevent damage and animal abundance that could hypothetically apply to the
unacceptable browsing levels on timber by wallabies) and effect of browsing by swamp wallabies on forest regeneration in
Australia. In A, animal abundance must be greatly reduced to observe
interested in determining threshold values of browsing that
significant reduction in browsing damage. In B, animal abundance and
would indicate critical stress to the system’s ability to pro- browsing damage vary directly and proportionally. In C, a small reduc-
duce new timber. The performance indicator is the level of tion in animal abundance leads to a disproportionately large reduction in
browsing that reduces regeneration of trees below the level at browsing damage. (Reprinted by permission from Wiley, from Di
Stefano, J., 2004. The importance of ecological research for ecosystem
which the stand can regenerate, a classic “threshold value.”
management: the case of browsing by swamp wallabies (Wallabia
Here production and risk are linked in the same indicator, bicolor) in commercially harvested native forests. Ecological Manage-
regeneration rate. Thus, Di Stefano noted, “Objective determent and Restoration, 5, 61–67. # 2004 Ecological Society of
mination of an acceptable browsing level provides a scientif- Australia)
ically defensible trigger for management action and requires
linking early damage levels with an accepted regeneration
information that could produce effective adaptive manage-
standard. If browsing results in a failure to meet the regener-
ment. The question is, will they? That query is not unique to
ation standard, browsing may be considered unacceptable”
Australia. It must be addressed in the global conservation
(Di Stefano 2004:63).
effort if ecosystem management is to become a process that
The relationship between abundance of wallabies and
can advance conservation strategy and long-term ecosystem
browsing damage could take several forms, as shown in
sustainability.
Fig. 9.15. If the relationship follows curve A, reducing
wallabies from abundance X to abundance Y would have
little effect on the amount of damage done by browsing. If Point of Engagement Question
curve B, the relationship is direct and proportional, such that Consider two dimensions of “Di Stefano’s Dilemma.”
every unit of reduction in wallabies results in a proportional First, design an experiment that would determine the
unit reduction in browsing damage. If curve C, a small relationship between stand regeneration and wallaby
reduction in wallabies leads to a disproportionately large browsing. Second, design a model of organizational
reduction in browsing damage. The goal of the experiment structure in the Victorian DSE that would make it
would be to apply different levels of wallaby removal in easier for managers and researchers to work together
different areas, measure the associated regeneration response, in designing the experiment and for transferring
and see which pattern the data fit. and applying the data gained from it to management
Because government agencies employ both managers and action.
ecologists, they are in a position to facilitate exchange of
9.3 Scientific Foundations of Ecosystem-Based Management 377
9.3 Scientific Foundations National Park, Big Cypress National Preserve, Biscayne
of Ecosystem-Based Management National Park, three water conservation areas (WCAs), and
numerous smaller state and private reserves protect about
9.3.1 The Problem of Location – Where Is 67% of the land area of the original ecosystem. But, although
the Ecosystem? the majority of the Everglades land area was protected, its
water flows were not. The construction of 2200 km of canals
An adaptive management approach employing specific per- and levees, over 40 pumps and spillways, and impoundment
formance indicators is an important component of effective of neighboring Lake Okeechobee to the north permanently
EBM. But even with a well-designed program of adaptive altered the amount and timing of water through the ecosys-
management, EBM will remain challenging because it tem, causing long-term, severe degradation (Lockwood and
attempts to manage a constantly changing entity, an ecosys- Fenn 2000). A long-embattled, but finally approved restora-
tem. To understand how ecosystems change and how such tion plan will require a multi-billion dollar effort spent over a
change might be managed, an ecosystem manager must period of more than 30 years to rectify current deficiencies in
determine boundaries within which adaptive management water flow. The Everglades is representative of one of the
will operate, as well as what it can investigate. These include most important and chronic problems in conservation and
determinations of: (1) physical boundaries of the system to be EBM, the determination of the actual ecosystem that sustains
studied, fundamental landscape units within the system, and a protected area. This problem has become the subject of
spatial and temporal scales at which they should be studied; increasing attention and recent research.
(2) meaningful ecological models of the system;
(3) methodologies for collection and monitoring of relevant
data at scales appropriate to the system and its model(s); 9.3.2 Do Protected Areas Protect Ecosystems?
(4) methods to identify, measure, and manage the most
important processes, both natural and anthropogenic, affect- Worldwide conservation effort incorporates establishment
ing transfer of matter and energy in the system; (5) how and monitoring of protected areas as one of the cornerstones
ecological processes interact with landscape processes and of global conservation strategy. However, almost all
scales; and (6) final adaptive management approaches. The protected areas, both large ones like the Everglades and
problem to be solved is how to design and conduct small- those that are much smaller, have been and continue to be
scale experimental manipulations within the system to test established under explicit political and social constraints
predictions of the model(s) and then determine likely which always include considerations of human welfare. As
responses to natural disturbances and management practices a result, a given protected area is almost never as large as
based on these experimental results. conservationists would like it to be. The protected area itself
Just as a general definition of ecosystem can be elusive, the is usually only a subset of a larger ecosystem, and therefore
particular limits of an individual ecosystem to be managed vulnerable to changes that may occur in unprotected portions
can be hard to define, even when there are no jurisdictional of the ecosystem within which the protected area and its
constraints and the best available scientific information is species interact. As a result, many protected areas worldwide
employed. Ecosystems have notoriously “leaky” boundaries. have undergone degradation. The causes are manifold, but
They are open systems, not closed to gains or losses in terms often linked to human activity. Ecological processes like
of organisms, matter, and energy. Different agencies and disturbance regimes (for example, frequency of fires) are
individuals approach the problem of ecosystem definition in often altered within protected areas. Non-native species
a variety of ways, depending on ecosystem characteristics have entered the ecosystems, sometimes leading directly to
and management objectives. extinction of native species. The problem to be solved today
Traditional political delineations are the easiest way to is how to maintain, improve or restore the ecological condi-
delimit boundaries of an ecosystem and have the attraction tion of protected areas despite changes on the lands
of placing the management area under a single administrative surrounding them.
jurisdiction, or at least under a group of related jurisdictions. To address this problem, conservationists have developed
Unfortunately, this method is almost never ecologically techniques of delineating protected area-centered
meaningful because ecological processes rarely match ecosystems (PACEs). PACES can be defined, on a case-by-
borders of management jurisdictions. Examples of the case (protected area-by-protected area) basis, and threats to
shortcomings of politically-based ecosystem designations them identified, by using comprehensive scientific methods
are as numerous as they are depressing. One of the most to map and analyze land-use changes in and around the
tragic is the case of the Everglades ecosystem of south protected area. Hansen et al. (2011) recently applied this
Florida (USA). The combined holdings of Everglades approach to 13 US national parks by evaluating four critical
ecological elements that determine size and configuration of a resources for these species. For example, the first and most
functional PACE (Fig. 9.16); namely ecological flows, effec- famous national park in the United States, Yellowstone,
tive size of the protected area, crucial habitats, and degree, contains no true winter range for two of its most charismatic
distribution and penetration distances of various human- mammal species, elk (Cervus canadensis) and bison (Bison
induced edge effects. Ecological flows include movements bison). Both would normally move out of the park to lower
of material, organisms, and energy that are waterborne or elevations during winter but are prevented from doing so by
airborne, as well as movements of disturbances such as fire. hunting at park borders or removal from adjacent private land
In its original form, such flows sustaining ecological pro- due to health concerns related to their potential for transmit-
cesses in a protected area may have been well-connected to ting disease to domestic livestock. In addition to these kinds
its ecosystem in the surrounding landscape (Fig. 9.16a), but of specific dilemmas, more general kinds of problems
the effective size of the surrounding supporting ecosystem associated with human-induced edge effects (Fig. 9.16e)
may be reduced over time by human activities which disrupt may be magnified along borders of a protected area, reducing
these flows (Fig. 9.16b). Actual forms and pathways of the amount of interior habitat – needed by some species,
ecological flows for a given protected area must be defined especially habitat specialists, as we saw in Chap. 7.
in their present condition and are affected by land use outside Using all of these criteria, combined with scientific
the officially designated protected area (Fig. 9.16c). Crucial mapping technology, Hansen et al. (2011) created boundaries
habitats include season-specific use areas, population of the PACE associated with each national park by
sources areas, movement paths, or some portions of annual identifying what maps needed to be made, what data were
home ranges for populations within protected areas. For needed to construct the maps, and what GIS layer (attribute-
many animals, especially migratory species, these crucial specific map) would be the result (Fig. 9.17). The resulting
habitats exist outside the protected area (Fig. 9.16d) but are PACEs identified averaged 6.7 times larger than the parks in
essential to complete life cycles and provide necessary upper watersheds and 44.6 times larger than those in middle
Fig. 9.16 Schematic illustration of the process of defining a protected exist outside the park, and are needed to complete life cycles and
area-centered ecosystem. (a) In its original form, a protected area may provide resources essential for such species. (e) Edge effects may be
have been well-connected to its ecosystem in the surrounding landscape magnified along the borders of a protected area, such as a national park,
in flows of energy, materials, and organisms. (b) The effective size of the reducing the amount of true interior habitat needed by some species,
surrounding supporting ecosystem may be reduced over time by human especially habitat specialists (Chap. 7). (Reprinted by permission from
activities which disrupt these flows. (c) Actual forms and pathways of Hansen, A.J., Davis, C.R., Piekielek, N., Gross, J., Theobald, D.M.,
these ecological flows must be defined in their present condition and are Goetz, S., Melton, F., DeFries, R., 2011. Delineating the Ecosystems
affected by land use outside the officially designated protected area. (d) Containing Protected Areas for Monitoring and Management. BioSci-
Crucial habitats for many species, especially migratory species, often ence 61, 363–373. # 2011 American Institute of Biological Sciences)
Fig. 9.17 Framework for delineating protected area-centered Piekielek, N., Gross, J., Theobald, D.M., Goetz, S., Melton, F., DeFries,
ecosystems (PACEs) using data available for the United States. R., 2011. Delineating the Ecosystems Containing Protected Areas for
Abbreviations: LANDFIRE, Landscape Fire and Resource Management Monitoring and Management. BioScience 61, 363–373. # 2011 Amer-
Planning Tools Project; PA, protected area; SAR, stock assessment ican Institute of Biological Sciences)
review. (Reprinted by permission from Hansen, A.J., Davis, C.R.,
watersheds (Fig. 9.18). Note in this figure just how much this 1999; Richardson and Gatti 1999). For example, managers
difference means when you can visualize it on an actual map! in India divided the country into 20 land resource regions
As Hansen et al. (2011) themselves noted, “The sizes of these (LRRs) and 186 land resource areas (LRAs) based on an
PACEs clearly emphasized the long-term reliance of park integrative assessment of differences in soil, rainfall, forest
biodiversity on surrounding landscapes. PACEs in the eastern cover, land use practices, water resources, and elevations.
United States were dominated by private lands with high rates This system was refined to 17 soil conservation regions
of land development, suggesting that they offer the greatest classified according to climate, rainfall, mean annual temper-
challenge for management. Delineating PACEs more broadly ature, elevation, watershed boundaries and land use
will enable monitoring, assessment, and conservation of (Table 9.6, Rama Mohan Rao et al. 1999). In Wisconsin
protected areas worldwide degraded by human activities in (USA), managers used watersheds as the fundamental unit
the environments surrounding them (Hansen et al. 2011). in an ecosystem management approach to restore drained
What about the problem of delineating ecosystems not agricultural wetlands (Richardson and Gatti 1999). Using
associated with protected areas? Are there techniques that GIS, they combined satellite imagery with information on
can be applied more generally, even with ecosystems of wetlands from state and federal wetland inventories to pro-
vastly different characteristics? Just as the relative position duce an integrated, digitized database that could be used to
of these US national parks in their surrounding watersheds identify wetlands and account for changes in wetland status
dramatically affected size of the real protected area required, over the previous 12 years. Wetlands were then ranked based
so watersheds are becoming, not only in this case but more on rates of soil loss and summed within a watershed. Resto-
systemically, an increasingly favored framework for ecosys- ration of upper-elevation wetlands proved to have the greatest
tem definition. effect on lowering water velocities to all downstream sites,
greatly reducing erosion and sediment delivery to down-
stream areas (Richardson and Gatti 1999).
9.3.3 Using Watersheds to Define Ecosystem Watersheds and wetlands are examples of an EBM
Limits, Boundaries, and Processes approach known as, sectoral management a category of
EBM methods that invoke spatial planning. Sectoral man-
In EBM, many have begun to define the ecosystems as agement is particularly helpful in marine systems where
collections of watersheds within a defined area or that system boundaries are three-dimensional and more difficult
empty into a common source (Rama Mohan Rao et al. to define. In such cases, both stresses and management
Fig. 9.18 Maps of protected area-centered ecosystems (PACEs) for Scenic and Recreational River. (Reprinted by permission from Hansen,
13 US national park units. Gradations in color outside parks indicate A.J., Davis, C.R., Piekielek, N., Gross, J., Theobald, D.M., Goetz, S.,
number of overlapping classification criteria. Sites with many Melton, F., DeFries, R., 2011. Delineating the Ecosystems Containing
overlapping criteria may be considered more important for monitoring Protected Areas for Monitoring and Management. BioScience
and management. Abbreviations: NP, National Park; NRA, National 61, 363–373. # 2011 American Institute of Biological Sciences)
Recreation Area; NRRA, National River and Resource Area; SRR,
jurisdiction may be location-specific, the latter often defined 9.3.4 Knowing the System– What Data Should
by international boundaries or other conventions. In a sec- Be Collected for EBM?
toral management approach that emphasizes jurisdictional
boundaries of different management entities, such as nation 9.3.4.1 General Considerations
states, each sector (national territory) regulates particular Ecosystem managers face difficult choices about which infor-
activities or projects taking place at a particular location mation to collect and how to interpret it, but they can take
(Douvere 2008) (Fig. 9.19). In some cases, different initiative to make informed decisions by collecting, monitor-
countries will establish a common watershed system across ing and reporting appropriate ecosystem data at appropriate
spatial scales that facilitate a higher degree of cooperative scales. From such information, they can determine the range
management for a central water body of interest. For exam- of fluctuations of ecosystem processes and components in
ple, Ontario, Canada has developed an integrated watershed- different systems, and whether existing data in their system
based approach to management of Lake Erie water quality, are within those ranges. Several important methodologies and
which has since been adopted by US states bordering that types of data – both old and new – exist to help achieve
lake (Great Lakes Commission 2019). ecosystem management goals.
Table 9.6 Major characteristics of soil conservation regions in India that serve as a basis for defining watershed-based management, one practical
approach to defining ecosystems for ecosystem management
Rainfall Mean Temp Growing period (degree
Region Climate (cm) ( C) days)
Glacier Cold arid 0–100 <20 0–90
Karewas Cold semiarid (dry) to humid and 60–250 20.0–22.5 90–300
perhumid
Shiwalik Semiarid to humid and perhumid 20–150 20.0–27.5 120–300
Indogangetic Plain Subhumid dry to subhumid moist 100–150 22.5–27.5 180–210
Arid Typic/arid to hyperarid 5–50 25.0–27.5 0–90
West Alluvial Plain Superarid dry to semiarid moist 50–150 25.0+ 90–150
Beehar Semiarid dry to moist 40–150 22.5–27.5 120–180
Southern Malwa Semiarid moist to subhumid dry 75–150 22.5–27.5 120–150
Plateau Semiarid dry to semiarid moist 75–150 20.0–27.5 90–150
Chalka Arid (typic) to semiarid moist 50–250 20.0–27.5 60–150
Western Ghat Subhumid to perhumid 100–250 25.0–27.5 240–270
Central Eastern Upland Subhumid dry to subhumid moist 75–150 22.5–27.5 150–180
Eastern Ghat Semiarid dry to subhumid dry 100–150 25.0–27.5 120–210
Diara Subhumid dry to subhumid moist 100–150 25.5–27.5 150–210
Sundurban and Eastern Subhumid dry to perhumid 100–150 20.0–27.5 210–300
Valley
North Eastern Hill Humid to perhumid 150–250 20.0–22.5 270–300
Island Humid to perhumid 160–300 20.0–28.0 240–300
Watershed boundaries and land use not included in this table. Based on data from Rama Mohan Rao et al. 1999
Fig. 9.19 Essential elements

needed to implement a spatial
management process in a target
ecosystem. Compare with the
elements of Integrated Ecosystem
Assessment (IEA) described
previously and displayed in
Fig. 9.10. (Design by K. DeVoss,
based on Ehler and Douvere 2009,
UNESCO)
One way to determine ecosystem health is by identifying National Park Service (NPS) scientists Steven Fancy and
the presence of ecological stress and the system’s response Robert Bennetts identified three critical elements needed for
(recall our earlier discussion of limit indicators as part of an effective long-term monitoring programs. These elements
“early warning system” for detecting system stress). Because were (1) relevance- attained by establishing clearly defined
ecosystems are successional in nature and often show signs of goals and objectives that actually matter to NPS management
stress only over long periods, long-term data are essential to and assessing these with carefully selected indicator variables
evaluate ecosystem condition. There are five kinds of long- -; (2) reliability – established by framing the data collection
term data available for ecosystems: (1) regularly collected in the context of conceptual models (Fig. 9.21) that accu-
data; (2) remotely-sensed data adaptable to GIS (3) archived rately describe ecosystem components and interactions – and
data from previous studies; (4) data from long-term natural (3) commitment- created by an agency’s determination to
repositories; and (5) data from preserved areas within the continue the program with solid funding support and
ecosystem not subjected to similar disturbance. pre-determined pathways to feed information gained from
monitoring back to management decision making that will
9.3.4.2 Regularly Collected Data demonstrate the value of the monitoring program. Where
Regularly collected data, sometimes incorrectly called “con- long-term sampling units do not exist in an ecosystem,
tinuously collected data,” are obtained from surveys or managers should establish them using standard survey
samples that measure the same variables, at the same methods within a well-conceived (and documented) design
locations, at regular intervals. They are a critical component and begin regular sampling and monitoring. Random or
of the larger realm of ecosystem monitoring, an essential tool systematic sampling points can be chosen by using a land-
for managing uncertainty in attempting to describe or predict scape grid to ensure that samples are representative of the
the behavior of ecosystem processes. For example, in specific entire area and that adequate numbers of sites are sampled.
regions and ecosystems, vegetation data are collected at Grid sampling can provide large amounts of data, but any
annual or otherwise regular intervals along permanently particular grid assumes that the sampled environment is rela-
established transects. The historic “Parker Transects” of tively homogeneous at the sampling scale. Where the envi-
Yellowstone National Park have provided estimates of ronment is obviously heterogeneous, multiple grids may be
range condition since 1954 (Fig. 9.20), although some necessary, one for each kind or category of landscape or
measurements began as early as the 1930s (Coughenour ecosystem encountered. As grids within categories or sam-
et al. 1994). Measurements taken inside and outside of pled points within grids multiply, sampling becomes more
long-established enclosures (wire pens that exclude labor intensive and costly. And in remote areas, randomly
ungulates) have helped to determine long-term effects of selected sampling points may be hard to reach. These and
grazing and browsing on plant communities, as McInnes other factors create tension between the number of variables
et al. (1992) did in Isle Royale National Park (USA) in a sampled and the number of points sampled. Sampling more
classic study evaluating effects of moose (Alces alces) brows- variables provides more information, but the increase in time
ing on forest ecosystems. investment per point reduces the sample size and reliability of
Permanent sampling sites, like the Parker Transects, observed differences.
revisited regularly over time, are powerful tools for ecosys- Today long-term, regularly collected data acquisition and
tem monitoring and, as a result, ecosystem management. analysis are facilitated by remote sensing techniques, contin-
Revisiting the same sites not only detects change over time uously operating recording equipment (“data loggers”) and
but increases the precision of the estimate of change in the GIS. But technology employed without insight will not
measured variable, and so increases the power to detect improve management decisions. The actual survey design is
temporal change. Managers need such precise, longer-term a foundational element of the scientific credibility of the data
data to understand and identify changes in ecosystems and derived from the survey, the method employed to do the
determine whether such changes reflect variability in the survey is critical. Modern survey techniques now usually
system or undesirable human influence (Fancy and Bennetts develop, in advance, a probability design in which each
2012). Although reliable long-term monitoring data are in unit of the target population in the survey has a known,
increasingly high demand and undeniably important to man- non-zero probability of being included in the sample, and
agement decision-making, the record of establishing and there is some random component to the selection of sampled
maintaining such long term monitoring has been poor sites (Fancy and Bennetts 2012). Surveys that incorporate
(Lindenmeyer and Likens 2009), and, of those that have probability designs use a spatially balanced sampling design
been established, many have been hurriedly planned, insuffi- because sample sites that are spatially balanced (more or less
ciently funded, and prematurely ended (Fancy and Bennetts evenly dispersed over the area or resource being sampled)
2012). In a comprehensive review of current and past long- more efficiently cover the sampling area and more represen-
term monitoring programs employed in US National Parks, tative of the resource’s distribution than simple random
Fig. 9.20 The historic Blacktail Sage Belt #2 transect in Yellowstone long periods of time provide quantitative documentation of change in
National Park. (a) 1958 data form showing mapping of aerial extent of ecosystems. (Historic photo and data from Denton and Kittams 1958;
sagebrush (Artemisia spp.) present, height of plants, seedlings, and dead 2008 photo courtesy of Art Sikkink. Reprinted under CC-BY-4.0 Inter-
shrubs; (b) 2008 data form showing mapping of the aerial extent of national, from Sikkink, P.G., 2011. Yellowstone Sage Belts 1958 to
sagebrush and other shrubs, species present, height of plants, seedlings, 2008: 50 Years of Change in the Big Sagebrush (Artemisia tridentata)
and dead shrubs; (c) 1958 photo of belt transect corresponding to 1958 Communities of Yellowstone National Park. Natural Resources and
sample form; and (d) 2008 photo corresponding to 2008 sample form. Environmental Issues 17, 16. # 2011 Utah State University)
Regularly collected data like that obtained from these transects over
sampling. Today one widely used approach to generate a sequence for sampling grid points. Through randomization
probability design ensuring spatial balance is that of of addresses, systematic sampling now results in a spatially
generalized random tessellation stratified (GRTS) samples. well-balanced random sample in which the probability of
Using a GRTS approach, investigators first create grids at selecting any particular address can be determined (Stevens
different spatial scales (“hierarchies”) and assign an and Olsen 2004).
“address” to each grid point as a way of uniquely designating Four general levels of analysis that typically occur in a
it. Addresses are then randomly sorted to determine a well-planned monitoring program include (1) descriptive and
Moderate grazing
Cedar/Juniper
Cedar/Juniper
Woodland or
Establishment
Savanna
Conversion to
, ti g,
me
Li esc
agricultural lands may
fire nin
pr
gh rib
occur directly from
ed hin
t g ed
Mo /low
no
native grasslands or any
ra
rib l t
de int
zin fire
sc ica
vegetative state
ra en
g,
te si
pre chan
gr ty
Indicates little or no possibility
az fir
Me
of restoration to previous state
in e
g,
o fire
zing, n
No gra
- Lowered diversity
Native Grassland and ANPP
Abundant annuals, Herbicide, planting, and time
Community - Loss of fire
cacti, and bare soils
Composition species
- High fuel loads
Grazing followed by fire

fire estin ing,
ble
z
po g,
gra
ssi
ht
fire g,
Pla
Lig
r
no razin
ntin
g
re
avy
g, t
ic fi
Restored
ime
He
oph
Grasslands
He no
fir
av t p
astr
e
y g os
Cat
ra sib
zin le
g,
- Deneutered soils
Increases
- Loss of plant
annuals
cover
(Opuntia sp.)
- Eroded soils
Fig. 9.21 An example of a conceptual model developed by the US of a grassland community to agricultural land. (Reprinted by permission
National Park Service (NPS) Southern Plains Inventory and Monitoring from Cambridge University Press, from S.G., Bennetts, R.E., 2012.
Network for managing plant species composition of NPS units in the Institutionalizing an effective long-term monitoring program in the US
NPS’s Southern Great Plains region of the United States. Note the National Park Service, in: Gitzen, R.A., Millspaugh, J.J., Cooper, A.B.,
model’s three potential pathways for changing the species composition Licht, D.S. (Eds.), Design and Analysis of Long-Term Ecological Mon-
using the interactions of fire and grazing, two manageable forms of itoring Studies. Cambridge University Press, Cambridge, pp. 481–497.
ecosystem disturbance. A fourth pathway (upper left) depicts conversion Modified by K. DeVoss)
summary statistics; (2) determination of the current status of they have made about the ecosystem and about cause and
the resource being measured; (3) identification of changes or effect relationships between management actions and ecosys-
trends in the resource over time (Fig. 9.22); and (4) synthesis tem responses (Manley et al. 2000).
of status and trends in one resource with other resources to
identify possible changes in overall ecosystem structure and 9.3.4.3 Ecosystem Management and Geographic
function (Fancy and Bennetts 2012). Such analysis is critical Information Systems – How Technology
to crafting effective decisions and management actions. In Enables Management Purpose
fact, one of the most important functions of ecosystem moni- and Strategy
toring is to determine the success of specific management In a practical sense, EBM was not possible prior to the
actions toward prescribed objectives based on the achieve- development of remote sensing, whereby airplanes flying
ment of those objectives. Thus, effective monitoring informs across a large spatial domain or satellites orbiting the Earth
managers (and modelers) about the validity of assumptions could provide information on a multitude of environmental
Watershed tour and Microbial source

10,000 Persistent summertime peaks
public meeting tracking study
above water quality standards
initiated
trigger notification and
monitoring frequency
Most Probable Number (MPN) 1,000
100
First observed
exceedance at
10 downstream end of
Narrows. MPN below
standards next day.
1 Outhouse
construction begins
upstream from park.
0.1
4/ 06
7/ /06
10 /06
2/ /06
5/ 07
8/ /07
12 /07
3/ /07
6/ /08
9/ /08
1/ 08
4/ 09
7/ /09
11 /09
2/ 09
5/ 10
8/ /10
0
/1
1/
5/
/
5/
/
9/
11
20
16
24
/2
11
19
27
15
24
/1
20
28
1/
/2
2A chronic 2A acute Pasture Sites Non-Pasture Sites
Fig. 9.22 Estimates of population levels (y axis, Most Probable Num- initiation of a study to track the source of increased E. coli. (Reprinted
ber (MPN)) of the bacterium Escherichia coli at monitoring sites on the by permission from Cambridge University Press, from Fancy, S.G.,
North Fork of the Virgin River, Utah, USA. Horizontal lines indicate Bennetts, R.E., 2012. Institutionalizing an effective long-term monitor-
water quality standards (“exceedance thresholds”) in relation to poten- ing program in the US National Park Service, in: Gitzen, R.A.,
tial dangers to human health. E. coli levels between 100 and 1000 MPN Millspaugh, J.J., Cooper, A.B., Licht, D.S. (Eds.), Design and Analysis
trigger public notification and increased monitoring. Levels significantly of Long-Term Ecological Monitoring Studies. Cambridge University
above 1000 MPN trigger a public meeting and watershed tour and Press, Cambridge, pp. 481–497. Modified by K. DeVoss)
variables. The longest continous space-based source of imag- (2) determining and displaying union and intersection of
ery for analysis of biological data has been collected via the different geographic and biological variables through virtual,
joint NASA/USGS Landsat Program. Every day, computer generated overlay maps; (3) organizing and
Landsat satellites capture high-resolution images and displaying distribution of geographic, climatic, or biologic
are arranged in orbits to systematically photograph all parts patterns at large scales (e.g., regional, continental, or world),
of the Earth’s surface at regular intervals (Fig. 9.23). When (4) converting raw remote sensing data to digital information
these photos are enhanced and digitized, or when high reso- that can be redisplayed as an interpretive map using pre-
lution color and multi-spectral satellite imagery is processed, scribed classification procedures; (5) performing statistical
they yield vast amounts of information, especially when analyses of geographic information; and (6) providing “deci-
integrated through Geographic Information System (GIS) sion support systems” that use pre-programmed decision
technologies, sometimes also referred to as Geographic Infor- rules to assist managers in making decisions about land use
mation Science. or resource allocation, visually represented by “multi-criteria
A GIS is a framework for gathering, managing, analyzing, suitability maps” that can indicate the best management
and visualizing different types of spatial data, including data practices on particular land areas, such as the kind used by
layers developed from remotely sensed data. Often The Nature Conservancy for aquatic ecosystem conservation
individuals use software platforms such as ArcGIS or QGIS as described in Chap. 9.
to solve complex problems in spatial analysis. Among the To display such unions and intersections, GIS programs
most common and important uses of GIS are: (1) creating can construct maps of the same area for different variables
data management systems for geographic information that (for example soils, vegetation, and elevation), and then create
allow users to enter “attribute data” (e.g., elevation, soil a single virtual overlay map that shows the union and inter-
type, vegetative cover, land use, and other variables) into section of different variables (“attributes”) with one another
files that can then be manipulated to display such data in (Fig. 9.24). Using such a map, potential habitats for species
new ways that are geographically or spatially sensitive; whose environmental tolerances require a union of multiple
Fig. 9.23 Satellite photograph of Yellowstone National Park, characteristics on local, regional, or global scales. (Landsat imagery
Wyoming (USA) taken on July 22, 1988, illustrating the type of courtesy of NASA Goddard Space Flight Center and US Geological
remotely sensed data that can be used by a geographic information Survey)
system (GIS) to analyze ecosystem, landscape and habitat
variable states (for example, vegetation, elevation, and mois-

ture) can then be identified, as they were by Rondinini et al. Point of Engagement Question
(2005) to develop range maps for African vertebrates Ecosystem management developed partly in response
(Chap. 7). to the increasing complexity of environmental
Satellite images or aerial photographs can be used to problems and to shifting public attitudes toward the
identify vegetation or land-cover types based on the spectra value of ecosystems. Was its development also
they emit. From this information, an appropriately facilitated by development of remote sensing and GIS
constructed GIS program can determine the proportion and technologies occurring at about the same time? How, in
quantity of areas associated with different land-use practices your view, did technological opportunity, management
or habitat types, a first step in ecosystem inventory. Recall need, and public sentiment interact to facilitate devel-
that the Gap Analysis Program (GAP, Chap. 7) is employed opment of the ecosystem management concept? Pro-
in biodiversity protection by integrating maps of natural vide an example.
vegetation with distributions of species, as well as with land
use jurisdiction and practices. For ecosystem management,
9.3.4.4 Archived Data and Historical
this application can be both practical and specific. For exam-
“Experiments”
ple, by creating an overlay map showing union and intersec-
Another important long-term comparison and evaluation is
tion of land stewardship categories and ecosystem types
the examination of archived data available from past studies
integrated with rare and endangered species distributions,
of ecosystems. In countries where all or part of a major
GIS can be used to determine the area of each ecosystem
ecosystem have a long history of public ownership and
type associated with each stewardship category and likely
agency jurisdiction, there may also be a long history of
impacts of land use practices on threatened species (e.g.,
ecological investigation. Many such investigations take the
Shinneman et al. 2000).
form of unpublished reports, unprocessed data in agency
Fig. 9.24 Diagrammatic representation of sequential steps commonly often employed in developing habitat suitability models (HSMs) as
employed in a Geographic Information Systems (GIS) analysis. In this described for species like the giant panda (Ailuropoda melanoleuca) in
example, environmental characteristics (attributes) are reclassified rela- Chap. 7. (Map images are the intellectual property of Esri and is used
tive to a predetermined scale of environmental suitability (for example, herein with permission. Copyright # 2019 Esri and its licensors. All
habitat suitability for a species of conservation concern) and then given rights reserved)
area-specific (“cell”) weights based on suitability scores, a technique
files, personal journals, historical photos, herbarium records compiled over hundreds or thousands of years. Such
that document location and occurrence of plant species, soil sediments are repositories of pollen grains dispersed annually
surveys, and geologic maps, past aerial photos, as well as by plants. Pollen grains are resistant to decay, especially in
peer-reviewed published literature. Archived information conditions associated with bog sediments, and distinctive by
will vary in quality and require extra effort in analysis and species. Their proportional abundance in sediment strata,
interpretation, but such data can yield unique and valuable appropriately corrected for differences in pollen production
insights. Effort employed in search and interpretation of past by different species, can provide an index of proportional
data and records (often referred to as data mining or mining abundance of plant species around the lake or bog. Properly
of legacy data) is therefore essential for ecosystem inventory, extracted, a core of sediment can provide a “profile” of the
as well as for establishing ecosystem baselines for compari- abundance of different species of pollen deposited over time.
son to current conditions (Fig. 9.25). As abundance of pollen of different species changes, such
changes can permit a manager to make inferences about
9.3.4.5 Data from Long-Term Natural Repositories changes in the ecosystem, and even estimate the time period
and Preserved Areas associated with such changes if the rate of sediment deposi-
Not all archives are found in file cabinets, field journals, or tion can be estimated. Notice, for example, how relative
old photo albums. An ecosystem keeps its own data, if one abundance of different pollen types changes in the sediments
knows where to look. Lake and bog sediments, for example, of a Panama lake (Fig. 9.26). Sediments also keep records of
provide long-term records of biological change in ecosystems material that precipitates from surrounding water and air.
Fig. 9.25 Effects of a fire on sagebrush (Artemisia spp.) distribution in collection, 2008 photo courtesy of Art Sikkink. Reprinted under CC-
Yellowstone National Park (USA). (a) Sagebrush growth prior to the fire BY-4.0 International, from Sikkink, P.G., 2011. Yellowstone Sage Belts
in 1988; (b) Young sagebrush regrowth at the same site 6 years after the 1958 to 2008: 50 Years of Change in the Big Sagebrush (Artemisia
fire in 1994; (c) Mature sagebrush at same site 14 years later (2008); and tridentata) Communities of Yellowstone National Park. Natural
(d) sagebrush growth on an adjacent site in the same year (2008). Resources and Environmental Issues 17, 16. # 2011 Utah State
Analysis of historical photos of the same sites can provide important University
insights on ecosystem change over time. (Photos from YNP archive
Two such precipitates are ash and charcoal, substances pro- pollution, water pollution, or invasive species may affect all
duced by fires. From the abundance and depth of these sites regardless of their “protected” status. Multiple types and
materials in lake and bog sediments, one can infer the relative levels of disturbances may affect unprotected sites, so
frequency of fire in the surrounding ecosystem over an differences between them and protected areas are unlikely
extended period of time, and whether that is similar to or to be traceable to a single cause. And protected, undisturbed
different from current fire frequencies. sites are often smaller than surrounding unprotected areas.
A special category of “long-term natural repositories” is Ecological structure and function may be adversely affected
sometimes found in relatively undisturbed areas, which can by fragmentation and edge influences (Chap. 7), which are
provide baseline information used to compare responses in more pronounced in smaller reserves.
systems stressed by varying kinds of disturbance. Nature
preserves or other protected areas can serve as controls or
“before impact” sites when evaluating ecosystem status and 9.4 Implementing Management Decisions –
change; what some ecologists call “time-control substitutes” Tools of Ecosystem Management
(Loehle 1991). Ecosystem structure and function in these
sites can be compared to otherwise analogous disturbed 9.4.1 Ecosystem Modeling
sites to determine how the system has changed over time in
response to disturbances that affect it. Such comparisons can Modeling of ecosystems is critical for management, even if
be helpful but require careful interpretation and many models fail to make perfectly accurate predictions about
qualifiers. Regional sources of disturbance, such as air ecosystem behavior (recall the model used (Fig. 9.12) in
9.4 Implementing Management Decisions –Tools of Ecosystem Management 389
Pollen and charcoal in Pollen and spores of plants

Lake Wodehouse indicate a associated with disturbance
human presence and increase after 3,900 BP.
agriculture around the lake
) 2)
beginning after 3,900 BP. Corn pollen appears in Particulate carbon also
r e o.
po N
sediments after 3,900 BP. increases substantially
)
re
(2 re
po
e o
after 3,900 BP.
p
(3
ea (3
e
ac e
ea
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) the
pi M ce
/M e
ac
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BP e
Particulate carbon
or
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20 (%) 50 (%) 50 (%) 20 40 10

Lake Wodehouse Total pollen (%) Concentration
sediment core (104 items/cm3)
Fig. 9.26 Pollen and particulate carbon in sediments of Lake Colinvaux, P.A., 1994. Tropical Forest Disturbance: Paleoecological
Wodehouse, Panama, dating to 3900 years before the present Records from Darien, Panama. Ecology 75, 1761–1768. # 1994 Eco-
(BP). (Reprinted by permission from Wiley, from Bush, M.B., logical Society of America)
our discussion of DPSIR). Ecosystem models take a variety defined in the model by a unique set of interactive processes,
of forms. A conceptual model is a visual or narrative sum- components, and structures. Level Two, the “Sphere Model,”
mary that describes or identifies important components of a identified the key processes in each of the five spheres
system and possible interactions among them. Conceptual responsible for transferring material, energy and information.
models can illustrate interconnectedness of ecological pro- Level Three, the “Processes Model,” depicted mechanics of
cesses, both as they occur in nature and as they respond to each of the processes operating in the individual sphere
anthropogenic influences. They can identify how major models. The depiction of the processes consisted of two
drivers and stressors will affect ecosystem components and parts – the essential elements and outcomes of each process
provide a framework for communication among scientists and the affectors that influenced the process. Manley et al.
and managers from diverse disciplines (Allen and Hoekstra defined affectors as “actions, consisting primarily of human
1992). The latter function is especially important in avoiding activities that generate a change in the value of state variables
the disconnect common between managers and researchers (e.g., elements, outcomes, or process operation)” (Manley
illustrated in our earlier example of the management of the et al. 2000:144). Affectors are more or less synonymous
swamp wallaby in Australian forests. with what have earlier in the chapter been described as
Conceptual models play important roles in the previously stressors if they have negative effects, or, alternatively,
discussed work of ecosystem monitoring. Patricia Manley of subsidies (to the ecosystem outputs and services) if their
the US Forest Service and her colleagues created a concep- effects are positive.
tual model of ecosystem processes in the Sierra Nevada Although models differ in detail and purpose, managers
Mountains of the US state of California organized in three must define values of five types of entities to build a working
model “levels,” the Ecosystem Model, the Sphere Model, and model. Stocks are amounts or levels of a variable of interest
the Processes Model. Level One, the overall “Ecosystem that the model counts or monitors. Sources and sinks are
Model,” consisted of five “spheres” including, as one would entities from which the stock originates (sources) or into
expect, the atmosphere, hydrosphere, biosphere, and litho- which the stock is absorbed (sinks). Flows, usually expressed
sphere (soil), as well as the “sociocultural sphere,” making as equations, determine rates of movement of stocks to and
humans and their activities an explicit and foundational com- from sources and sinks or from one stock to another (e.g., the
ponent of the model. The each of the five spheres were conversion of plant biomass into animal biomass).
Parameters or converters are values of variables used to ecosystem components, values of the parameters, and paths
determine rates of flow. Connectors show the path through through which the stocks are transferred, then, components of
which material is transferred from one stock to another, or to interest in the ecosystem can be modeled. You can see how
and from sources and sinks. If a manager can accurately these various components interact in a simple, but real exam-
determine initial value of pertinent stocks, the conceptual ple of ecosystem interactions in Information Box 9.2.
framework (often, equations) that regulate change in stocks
or the rate at which stocks move or are converted to other
Information Box 9.2: Stocks and Flows in Limpets Information Box 9.2 (continued)
and Seaweed the equations) are sufficient to track changes in their
As a first step in understanding the basics of an ecosys- populations (represented in the model as biomass) and
tem model, take a classic example of a model system other variables which affect them, which you can track
comprised of two stocks, populations of seaweed and in the illustration as “runs” in the model (SIM1 –
their primary herbivore, limpets (a gastropod mollusk). SIM4). Boxes (L and S) represent populations (bio-
An estimation of initial levels of these stocks, knowl- mass or “stocks”) of limpets and seaweed in the sys-
edge of values of six parameters (reproduction rates of tem. K1 – K5 and B are parameters (constants)
limpets and seaweeds, effect of limpet feeding on sea- affecting rates of flow (dL/dt and dS/dt) between stocks
weed, limpet death rates, and density dependent and ecological sinks (clouds) (Brennan et al. 1970;
constraints on limpet and seaweed population growth) Information Box Fig. 9.2).
and an understanding of connections between parame-
ter values and rates of change in stocks (expressed in
Information Box Fig. 9.2 Conceptual illustration, equations, parameter values, and simulation results (SIM1-SIM4) of a model of limpet
herbivory on seaweed. (Equations and parameters derived from Brennan et al. 1970. Drawing by J. D. Schmeling, modified by K. DeVoss)
(continued)
Models that attempt to address all components of an heterogeneity. Large, contiguous fires make landscapes
ecosystem are more complex than this example. However, more homogeneous, particularly if the fire is hot and spreads
models that make predictions about single effects or states rapidly.
may be much simpler. For example, in the Serengeti, Ecosystem managers can use fire to maintain or enhance
Wolanski et al. (1999), after conducting extensive field stud- biodiversity, increase habitat heterogeneity, increase plant
ies, hypothesized that water quality and quantity were the nutrient uptake, create conditions attractive to particular spe-
dominant forces driving ecological events. Background stud- cies or other desired outcomes on a site-specific basis, or
ies established that migration of ungulates (hooved permit naturally-ignited fires to burn over wider areas to
mammals) was related to level of rainfall and river flows, create such effects throughout the ecosystem. In managing
but these variables still did not predict animal movements fire, managers can basically manipulate four variables. The
with precision. Further investigations revealed that water first is the type of fire to be used. For example, managers
quality, specifically salinity, was the most accurate predictor could ignite a surface or a canopy fire in a forest, with each
of ungulate movements. Where water was fresh, wildlife type producing radically different risks and results. The sec-
remained. Where it became saline, they left. And variation ond manageable variable is fire intensity. Managers can
in salinity by decade was the most important factor determin- affect amount of heat generated by a fire by their choice of
ing discontinuity between grasslands and wooded savannas. site, fuel types and accumulations that are present, and envi-
Modelers, using the value of average salinity from a single ronmental conditions under which the fire is set. Third,
lake, were able to predict movement of animals from managers can manipulate fire frequency. For example, in
grasslands to woodlands within 1 week when salinity values North America, a tallgrass prairie burned every year will
rose above a pre-determined threshold (Wolanski et al. have a different species composition and physical structure
1999). than one burned every 3 years or every 5 years. Management
Modeling and monitoring are connected because decisions about fire frequency will have important short-term
modeling identifies critical variables that are the best indices effects on species composition on burned sites, and poten-
of changes in status or function of an ecosystem, and so make tially long-term effects on landscape composition at larger
monitoring efforts more efficient and cost effective. By scales. Finally, managers can alter fire timing, usually
knowing what to sample and monitor, managers can reduce through their choice of which season to set the fire. Again
waste of time and money spent monitoring variables that are to use North American tallgrass prairies as an example, early
not sensitive (or less sensitive) to ecosystem processes. spring fires ignited before germination of new, living plant
A model can tell a manager what processes are important biomass favor establishment of a relatively small number of
to the ecosystem’s structure, function, and production, but the “warm season” grass species such as big bluestem
model itself does not manage anything. It remains for the (Andropogon gerardii), Indian grass (Sorghastrum nutans),
manager to take action in manipulating ecosystem processes and switchgrass (Panicum virgatum) because the resulting
to achieve predetermined objectives. What ecosystem pro- bare, blackened soil readily absorbs heat, increasing soil
cesses are amenable to management? temperatures that favor germination of these and other spe-
cies of grasses that germinate best in warmer soils. Summer
burns, which burn living plants with higher moisture content
9.4.2 Managing Ecosystem Processes (and therefore burn less completely and intensely), leave
more plant material on the soil, reducing post-fire soil
9.4.2.1 Fire temperatures and favoring establishment of more diverse
Ecological effects of fire vary, but in most terrestrial communities of “cool season” grasses and forbs.
ecosystems, fire can lead to increases in: (1) habitat heteroge- Even if fire is allowed to burn without suppression
neity and amount of edge; (2) plant and animal diversity; throughout the system, it is unlikely to produce a stable
(3) nutrient uptake by plants, especially graminoids and pattern of landscape structure or to necessarily replicate pre-
forbs; (4) loss of nutrients from soil; (5) rates of erosion and vious historical landscape patterns. Although site-specific
surface runoff; (6) rates of streamflow; (7) grazing and application of fire can be beneficial in fragmented habitats,
browsing of burned areas by ungulates; and (8) establishment regional or ecosystem-wide use of prescribed fire is unlikely
of early successional species (Knight and Wallace 1989, in the foreseeable future because of the lack of appropriate
Leach and Givnish 1996, Van Dyke and Darragh 2006). All technology, insufficient financial and human resources in any
but the first of these effects are usually of short duration, one agency to control large-scale fires, and strong public and
generally lasting only 1–3 years. In contrast, increases in political opposition to large-scale fires as a management
habitat heterogeneity may be a long-term effect, although technique. Even the choice to allow fires to burn in relatively
magnitude of effect varies with scale and intensity of the uninhabited ecosystems is problematic because large fires
fire. Small, dispersed fires tend to increase habitat can spread to inhabited areas, resulting in loss of human life
and property, and cannot be effectively controlled if they associated landforms, and sizes of individual trees. Infrequent
reach certain critical sizes. Although fire has been a powerful but large-scale flooding can shift a population of trees from
force in shaping ecosystems, its current applications in man- an uneven-aged stand to an even-aged one because the imme-
agement will likely remain limited to individual, relatively diate post-flood period synchronizes germination of new
small sites. individuals in the flooded area. Floods also move downed
trees, limbs, and brush into stream channels, increasing chan-
9.4.2.2 Water Flow nel structure, surface area, and roughness, promoting sedi-
ment retention, and increasing stability of the channel
Direct Manipulation surface. Such woody biomass in streams increases inverte-
Today most wetland and riparian systems are managed on a brate activity, provides cover for fishes, and increases habitat
site-specific basis, but processes that control them –especially diversity of the stream channel (Michener et al. 1998).
water flow – extend over vast areas. As we saw in Chap. 9, Most stream species are adapted to flow variations and
flow management and regulation are among the most impor- decline in abundance if variation is reduced or eliminated
tant tools used in stream habitat conservation today. Their use (Sparks 1995). Such species may use variations in flow as
becomes even more important when conservation efforts cues for breeding and feeding activity because optimal
shift from habitats to ecosystems, especially stream conditions for both are different and do not usually occur at
ecosystems. the same time. If variations in flow cease, breeding and
Most lake systems remain relatively constant in water feeding activities may be curtailed or aborted. If variations
volume and flow, but streams experience seasonal flow in flow cease, breeding and feeding activities may be
variations that significantly affect stream ecosystems and curtailed or aborted, as in species like the black stilt, least
associated terrestrial ecosystems. Variation in water flow is tern and piping plover (Chap. 8).
one of the most important determinants of ecosystem struc- In terms of human safety, property, and economics, floods
ture and function. Changes in water flow promote exchanges are negative events. Thus, managers have tried to control or
of nutrients among habitats and enhance system productivity. eliminate flooding through impoundments that reduce
A flood pulse also can provide a dimension of “seasonality” fluctuations in flow; to construct levees and other physical
to environments that are otherwise unseasonal, such as tropi- structures that reduce the size of the floodplain; and, on
cal rainforests, making them more productive and diverse larger streams, to physically remove downed timber and
(Sparks 1995). Systems that experience flow variations other woody debris from stream channels to increase ease,
include some of the most species-rich places on earth, such speed, and safety of navigation. In a context of ecosystem
as the Amazon rainforest, the papyrus marshes of the Nile, management, these are legitimate concerns. However,
the swamps of the Okavango River in Botswana, and the if floods are eliminated, flood-adapted components of the
shallow wetlands and lakes of the Gran Pantanal of the system will decline because flood pulses are the primary
Paraguay River in South America. These areas also have source of energy in many freshwater aquatic systems,
traditionally supported important commercial fisheries facilitating creation of habitat structure, nutrient exchange,
(Welcomme 1985). and organism movement (Sparks 1995). Flooding also
In stream ecosystems, natural flooding over a historical renews floodplain soil by depositing sediment and nutrients
floodplain increases habitat heterogeneity during floods otherwise lost through erosion, depleted by crops, or
because differences in topography in the floodplain, even sequestered in soil.
small depressions, will hold water longer and at greater If flow is restricted to a narrower channel such that the
depths, providing habitat for waterfowl, amphibians, and floodplain cannot receive and store the floodwaters, flood
other terrestrial species that use ephemerally flooded areas. heights and damages will increase at other locations. It is
The increasing use of “laser leveling” (establishing a single noteworthy that many dams on major rivers were built to aid
slope across a field to provide irrigation to crops from navigation by maintaining river depth at a sufficiently high-
floodwaters) eliminates small depressions and associated level during periods of lowest flow, not to stop floods (Sparks
habitat heterogeneity, with predictable declines in species 1995). Under an ecosystem management approach, managers
diversity. would not only manipulate flows of smaller streams in local
Flooding lowers total biomass production over the short watersheds, but larger dams also could be removed to restore
term but does not always produce long-term effects. In the natural flow regimes in larger watershed systems. Such
southeastern US, large, infrequent floods regulate the devel- possibilities are no longer hypothetical. As noted in
opment of longleaf pine forests through differential mortality Chap. 8, hydroelectric dams have been destroyed in Maine
via complex interactions of forest landscape position, (USA) to restore migration of anadromous fish species, and
the removal of several dams in the US Pacific Northwest is applications, but as tools in comprehensive ecosystem-
now being considered. based management approaches that affect basic ecological
processes, including the ability of vegetation to influence
Indirect Manipulation: Using Vegetation Management surface runoff and soil absorption of water to reduce the
to Manage Water Flow negative effects of stream flow changes induced by climate
Faced with the reality of increasing temperatures as well as an change events.
increasing frequency of extreme wet and dry years in the
southern Appalachian forests of the United States, US Forest
Service (USFS) Research Project Leader and Ecologist Chelcy 9.4.3 Managing Nature’s Ecosystem Engineers:
Ford Miniat (formerly Chelcy Ford) and her colleagues at the Herbivores and Herbivory
USFS Southern Research Station examined the possibility of
managing land use to affect water flow by manipulating forest Ecosystem engineers, when the term is applied to non-human
plantings. Ford Miniat and her colleagues explain, “Many tree species, are species “whose presence or activity alters its
species differ considerably in the amount of annual and sea- physical surroundings or changes the flow of resources,
sonal water use via transpiration and interception. . .For exam- thereby creating or modifying habitats and influencing all
ple, in some geographic areas with mild winters, a shift from associated species” (Crain and Bertness 2006). One obvious
deciduous to evergreen forests is likely to result in greater forest example is the beaver (Castor canadensis) whose activities in
water use due to greater year-round transpiration and intercep- building dams to impede the flow of water in streams creates
tion. . .Forests are unique among land uses because they are ponds immediately upstream of the dam, radically changing
long-lived, relatively stable, and respond to climate through ET the stream’s habitat.
[evapotranspiration]; yet their structure and function can be Other ecosystem engineer species may be less obvious,
substantially altered by forest management. . .” (Ford et al. but more pervasive in their effects. Large herbivores can
2011:2050). exert profound controlling influences on ecosystem
Ford Miniat’s team set out to identify if and by how much components, structure and function. In many cases, ecosys-
land use could affect levels of streamflow for any given tem management is impossible without herbivore manage-
amount of precipitation, then identify management treatments ment, and managers who fail to manage the ecosystem’s
that would mitigate against extremes in annual precipitation. herbivores will often find that the herbivores will manage
They determined that forest management could affect stream the ecosystem for them, and not always to their intended
flow and mitigate effects of climate warming and associated ends. Herbivores affect ecosystem processes primarily
climate extremes (Chap 4). Management strategies employed through regulation of habitat, regulation of energy flow,
caused streamflow to increase, although effects were tempo- regulation of plant nutrient cycling, and effects on plant
rary. Different management treatments produced different nutrition. Both browsers (herbivores feeding on woody veg-
stream flow responses. “. . .For example,” noted Ford et al., etation) and grazers (herbivores that feed on herbaceous
“compared to an unmanaged catchment, managing a catch- vegetation) achieve these results, although not always
ment with a species conversion treatment decreased the appar- through the same means.
ent frequency of observed extreme wet-event years by a factor Browsers often prevent vertical development of sapling
of seven . . .The choice of species in this type of management vegetation of preferred food species, contributing to spread-
treatment is a critical determinant of the magnitude of ing, shrubby growth forms that keep the affected plant bio-
response. . .” (Ford et al. 2011: 2064–2065). mass within their reach (McNaughton et al. 1988). In
Response of streamflow in experimentally treated northern boreal forests in Europe and North America,
watersheds was extremely predictable (Fig. 9.27), indicating moose prevent saplings of preferred species from growing
that forest management could be an effective method of into the tree canopy, resulting in a forest with fewer canopy
mitigating changes in stream flow associated with climate trees (consisting mostly of non-preferred species) and a well-
change. Overall, Ford et al. concluded, “We found that developed understory of shrubs and herbs, all within reach of
streamflow response to climate was affected by nearly all moose (McInnes et al. 1992)! In fact, light to moderate
land uses examined, supporting our hypothesis that climate browsing leads to increased production efficiencies (higher
impacts may either be mitigated or exacerbated by forest rates of production per biomass) in shrubs and saplings that
management practices” (Ford et al. 2011: 2062). are browsed (Fig. 9.28). Through such browsing, moose also
Here we have a clear example of how long-standing, affect quantity and quality of litter and soil nutrients, driving
traditional forest management practices (vegetation manage- a set of ecological interactions between browse, litter quality,
ment), especially management of tree species of commercial and soil nutrients (Fig. 9.29) (McInnes et al.1992). Similar
value in managed forests, can be creatively adapted and effects are seen in mixed deciduous-coniferous forests, where
employed, not in their traditional “resource management” moose typically browse preferentially on deciduous
Fig. 9.27 Predicted streamflow

in watersheds subject to different
forest management treatments
(Q^ T ) compared to observed
streamflow (QT). In (a) (Species
conversion (S)), the treated area
was subjected to a prescribed
burn. In (b) (Species conversion
(N)), all woody vegetation in the
treated area was cut in a single
year. In (e) (Coppice), “coppice”
refers to a management treatment
in which woody vegetation is
periodically cut back to ground
level to stimulate growth, often
with the objective of providing
firewood or timber. (Reprinted by
Ford, C.R., Laseter, S.H., Swank,
W.T., Vose, J.M., 2011. Can
forest management be used to
sustain water-based ecosystem
services in the face of climate
change? Ecological Applications
21, 2049–2067. # 2011
Ecological Society of America)
hardwoods. This feeding pattern not only changes forest Like browsers, grazing species affect a variety of
composition, but reduces nitrogen mineralization, nitrogen components of ecosystem structure and function through
inputs, and primary productivity because the browsing their effects on vegetation. In some systems, grazers can
reduces quantity and quality of litter returned to the soil remove up to 40% of standing biomass, significantly reduc-
(Pastor et al. 1993). In Africa, mixed feeders, such as ing ecosystem production. In other systems, grazers may
elephants (Loxodontai africana), that combine browsing initiate changes in plant morphology and physiology that
and grazing, regulate abundance of woody browse and under- lead to higher levels of plant productivity. For example, in
lying grasses, often by pushing over trees or stripping their African grazed systems, dominant grasses are dwarfed, low
bark, reducing abundance of woody vegetation and creating growing forms with short internodes, while in ungrazed
openings grasses then invade (McNaughton et al. 1988). In systems in the same area dominant grasses are tall growing
North America, elk can, at high densities, suppress height and species (McNaughton et al. 1988). When grazed, many plants
survivorship of quaking aspen (Populus tremuloides) and respond by increasing biomass concentration in their tissues
willow (Salix spp.) as well as many conifer species through (the ratio of mass to volume, often measured in mg of plant
browsing (Kay and Wagner 1994; Romme et al. 1995). biomass per cm3), creating more “biomass per bite” for
Table 9.7 A summary of important effects of herbivory on plant

growth and metabolism
Effects of herbivory on plants
1. Photosynthetic rates increase in the remaining tissue.
2. Older tissues, functioning at levels below maximum photosynthetic
level, are removed.
3. The active photosynthetic period of residual tissue is prolonged as
the rate of leaf senescence is reduced.
4. Substrates are circulated through the plant.
5. Removal of overshadowing tissue intensifies light on potentially
more active underlying tissues.
6. Increased leaf growth and tillering result from the division and
elongation of cells; the activation of remaining meristems increase due
to the plant’s hormonal response; and growth is also promoted by
Fig. 9.28 The relationship of browsing to production efficiency. Note chemicals in ruminant saliva.
that light-to-moderate browsing by moose stimulates higher ratios of 7. Transpiration surface is reduced. Consequently, soil moisture
plant production per unit of biomass in shrubs. (Reprinted by permission conservation increases.
from Springer Nature, from McInnes, P.F., Naiman, R.J., Pastor, J.,
8. Nutrients are recycled from dung and urine.
Cohen, Y., 1992. Effects of Moose Browsing on Vegetation and Litter
of the Boreal Forest, Isle Royale, Michigan, USA. Ecology 73, 2059– Design by M. J. Bigelow based on concepts from McNaughton 1979
2075. # 1992 Ecological Society of America. Modified by M. J.
Bigelow)
Fig. 9.29 Relationship between levels of moose browsing, preferred

and unpreferred species of browse, litter quality, and soil nutrients.
Arrows indicate direction of interaction. + signs indicate positive effects.
Minus signs indicate negative effects. (Reprinted by permission from
Springer Nature, from McInnes, P.F., Naiman, R.J., Pastor, J., Cohen,
Y., 1992. Effects of Moose Browsing on Vegetation and Litter of the
Boreal Forest, Isle Royale, Michigan, USA. Ecology 73, 2059–2075. #
1992 Ecological Society of America. Modified by M. J. Bigelow)
Fig. 9.30 Response of plant productivity to grazing by African large
mammals. Grazing produces a response function of increased produc-
herbivores. Gregarious herbivores exploit this response in tivity (g/m2/day) in grazed vegetation, with greatest increases associated
plants by actively creating “grazing lawns”- intensively with intermediate levels of grazing intensity, 1 – g/ng, where g is the
grazed areas within the ecosystem where the herbivores’ biomass in grazed areas unprotected by fencing and ng is the biomass in
a permanent exclosure in which no grazing occurs. Grazing as an
own grazing activity keeps plant heights low and biomass optimization process: grass-ungulate relationships in the Serengeti.
concentration high. Although this reduces total plant biomass (Reprinted by permission from University of Chicago Press Journals,
density in the grazing lawn compared to ungrazed areas, it from McNaughton, S.J., 1979. Grazing as an Optimization Process:
elevates foraging efficiency because of increases generated in Grass-Ungulate Relationships in the Serengeti. The American Naturalist
113, 691–703. # 1979 The University of Chicago. Modified by M. J.
biomass concentration (McNaughton 1984). Grazing also Bigelow)
tends to raise photosynthetic rates in plants, increase rates
of nutrient allocation to growing plant tissues, increase
growth rates in plants, and produce other effects that often grazed systems in the Serengeti-Mara regions of Tanzania
benefit herbivores (Table 9.7). Light to moderate grazing by and Kenya, McNaughton (1979) determined that over
native herbivores produces positive responses in plants in two-thirds (r2 ¼ 0.69) of variations in plant productivity
growth rates, metabolic efficiency, and nutrient could be explained by grazing intensity alone (Fig. 9.30),
concentrations (McNaughton 1979). In fact, in one of the producing a response like that of browsing as previously
earliest but most well-designed experimental analysis of discussed (Fig. 9.28).
9.5 Creating and Managing Governance roundtable discussion. This vision must not only incorporate
Systems of Ecosystem-Based ecological science, but also social, economic, and
Management community-based perspectives. Nevertheless, science
remains the foundation of EBM because it gives all
9.5.1 Managing Through Collaboration participants shared understanding of how the ecosystem actu-
with Stakeholders ally functions. To see how science can be successfully
integrated with and act to inform stakeholder interests, con-
As we have noted earlier, EBM, compared to traditional sider an example from northeast Australia.
resource management, not only requires a different frame-
work of management actions to be effective, but a different
kind of managing process and managing governance. 9.5.2 Linking Interest with Identification:
Because EBM involves management of ecological processes Australia’s Tully-Murray Watershed
over large landscape scales that routinely exceed jurisdictions
of individual agencies, management decisions must incorpo- As we saw earlier in solving the problem of locating and
rate decision-making strategies that involve all agencies with defining ecosystems, managers worldwide use watersheds,
jurisdiction over lands or processes in the ecosystem, private catchment basins, and wetlands as units that can give
landowners within or adjacent to the system, and ecosystems defined spatial boundaries. Where and when
non-residents who use the system on a seasonal basis or this is the case, ecosystem managers will also often unify
who have specific and vested interests in the system. To proposed EBM modeling and management around water
accomplish this, EBM must integrate both ecological and quality planning goals intended to mitigate pollutant
social information at multiple levels and then coordinate run-off, including riparian and wetland restoration. In north-
management activities on separately owned and diversely eastern coastal Australia, such planning is of critical impor-
governed lands, often with conflicting mandates. To deter- tance to one of the world’s most unique marine ecosystems,
mine which management actions to choose, managers must the Great Barrier Reef (GBR). One of the most important
support ongoing dialogue with the community to evaluate elements in effective EBM is stakeholder identification and
different management and policy options. In doing so, they subsequent linking of stakeholder groups to both problems
must build community understanding of ecological issues, and solutions in EBM. We will see how linkage of identifi-
and then demonstrate that they are choosing management cation and interests are effectively related in the following
actions informed by meaningful deliberation with example.
stakeholders. The GBR has been degraded by declining water quality,
To accomplish these things, it is essential to identify and which has been caused by pollutants in surface water runoff
include non-agency stakeholders in decision-making pro- from terrestrial systems. One of the largest contributors to
cesses. But exactly what are stakeholders and who are such pollution is agricultural run-off, but agriculture in north-
stakeholders? We can address the “what” by defining eastern coastal Australia is a major industry employing or
stakeholders as individuals or groups possessing vested affecting livelihoods of millions of people. What kinds of
interests in the persistence, health, products, state, or tools can be developed to assess trade-offs between economic
services of a system to whom managers have legitimate and welfare and protection of GBR in land use? Using the frame-
defined functional, ethical or legal obligations. But stake- work for identifying and evaluating ecosystem services
holder identification is not enough. Managers must form provided by the Millennium Ecosystem Assessment, James
partnerships with stakeholders to ensure their involvement Butler and his colleagues identified and assessed trade-offs
in decision making and create community support for man- between ecosystem services and stakeholders linked to both
agement actions. We should understand partnerships in this the GBR and commercial agriculture in the combined
context to mean dynamic relationships among actors, based watersheds of the Tully and Murray Rivers (the Tully-Murray
on mutually agreed upon objectives, pursued through an Catchment) in northeastern Australia (Fig. 9.31) (Butler et al.
understanding of division of labor based on the respective 2013).
comparative advantage of each member (Brinkerhoff 2002), In this region, dissolved inorganic nitrogen (DIN) in sur-
not casual associations of people who happen to attend the face and groundwater, derived from fertilizers applied in
same meetings. sugarcane and banana production, enters marine waters and
Stakeholder involvement and partnership is an attractive degrades reef ecosystems. Pesticides and sediment pollution,
vision to contemplate, but a difficult reality to create. To be also derived from agriculture, contribute additional threats.
successful, EBM must achieve collaboration that leads to a The Tully Water Quality Improvement Plan (WQIP) was
common vision of desired future conditions, not just a designed as an EBM approach that would focus on reducing
Fig. 9.31 The Tully-Murray Catchment (floodplain) in northeastern Grieken, M.E., Lawson, T., Bruce, C., Kroon, F.J., Brodie, J.E., 2013.
Australia, where agricultural production and conservation of the Great An analysis of trade-offs between multiple ecosystem services and
Barrier Reef are linked through water quality regulation at the ecosystem stakeholders linked to land use and water quality management in the
level. (Reprinted by permission from Elsevier, from Butler, J.R.A., Great Barrier Reef, Australia. Agriculture, Ecosystems and Environment
Wong, G.Y., Metcalfe, D.J., Honzak, M., Pert, P.L., Rao, N., van 180, 176–191. # 2011 Elsevier B.V.)
Fig. 9.32 Ecosystem functions and services identified in the Tully- 2013. An analysis of trade-offs between multiple ecosystem services
Murray floodplain and adjacent Great Barrier Reef linked by water and stakeholders linked to land use and water quality management in the
quality regulation. (Reprinted by permission from Elsevier, from Butler, Great Barrier Reef, Australia. Agriculture, Ecosystems and Environment
J.R.A., Wong, G.Y., Metcalfe, D.J., Honzak, M., Pert, P.L., Rao, N., 180, 176–191. # 2011 Elsevier B.V.)
van Grieken, M.E., Lawson, T., Bruce, C., Kroon, F.J., Brodie, J.E.,
DIN runoff by applying agricultural best management clearance that may be done outside of protected areas. In
practices such as zero tillage, split fertilizer applications and their first scenario (Scenario 1), Butler et al. (2013) assumed
restoration of riparian vegetation and wetlands to areas cur- that there was no VMA and therefore no regulation. Under
rently contributing pollutants. As a first (scoping) step, these conditions, they assumed that all native vegetation and
managers identified ecosystem functions and services in the remaining forest fragments would be removed for more eco-
Tully-Murray floodplain and adjacent GBR which were or nomically profitable uses, all revegetation done since the
could be linked through the regulation of water quality passage of the VMA would be absent, and all plantation
(Fig. 9.32). forests would be converted to livestock grazing areas. Sce-
To assess trade-offs between ecosystem services, Butler nario 2 assumed present conditions. Scenario 3, which set
et al. (2013) created potential development scenarios (eco- priorities on reducing fertilizer and sediment pollution,
system models) to estimate relative changes in ecosystem maintained 2007 protected areas, replaced cropland and
services that were likely to occur in each scenario, grazing areas with riparian vegetation in sensitive areas, and
intentionally separating and separately analyzing effects on restored selected wetlands. Scenario 4, the most
ecosystem services in the Tully-Murray floodplain from conservation-oriented plan, replaced land devoted to sugar-
those associated with GBR for each major stakeholder cane, banana plantations and grazing with forestry. All
group and the associated management bodies responsible forests within protected areas and all vegetation regrowth
for protecting and sustaining that service (Fig. 9.33). Their were converted to native vegetation. Wetlands were restored
model construction process consisted of (1) developing alter- to pre-European extents. Riparian vegetation was maintained
native plausible land use scenarios for the Tully-Murray as in Scenario 2 to reduce fertilizer and sediment pollution,
floodplain; (2) measuring the status of floodplain ecosystem with protected areas with previously established boundaries
services associated with each scenario, and (3) estimating (Butler et al. Butler 2013).
resulting trends in GBR ecosystem services under each sce- Scenario 1 (no VMA) yielded the highest agricultural
nario (Butler et al. 2013). revenue of AUS $120 million per year, and the highest total
Management of terrestrial vegetation in the floodplain had N output of 1999 tonnes (1000 kg or 2200 pounds, larger
been governed by the Vegetation Management Act (VMA), a than a US “ton”) per year, resulting in maximum values for
law which regulated the type and amount of vegetation food and fiber production and lowest values for water quality
9.5 Creating and Managing Governance Systems of Ecosystem-Based Management 399
Fig. 9.33 Primary stakeholders, and statutory (solid line) and E., Lawson, T., Bruce, C., Kroon, F.J., Brodie, J.E., 2013. An analysis
non-statutory (dashed line) bodies responsible for water quality regula- of trade-offs between multiple ecosystem services and stakeholders
tion in the Tully-Murray watershed and adjacent Great Barrier Reef linked to land use and water quality management in the Great Barrier
relative to their role in managing ecosystem functions and services. Reef, Australia. Agriculture, Ecosystems and Environment 180, 176–
(Reprinted by permission from Elsevier, from Butler, J.R.A., Wong, 191. # 2011 Elsevier B.V.)
G.Y., Metcalfe, D.J., Honzak, M., Pert, P.L., Rao, N., van Grieken, M.
regulation among the scenarios. Under Present Day (Scenario maintained or increased. The most marked negative trade-off
2) conditions, models predicted that annual agricultural reve- occurred between farmers (for food and fiber production),
nue would decrease to AUS $113 million per year with who lost revenue, and GBR tourists, tour operators, and
corresponding decreases in total DIN (fertilizer) output, recreational and commercial fishermen, whose revenues or
resulting in a slight decline in food and fiber production and revenue-generating spending increased. As Butler et al.
a slight increase in water quality. In Scenario 3, agricultural noted, “The identification of linked ecological functions and
revenue decreased more than in Scenario 2, as did total DIN stakeholders. . .allows an assessment of the design of existing
output, resulting in further declines in food and fiber produc- water quality management. ‘Scale mis-matches’ occur when
tion and some additional increase in water quality. In Sce- the spatial scale of management is not aligned with the
nario 4, (Native Forestry), agricultural revenues decreased to ecosystem processes it aims to manage, resulting in socio-
AUS $37 million per year, resulting in the lowest levels of ecological system disruption and loss of resilience. . .” (But-
food and fiber production of all scenarios. Total N declined ler 2013:187). Overall, this analysis revealed strong negative
markedly to 596 tonnes per year, maximizing water quality. correlation between water quality regulation and agricultural
This reduction brought total N to the desired threshold for production. In contrast, it showed positive correlations
reducing fertilizer pollution in GBR, resulting in improved between water quality regulation, recreational and commer-
performance of marine GBR services. Residential and com- cial fisheries, and biodiversity.
mercial fisheries production, pest regulation, and biodiversity The effort of Butler et al. (2013) not only shows how to
increased to maximum levels, and marine ecosystem services connect proposed management actions with predicted
reached 91% of projected maximums (Butler 2013). outcomes, but how to connect outcomes to their ecological
For conservation, the most optimal scenario was Native effects and relate their benefits and costs to specific stake-
Forestry (Scenario 4). Here, although agricultural revenues holder groups who depend on an ecosystem for particular
fell, tourism and commercial fisheries revenue values were goods and services. Now the question is: what kind of
ecosystem-based governance structures make these recommended reductions in harvest were always delayed,
connections part of the management decision-making pro- but recommendations for increased harvests were
cess, and how can these kinds of structures be created? implemented immediately. Similarly, recommendations for
decreasing catches were only partially implemented, but
recommendations for increasing catches were fully
9.5.3 Costs of Bad Governance: Managing implemented. Brown et al. (2012) noted, for instance, that,
Time from Recommendation to Action in past management of Atlantic Canadian ground-fish stocks,
administrators and managers decreased catches by only 50%
Does the timing of ecosystem-based management decisions of the amount recommended by scientists, but always
matter? EBM affects human beings and their activities, and followed recommendations for increasing catches by the
so requires multiple agency coordination and agency full amount. As Brown et al. expressed in their own words,
approvals, as well as political processes that can extend “We found that management delays are pervasive and com-
over long periods to ensure participation and representation monly recognized by fishery scientists, although their cause
of all stakeholders. Let us take one example of the effect of was rarely reported. Delays in acting on population declines
delayed management decisions on ecosystem behavior in a may be caused by sociopolitical, institutional, and ecological
well-studied field: marine fisheries. factors... Longer delays, of up to 25 years occurred when
Christopher Brown of the University of Queensland social pressure meant managers were reluctant to reduce
(Australia) and his colleagues examined how delays in man- harvest limits despite evidence of population
agement interacted with environmentally driven changes in declines. . .Delays in management action resulted in collapse
population growth to affect sustainability and viability of of the fish population” (Brown et al. 2012: 302).
marine fisheries managed by international governing bodies Causes of delays were varied, but always political
(Brown et al. 2012). They found that delays implementing (Table 9.8). Effects of delays were negative, in some cases
scientifically-based recommendations for globally catastrophic. You can see the outcome of the delays
administered marine fisheries were pervasive, and political graphically in Fig. 9.34. Using a simulation model
considerations, usually to appease stakeholder groups, constructed specifically for these fisheries data sets, Brown
influenced decision timing (Table 9.8). Actions on et al. (2012) found that, for a fish population whose
Table 9.8 Examples of delays in implementing management action on harvest changes for fisheries as recommended by scientists
Species/fishery Delay Cause Reference
Multispecies lake fishery of piscivores and 15–25 years, ongoing (as of 2008) effort reductions delayed to reduce short- de Leeuw
benthivores; Lake Ijsselmeer, the term losses in profit to fishers et al.
Netherlands (2008)
Australian Commonwealth fisheries 3–5 years policy requirement for turnover in strategic Smith et al.
assessments (2008)
North Pacific Minke Whale (Balaenoptera 12 years scientific uncertainty led to a long process to Punt and
acutorostrata, Balaenopteridae) calculate catch limits Donovan
(2007)
Whales 5 years time period required between reviews of Punt and
management strategy by the International Donovan
Whaling Commission (2007)
Orange roughy (Hoplostethus atlanticus, 5–7 years biological uncertainty meant a lengthy Bax et al.
Trachichthyidae); South-East Australia debate over sustainable catch limits (2003)
European eel (Anguilla anguilla, Not reported, but ICES has not clear from report ICES
Anguillidae); multiple river basins in repeatedly recommended that a (2005)
Europe recovery plan is needed
Herring (Clupea harengus, Cluperidae); 5 years not specified ICES
North Sea and Skagerrak (ICES Subareas (2008)
IV and IIIa)
Atlantic cod (Gadus morhua, Gadidae); 8 years, ongoing (as of 2009) not specified ICES
North Sea and Skagerrak (ICES Subareas (2008)
IV and IIIa)
Elasmobrachs, North Sea (ICES Subarea 2 years stock assessments provided every two years ICES
IV) (2008)
Reprinted by permission from Wiley, from Brown, C.J., Fulton, E.A., Possingham, H.P., Richardson, A.J., 2012. How long can fisheries
management delay action in response to ecosystem and climate change? Ecological Applications 22, 298–310. # 2012 Ecological Society of
America
Fig. 9.34 A hypothetical fished population simulated with environ- harvest on fished population, leading to eventual collapse of the fished
mental effects on the intrinsic growth rate (r) at time t. (a) Productivity population. (Reprinted by permission from Wiley, from Brown, C.J.,
(r) over the simulation period (solid line); r set for mean decline of 50% Fulton, E.A., Possingham, H.P., Richardson, A.J., 2012. How long can
over 50 years (dotted line). (b) Simulation with no management delay in fisheries management delay action in response to ecosystem and climate
implementing reduction in harvest. Catches (dotted line) closely follow change? Ecological Applications 22, 298–310. # 2012 Ecological
biomass (solid line) of fished population. (c) Ten-year management Society of America)
delay between recommendation and implementation of reduction in
productivity was declining (which is the state of almost all

commercially fished marine species in the world today)
(Fig. 9.34a), management action to reduce harvests
implemented without delay allowed the fishery to continue
at reduced but sustainable levels (Fig. 9.34b). In contrast, if
harvest reductions were delayed by 10-years (not an uncom-
mon delay in international fisheries management), the popu-
lation eventually crashed, and so did the fishery (Fig. 9.34c).
In all cases in which their model was applied, delays in acting
on scientific advice to reduce harvest increased the probabil-
ity that the fishery would collapse (Fig. 9.35).
After this review of multiple fishery studies, Brown et al.
(2012) found no cases in which managers took action to
reduce delays. Yet such initiative should have been and
must be viewed as an essential element in all forms of
EBM. Ecosystem managers must manage the timing of
implementation of management actions in order to achieve
the intended outcome of management actions. Managers can Fig. 9.35 Probability of collapse of a fished population increases with
longer delays between recommendation and implementation of harvest
take part in one or both of two strategies to reduce delays. reduction when the population is experiencing declining growth (solid
First, they can play a political role in advocating stronger line) or stable growth (dashed line). Under declining growth, the proba-
top-down governance and enforcement of harvest changes. bility of collapse approaches certainty with delays of more than 10 years.
Second, they can take part in creating incentives for sustain- (Reprinted by permission from Wiley, from Brown, C.J., Fulton, E.A.,
Possingham, H.P., Richardson, A.J., 2012. How long can fisheries
able fishing practices. On a global scale, those fisheries in management delay action in response to ecosystem and climate change?
which fishers, managers, and scientists cooperate to manage Ecological Applications 22, 298–310. # 2012 Ecological Society of
the harvest are fisheries that are less likely to be overexploited America)
(Brown et al. 2012). “Ideally,” Brown et al. noted, “delays in These and other threats directly affected megafauna spe-
management should be minimized to promote long-term cies in and around the GBR including dugongs, turtles, and
persistence and higher catches in fisheries impacted by envi- sharks, all of which have declined over time. Size and density
ronmental change. Alternatively, more conservative harvest of fish species targeted by recreational and commercial
limits are required to account for long delays in management fisheries were also trending downward. To combat such
action. Past history shows that such systems are challenging decline, the GBRMP underwent a major rezoning and trans-
to develop, but critical for long-term sustainability of fishing formation in governance toward stewardship of the large-
industries” (Brown et al. 2012:309). The problem is how to scale seascape, incorporating 70 bioregions (including
develop them. For a possible solution, we again examine the many non-reef habitats) under new governance structures.
Great Barrier Reef (GBR), a system that underwent massive The rezoning has been an ambitious effort to better manage
reorganization of its governance structures in order to sustain the GBR and associated ecosystems, especially to increase
the integrity of the ecosystem and the services it provided. their resilience to climate change, potential pollution from
terrestrial sources, and anthropogenic disturbances, while
maintaining the systems’ ability to provide essential ecosys-
9.5.4 Creating a Working Framework tem services (Olsson et al. 2008).
for Ecosystem-Based Governance: The This transition required a sequence of carefully executed
Case of the Great Barrier Reef steps (Table 9.9). Rezoning and transformation of gover-
nance began in 1998 at the initiative of the Great Barrier
As noted at the beginning of this chapter, one of the founda- Reef Marine Park Authority (GBRMPA), the primary gov-
tional problems of traditional resource management ernment agency responsible for management and protection
approaches was that decision-making processes were of the GBRMP. The GBRMPA initiated a major rezoning
compartmentalized jurisdictions focused on individual plan for the entire park, the Representative Areas Program
resources (such as timber, fisheries, soil, or wildlife). This (RAP), formulated “to systematically increase the protection
approach limited policy and management tools that were of biodiversity within the GBRMP by protecting representa-
developed and created incentives for over-exploiting tive examples of each type of habitat within a network of
resources that undermined system sustainability. Governance no-take areas” (Olsson et al. 2008:9491). The focus of the
structures developed under this approach could not respond RAP was on protecting biodiversity and maintaining ecosys-
to complexities of dynamic ecosystems or develop adaptive tem function and services rather than on the former, tradi-
capacities to cope with changes and uncertainties in those tional approach of separate, independent government
systems (Olsson et al. 2008). Single-resource approaches are agencies attempting to maximize yield of commercially
also easily overwhelmed by global economic drivers or important fisheries.
global patterns of environmental change like global climate Formal development and implementation of RAP could
change and cannot easily address complex and dynamic not be accomplished by agency edict. It required extended
relationships of linked social-ecological systems (ecosystems involvement and interaction with stakeholders and the
in which humans and their activities are deeply embedded). surrounding community. Such interaction took place in two
Solving these problems requires transition to a system-based phases. First, over a 3-month period from May through July
approach that integrates all processes that affect these 2002, GBRMPA sought community input for preparation of
resources and their use. a Draft Zoning Plan for the entire GBRMP. Through a
The Great Barrier Reef Marine Park (GBRMP) contains combination of expert opinion, stakeholder involvement,
one of the largest coral reef systems in the world, covering and analytical approaches, different options for no-take area
344,000 km2, an area about the size of the US state of networks were identified using a variety of planning tools to
California. Within the GBRMP, coral-dominated integrate biophysical, social, and economic information.
assemblages form discontinuous fringing reefs along the These efforts led to completion of the Draft Zoning Plan,
mainland coast and offshore islands, with most individual which then guided the second community participation
reefs located 30–200 km offshore (Olsson et al. 2008). As we phase, which solicited additional input on the proposed
saw in our study of the Tully-Murray catchment, the plan, and received 21,500 additional written submissions of
GBRMP, including the GBR itself, has long been showing suggested revision. Informed by these submissions and other
signs of degradation, primarily from runoff of sediment from input, the GBRMPA presented a Revised Zoning Plan by
land, over-harvesting of fishery resources, and increasing 2003 which increased the percentage of no-take areas in the
water temperatures associated with global climate change. GBRMP to 33% (more than six times the percentage of the
The various and diverse ecosystems associated with, but original plan) and included at least 20% of the area of each of
distinct from, the GBR were each subject to a unique array the 70 bioregions. In December 2003 the new zoning plan
of threats. Land-based pollution was diffuse, but often was submitted to the Australian parliament and passed into
associated with agriculture. law in July 2004. The Australian Government also agreed to
Table 9.9 Strategies used by the Great Barrier Reef Marine Park Authority (GBRMPA) (Australia) to facilitate transition from resource-based
management to ecosystem-based management
Strategies Actions Examples of barriers to change
Making internal organizational Establishing Senior Managers Forum and four Resource constraints
changes regional teams Inability to innovate or deal with surprise
Providing clear and transparent leadership at the Lack of direction, shared vision, engagement, trust,
relevant levels within the organization leadership, cross-sector cooperation, and communication
Communicating a shared vision and goals Having few leaders exacerbates vulnerability
Bridging science and policy Drawing on existing networks of scientists, Science is fragmented
managers and industry to Lack of scientific certainty
Promote dialogue Different perceptions and views among scientists and
Workshops and forums for synthesizing managers, lack of trust
knowledge
Communicating shared vision and goals
Changing public perceptions Clear, simple, and tailored stakeholder Different knowledge and interests among stakeholder
information from a groups
communication professional Low awareness of problems, threats, and ecological
Visualizing the entire GBR as an interconnected interactions
ecosystem
Creating a sense of urgency for conservation
Facilitating community Building trust with communities: personal Lack of trust
participation and public interactions and regional teams Conflicting views among key actor groups,
consultation Community information sessions misinformation
Recasting problems as opportunities Outreach to local communities difficult
Periodic updates on the rezoning process Lack of leadership
Innovative submission routines
Gaining political support Prepared for change: Politically expert staff, Change of people in power
timing actions, having relevant information ready Lack of support from key politicians
Briefing key players well before the new zoning Zoning plans can be stopped
plan Opposing views
Allying with other key actor groups
Pollsters for leverage and monitoring public
opinion
Reprinted by permission from the National Academy of Sciences, from Olsson, P., Folke, C., Hughes, T.P., 2008. Navigating the transition to
ecosystem-based management of the Great Barrier Reef, Australia Proceedings of the National Academy of Sciences 105, 9489–9494. # 2008.
National Academy of Sciences, USA
a structural adjustment package that provided compensation The GBRMPA’s restructuring of the governance of the
for commercial fishers and others adversely affected by the GBR ecosystem showed that policy development and imple-
new zoning (Olsson et al. 2008). mentation are complex, highly dynamic, and sometimes
This transition to a new EBM governance structure abrupt. After years of slow, seemingly ineffectual effort,
employed five strategies to implement RAP and the new transitions can occur rapidly if and when a policy window
zoning plan. These were (1) internal organizational changes, of opportunity appears. Taking advantage of such a window
(2) bridging communication and action between science and requires political skill because enabling legislation is a criti-
policy, (3) changing public perceptions, (4) facilitating public cal element for achieving adaptive co-management, along
consultation and community participation, and (5) gaining with effective interaction between science and policy.
political support (Table 9.9). Timing and political skill were Along with these elements, those committed to comprehen-
also vital. The zoning plan had to be submitted to the sive EBM must also be able to create more flexible manage-
Australian Parliament in December 2003 if it was going to ment institutions and more extensive public support if they
have any chance to become operational by July 2004, prior to wish to achieve effective and sustainable management of an
the upcoming federal election the same year. The chair of the ecosystem.
GBRMPA stated in retrospect, “I realized. . .that we needed
15 sitting days of Parliament, and a simple examination of the
parliamentary sitting timetable showed that if one wanted the 9.5.5 Relational Governance: Managing
15 sitting days to end, say, around midyear, then the plan had Stakeholder Interactions by Building
to be submitted to Parliament before the Parliament got up for Trust
the Christmas break. If one didn’t do that. . .one would have
been looking at October, November, which would have been EBM assumes interaction between managers and
slap-bang in the middle of a federal election” (quoted in stakeholders, and the case of governance transition in the
Olsson et al. 2008:9493). GBR provides an example in which these interactions
contributed to the intended outcome. But how do managers of knowledge about the system being managed (Weeks and
foster interactions with stakeholders that lead to cooperation Packard 1997; Olsen and Shindler 2010). But the most
and not conflict? Managers and their agencies have often important factor is trust. Bruce Shindler, natural resource
interacted with stakeholders using the so-called “knowl- and social scientist at Oregon State University and his
edge-deficient” model, which presumes that stakeholder colleagues studied the element of trust in stakeholder
opposition to proposed management action or agency policy interactions with federal agency decisions involving the man-
reflects a lack of scientific understanding. When managers agement of sagebrush (Artemisia spp.) ecosystems prevalent
make this assumption, they invest heavily in education in much of the intermountain western US, particularly on the
programs designed to “enlighten” stakeholders and the pub- issue of using prescribed fire as a management technique
lic. As noted earlier, it is vital for managers to build commu- (Shindler et al. 2011). What caused stakeholder trust of
nity understanding of ecological issues, but this is most agency managers to increase or decrease, and were these
effective when it is done interactively, and with respect for things managers could influence?
the often considerable experience and expertise that knowl- Rural and urban landowners in the same region in the state
edgeable stakeholders have with systems which they use or in of Utah were asked about six agency practices employed in
which they reside. Unfortunately, the knowledge-deficient managing sagebrush-juniper (Juniperus spp.) systems:
model of manager-stakeholder interaction does not always grazing, felling trees, prescribed fire, mowing, spraying
convey respect for stakeholders or foster two-way, give-and– herbicides and chaining (i.e., removing sagebrush by drag-
take communication essential for public acceptance of man- ging a heavy chain between two bulldozers). There were
agement actions. It fails not only because its foundational differences in perceptions between groups regarding what
premise of stakeholder ignorance is often false, but because were acceptable practices, with rural residents being more
public acceptance of management decisions and actions is a accepting of grazing and felling (Table 9.10). But a more
function of both technical understanding (of management telling question was posed in a separate questionnaire. “How
technique and of science) and personal experience (with the much do you trust federal agencies like the BLM [Bureau of
agency employing the technique). If such understanding is Land Management] or Forest Service to use these practices
effectively communicated, it gives all participants on rangelands in the Great Basin?” “Trust” ratings were less
(stakeholders and managers) shared and accurate understand- positive than “acceptability” ratings (Shindler et al. 2011).
ing of how the ecosystem actually functions. In contrast, if Put trust scores and acceptance scores side by side
stakeholder experience with a manager or agency is one of (Fig. 9.36), and you immediately see that trust levels were
disdain or disregard for stakeholder viewpoints or inputs, lower than acceptance levels for all six practices. Acceptabil-
stakeholders are unlikely to support agency decisions. ity of a management practice was not the most important
Recent studies in EBM in which stakeholder interests are objection to the practice. The most important objection was
prominent, such as ranching, forestry, and fishing, show that lack of trust in agency managers to use the technique prop-
public acceptance of management actions depends on a suite erly or employ it for the right reasons.
of factors, including political costs of change, past What contributed to this lack of trust? Agencies got low
relationships among stakeholders, beliefs about appropriate grades on engaging the public in making decisions and even
relationship between humans and nature, and differing types lower marks on effectively building trust and cooperation
Table 9.10 Acceptability of management options for fuel hazard reduction and sagebrush ecosystem restoration. Values are weighted percentages
categorized by urban (U) or rural (R) settings
Mowing
shrubs
Prescribed Livestock Felling and Herbicide Chaining
fire grazing1 trees1 grass1 application1 trees1
This practice. . . U R U R U R U R U R U R
Is a legitimate tool that land managers should be able to use 39 41 48 63 22 42 27 35 11 24 11 31
whenever they see fit
Should be done infrequently, only in carefully selected areas 45 40 31 18 42 31 34 35 34 36 24 33
Should not be considered because it creates too many negative 6 8 8 5 14 9 11 8 24 17 24 11
impacts
Is an unnecessary practice 4 5 4 4 9 9 9 11 20 14 23 13
Don’t know 7 6 10 10 14 9 19 11 12 10 19 12
1
Indicates a significant difference between urban and rural responses P 0.05; N = 1345. Reprinted by permission from Elsevier, from Shindler, B.,
Gordon, R., Brunson, M.W., Olsen, C., 2011. Public perceptions of sagebrush ecosystem management in the Great Basin. Rangeland Ecology &
Management 64, 335–343. # 2011 Elsevier B.V.
Livestock 89
Grazinga, b 62
88
Prescribed Fireb 61
78
Felling Treesa, b 64
Mowing Shrubs 78
& Grassesa 65
Herbicide 60
Applicationa 44
Chaining 60 Acceptability
Treea, b 51 Trust
0 10 20 30 40 50 60 70 80 90 100
Percent of Response
Fig. 9.36 Comparison of acceptability of management practices in trust, moderate trust, full trust). Significant differences between urban
sagebrush- (Atemisia spp.) dominated ecosystems in the Great Basin, and rural residents about acceptability (a) and trust (b) P 0.05.
USA, and trust in federal agencies to implement the practices appropri- χ2 analysis. (Reprinted by permission from Elsevier, from Shindler,
ately by rural and urban residents in the Great Basin. Values represent B., Gordon, R., Brunson, M.W., Olsen, C., 2011. et al. Public
weighted percentages from full respondent sample (N ¼ 1345). Accept- perceptions of sagebrush ecosystem management in the Great Basin.
ability scores are combined percentages of the first two categories in Rangeland Ecology & Management 64, 335–343. # 2011 Elsevier
Table 9.10. Trust scores based on a four-point scale (no trust, limited B.V.)
Table 9.11 Agreements with statements about interactions with management agencies
Percent agree (don’t know)
Statements about interactions Urban Rural
Agency information about projects usually provides a good explanation of options and consequences. 43 (38) 46 (25)
Federal managers use public input to help make decisions. 20 (40) 27 (20)
Federal managers effectively build trust and cooperation with local citizens. 17 (38) 21 (20)
There are few opportunities for citizens to participate in the agency planning process. 52 (32) 59 (18)
I am skeptical of information from federal management agencies. 51 (23) 61 (14)
Local agency staff are constrained from doing their jobs by government restrictions at the national level. 53 (37) 61 (28)
Response options were on a non-point scale from strongly disagree to strongly agree, with a don’t know option. Strongly agree and agree are
combined for presentation purposes. Values are weighted percentages. Significant differences were noted between urban and rural residents for all
statements (P 0.05. χ2 analysis; N ¼ 1345 and varies slightly by statement). (Reprinted by permission from Elsevier, from Shindler, B., Gordon,
R., Brunson, M.W., Olsen, C., 2011. et al. Public perceptions of sagebrush ecosystem management in the Great Basin. Rangeland Ecology &
Management 64, 335–343. # 2011 Elsevier B.V.)
with local citizens (Table 9.11), areas in which only 17% of to participatory processes. As a result, even though agencies
urban residents and 21% of rural residents thought agency had “events” such as hearings or agency “open house” days
personnel did a good job. Shindler et al. noted, “Overall, to hear from stakeholders on particular issues, most
respondents were critical of agency actions . . .Both partici- stakeholders found little satisfaction in such interactions.
pant groups gave agencies particularly low marks for trust Neither urban nor rural residents perceived that agency per-
building and their use of public input. At the same time, they sonnel wanted to include their views or listen to them when
saw few opportunities to participate and were skeptical of the they offered input.
information provided by the agencies. . .[overall], trust in Trust is built, sequentially, in four steps (Covey and
agency competence was the most highly correlated factor in Merrill 2006). The first is through integrity. Ecosystem
acceptance of each practice” (Shindler et al. 2011:338–339, managers must make statements consistent with their actions,
emphasis ours). and must demonstrate that their decision-making processes
Neither urban nor rural residents perceived management are fair, impartial, and without favoritism to any particular
agencies as inclusive in decision-making or fully committed individual or group. They must actively solicit public input
and show that such input can really affect the outcomes of the will not necessarily produce a quick fix, but rather become a
decision-making process. That is, they must show normal way of doing business – then the goal of trustworthy
stakeholders that their input is consequential, that it makes relations. . .is realistic” (Shindler et al. 2011: 341).
a real difference in decision making. Second, managers and
agencies must clearly communicate the intentions of pro-
posed management actions, explaining and identifying who 9.5.6 Stakeholders as Managers: Ecosystem
will reap the benefits and who will bear the costs, and show Management from the Bottom Up
that they have sensitivity and concern for both. Third,
managers and their agencies must demonstrate capability in Besides lack of trust, other recurring relational problems that
using the management techniques they advocate with skill develop in EBM often arise from the fact that management
and effectiveness. One prescribed burn that escapes its “pre- initiatives usually originate from government agencies and
scription” and turns into an uncontrolled wildfire will undo fail to effectively engage the public and its diverse stake-
years of good work in an agency’s heretofore well-run pre- holder communities. When this happens, agencies create ad
scribed burning program. And one of the best ways to show hoc groups for individual ecosystem management projects,
stakeholders that managers are capable is to invite them to but fail to form the kinds of genuine partnerships with
observe situations where management techniques are being stakeholders essential for enduring trust and lasting success.
implemented and, where possible and safe, include To be effective in the long-run, ecosystem management
stakeholders in the management action, such as, in a sage- groups would not only need to incorporate strong
brush ecosystem, planting native vegetation, removing partnerships, but also be more-or-less permanent, have their
non-native plants, or helping to construct fences that direct own budgets, and possess the authority to make meaningful
grazing livestock to appropriate areas. decisions about ecosystem management policies.
Finally, managers and agencies must build credibility, a Agencies have been traditionally reluctant to form such
track record of effective management and positive outcomes groups, fearing that they will lose power in the process.
over a period of many years. This is not easy, given the However, sometimes citizens and stakeholders tire of waiting
turnover of personnel common in government agencies, but for agencies to act and simply create radical reorganizations
transience in personnel can be overcome by faithfulness of stakeholder-agency relationships themselves. One exam-
to core values and policies, especially those that local ple has been the formation of “watershed councils” (yet
stakeholders understand, affirm, and help to form. That is, another example that watersheds are increasingly used world-
credibility must have an ethical dimension as well as a wide as effective units for defining ecosystem locations and
professional and technical dimension. Managers must dem- boundaries) which consist of groups of public and private
onstrate repeatedly that they have the specialized skills and stakeholders that address ecosystem needs within individual
expertise to carry out management actions effectively, but watersheds. For example, the US Pacific Rivers Council,
also demonstrate repeatedly that they have empathy with part of the Conservation Alliance, a coalition of outdoor-
stakeholder needs and values, because human beings judge related businesses committed to the preservation of natural
the world, and the people in it, from their own “sense of self” areas for habitat and recreational values, funded a compre-
which is based on their own needs, beliefs, and values. hensive study of Pacific salmon populations in the US Pacific
Managers must show stakeholders that their decisions reflect Northwest that produced methods and guidelines for
an understanding of these things in local situations and prioritizing these populations for conservation (Allendorf
contexts, and that, as managers, they communicate with et al. 1997).
stakeholders honestly in every situation. When stakeholders A more comprehensive initiative in privately-led EBM
view managers as honest, committed to lasting values efforts, also based on watersheds, emerged in Hawaii
associated with their agencies mission, and empathetic in (USA). On the Hawaiian Islands, upland forest watersheds
understanding the stakeholders’ perspective, managers will have value as reservoirs of biological diversity, as recharge
be perceived as trustworthy (Horton et al. 2015). areas critical for underground aquifers, and as sources of
Without trust in managers and their agencies, an EBM billions of gallons of surface water to agricultural, residential
approach, no matter how scientifically and technically and commercial sectors. However, the Hawaiian rainforest
sophisticated, cannot be implemented. But if trust is has been degraded and reduced to only 58% of its original
established and practiced, EBM can become part of the cover. It continues to be under pressure from development,
culture of resident human communities, and conservation increasing demand for water, and continued environmental
itself can then become a sustainable social entity in the degradation caused by feral and invasive alien species. State
preservation and management of the ecosystem. As Shindler government agencies have lacked sufficient resources to
et al. summarized, “when trust building is viewed as a con- address comprehensive ecosystem management of these
tinual process encompassing a suite of attributes – ones that watersheds, in part because of historically low investment
in forest protection at the state level. In addition, much established a pattern of building effective collaboration and
remaining forested landscape is in the hands of private management (Fig. 9.37) based on a recurring set of common
landowners. In this situation, landowners, principles and procedures. First, participants found that using
non-governmental organizations, and state and federal interdisciplinary science-based models to identify manage-
agencies in Hawaii have formed partnerships to conserve ment targets and strategies was most effective. Informed by
and manage the state’s forested watersheds by voicing their such models, participants gained a shared and accurate view
concerns, generating and implementing action plans and of the state of the system and the processes driving it and
petitioning for greater support (Gutrich et al. 2005). developed a common understanding of threats facing the
The first such collaboration to attempt an EBM effort was system. Note that this strategy illustrates in practice what
the East Maui Watershed Partnership. In this case, was described earlier in principle; the use of conceptual
stakeholders wanted to establish long-term protection of models as heuristic devices to improve communication
forests in the watershed by placing a fence around the water- among people of diverse backgrounds and expertise.
shed to exclude feral pigs and goats from higher elevations, Second, effective partnerships developed a common lan-
remove pigs and goats already in the forest by hunting and guage among stakeholders and decision makers for compar-
trapping, and remove or slow the spread of exotic plant ing various management options that were proposed. This
species. Other partnerships with similar objectives soon occurred through repeated and iterative stages of scientific
began to develop in other watersheds, supported by the analysis that informed stakeholder deliberations, and such
Hawaii State Legislature’s declaration of 2003 as the “Year deliberations in turn directed subsequent analysis.
of the Hawaiian Forest,” a move that brought attention, and A third requisite was trust, which, as we saw in the
some funding, to forest ecosystem management and protec- example of management of sagebrush ecosystems in the US
tion initiative. In this effort, watershed partnerships Great Basin, is the most important factor in gaining public
Fig. 9.37 A general framework of the public process for building Manopimoke, S., McCauley, D., Norton, B., Sabatier, P., Salzman, J.,
partnerships, achieving consensus, and implementing management Sasmitawidjaja, V., 2005. Science in the public process of ecosystem
plans among public and private stakeholders as employed in watershed management: lessons from Hawaii, Southeast Asia, Africa and the US
partnerships in Hawaii (USA). (Reprinted by permission from Elsevier, Mainland. Journal of Environmental Management 76, 197–209.
from Gutrich, J., Donovan, D., Finucane, M., Focht, W., Hitzhusen, F., # 2005 Elsevier Ltd)
support for government management actions. In this case,

trust proved an essential element to building consensus. too few species, even as they overlook biota that are
Watershed coalitions found that, to build trust, integration small, difficult to classify, or not appealing to public
of scientific input needed to be transparent and responsive to sentiments. But EBM remains technically and politi-
feedback from the public. Public trust of scientists was high cally challenging. To make progress, conservationists
when scientists were viewed as independent and objective, must develop a common, accepted, and operational
but low when scientists were perceived as members of a definition of the concept, devise practical ways to
coalition with a predetermined agenda. In general, when implement it under varying conditions, and create
government officials established high levels of trust among political support and legislative mandates to translate
non-government stakeholders, stakeholders were willing to ecosystem management concepts into enforced policy
accept agency analyses with little reservation or debate. If directives. Progress will necessitate conflict and
trust was low, stakeholders demanded independent analysis clarification of values. Individual agency jurisdictions
and took longer to reach agreement on management plans. must be replaced with permanent working groups or
Lack of trust undermined effective application of scientific boards with independent budgets, regional authority,
input. In the absence of trust, no consensus was achieved and and legal mandates. Without these, EBM will remain a
partnerships failed. compelling concept, but a frustrating and mostly unful-
Fourth, watershed partnerships had to give careful consid- filled practice.
eration to benefits and costs accrued to each stakeholder
group under different management options. Plans that called
for substantial costs to be borne by stakeholders who had
been excluded from the planning process were least likely to
succeed (Gutrich et al. 2005).
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Values and Ethics in Conservation
10
Future historians will find our century remarkable for its breadth of knowledge and narrowness of value
judgments. Never have humans known so much about, and valued so little in, the great chain of being.
Holmes Rolston III 1986:114
Keywords treated by biologists as a central object of scientific interest,

Instrumental value · Intrinsic value · Ecosystem valuation · but scientific interest is not the same as, nor does it require or
Primary environmental ethics · Secondary environmental impute, ethical value. A view of “ethics-free conservation”
ethics · Moral extensionism · Deep ecology · Ecocentrism · rests on perceiving conservation, specifically conservation
Rights of nature · Faith-based conservation biology, solely as a “regulatory science” whose purpose, in
the words of environmental legal scholar A. Dan Tarlock, is,
“to develop scientific standards that can be applied to regu-
latory criteria and then to develop management strategies to
Overview meet those standards” (Tarlock 1994:1130). In this view,
In this chapter, you will learn about: conservation biologists collect and analyze scientific data
relevant to questions policy makers and politicians consider
1. Characteristics that distinguish major categories of important to the development and enforcement of conserva-
value in conservation. tion law and policy. As conservation scientist Dennis Mur-
2. Philosophical, cultural, and religious traditions that phy explained, conservation biology “exists only because
affirm instrumental and intrinsic values of species, biological information is needed to guide policy decision
biodiversity, and natural objects. making” (Murphy 1990:203). Conservationists don’t need
3. The interaction of values and ethics with conserva- ethics. That comes from policy makers and legislators
tion problems to produce comprehensive and cultur- informed by public sentiments.
ally acceptable solutions to conservation dilemmas. A different view of conservation biology is espoused by
other conservationists, including those who founded conser-
vation biology as a modern scientific and academic disci-
pline. As discussed in Chap. 1, one of the founders of modern
conservation biology, Michael Soulé, defined it as a disci-
pline that is value laden and mission driven, a mission
organized around what Soulé called four “normative
10.1 Does Conservation Science Need postulates” that direct its intentions. These are (1) diversity
Conservation Ethics? of organisms is good, (2) ecological complexity is good,
(3) evolution is good, and (4) biotic diversity has intrinsic
10.1.1 Conservation Biology – Regulatory
value, regardless of its utilitarian value. These normative
Science or Value-Laden Mission?
postulates assume normative outcomes. As Soulé and others
wrote more recently, “We propose. . .that conservation
An honest discussion of ethics in conservation must begin
practitioners, whether in a public or private (nongovernmen-
with the admission that many conservationists claim there is
tal) employ, are . . .obligated to apply new biological
none, nor any need for one. The sheer variety of life is often

412 10 Values and Ethics in Conservation
knowledge in their work. Such a doctrine of “best conserva- furthers attainment of that end” (Daly 1999:694). To further
tion practices based on the best science” is tantamount to an the appreciation of the value of biodiversity, conservation
ethical obligation of biologists to adopt a higher standard for biology must acknowledge and explain the purpose of
management than is mandated by existing statutes and conserving it.
regulations,” (Soulé et al. 2005:175). This is what conserva- We, the authors, affirm this second view of conservation,
tion biologist Reed Noss referred to as the “overarching having found that seeing conservation biology only as a
normative assumption” in conservation biology, that “biodi- “regulatory science” is not accurate in concept, workable in
versity is good and ought to be preserved” (Noss 1999:117). practice, or faithful to conservation’s intellectual and histori-
Paul Angermeier of the US Geological Service put it this cal origins. The modern environmental movement itself,
way, “Conservation biology exists because conservationists from its inception, has always been characterized by four
believe that biotic diversity, ecological complexity, and evo- main goals – conservation, restoration, sustainability and
lution are intrinsically good. . .and ought to be conserved” equity. But every one of these is an ethical goal, not a
(Angermeier 2000:377). Environmental philosophers Dwight scientific hypothesis of analytical description of a state of
Barry and Max Oelschlaeger elaborate, “To deserve its title, nature. The study and practice of conservation biology
conservation biology must be ethically overt – that is, it must remains grounded in these goals. In this long-standing and
affirm its mission to be the protection of habitat and the increasingly important context of conservation, we will
preservation of biodiversity. Otherwise its name is as linguis- assume that there are legitimate ethical norms in conservation
tically dishonest as the (Orwellian) claim that ‘war is science and practice. Therefore, our interpretation of conser-
peace.’. . .conservation biology is not applied biology but vation ethics is what ethical philosophers would call a
rather hinges on an explicit evaluative judgment: Biodiver- cognitivist view –that reasoned ethical assertions can repre-
sity is good and should be preserved. Apart from such a value sent claims about ethical realities of right and wrong, not
judgment, one must wonder why there would be any reason merely expressions of attitudes of the persons stating them
to invest effort in conservation biology” (Barry and (an emotivist view) (Sarkar 2012). From this cognitivist
Oelschlaeger 1996:906). foundation, we assume that one of the most important roles
In this second view, it is conservation values, not of ethics and values in conservation is to discover what the
appointed officials, that drive conservation policy. Again fundamental purpose of conservation is. And evaluating
Angermeier explains the value-policy relationship with clar- competing values in conservation requires ethical judgment.
ity. “Because value-based policies limit conservation success What ethically legitimate goals and ends can exist for
more than does biological knowledge. . ., the most crucial conservation? An understanding of conservation ethics,
task facing conservationists is facilitating shifts in societal which direct conservation values (the basis for what we
values toward more respect for nature. . .To be effective, judge to be admirable or worthy in conservation effort) and
conservation biologists will need to openly profess their goals (the objective determined or desired from a particular
values and persuade others that natural biotic diversity conservation action or plan), is necessary for making man-
contributes significantly to the quality of human lives. agement decisions and explaining those decisions to the
Much of nature as we know it hinges on our success in public, as well as for personal and professional development
these endeavors” (Angermeier 2000:379). As conservation as a conservation practitioner. Values and ethics must be
policy expert Tim Clark put it, “No conservation biologist informed by science, but are not the same as science. Values
can escape being part of policy making” (Clark 2001:33, and ethics constitute, rather, another way of knowing and
emphasis his), which, as Clark goes on to point out, is “the understanding the world around us and of defining the
making of important decisions that affect the distribution of problems that conservation must solve.
values in society” (Clark 2001:33, definition from Laswell
and McDougal 1992:1269). Thus, to engage its mission,
conservation biology, and its leading professional society, 10.1.2 Value -- Property of Nature or Product
the Society for Conservation Biology, have always felt it of Thought? Problems of Plastic Trees
essential to engage in policy formation rooted in conservation
values, not simply have conservation policies dictated by In 1972, the city of Los Angeles, California (USA) made an
elected and appointed policy makers (Meine et al. innovative proposal for urban beautification. It planned to
2006:642). And it is precisely such values that have always line the median strip of a major boulevard with plastic trees
distinguished conservation biology as a mission-driven disci- constructed of factory-made “leaves” and “branches” wired
pline, a discipline that proceeds with a purpose. Indeed, the to plumbing pipes. The trees were “planted” in aggregate
noted ecological economist Herman Daly called conservation rock coated with epoxy, and after their installation, an
“a policy in the service of a purpose. . .To have purpose unknown person or persons added plastic birds (Tribe
means to serve an end, and value is imputed to whatever 1974). Despite the unnaturalness of plastic trees, proponents
10.1 Does Conservation Science Need Conservation Ethics? 413
of the plan could marshal a compelling argument: only plastic externally given,. . .rather than generated by reason, environ-
trees, they contended, could survive the soil-deficient, smog- mental policy makes a value judgment of enormous signifi-
ridden environment of downtown Los Angeles, and a plastic cance. And once that judgment has been made, any claim for
tree, however artificial, would be more appealing to LA the continued existence of threatened wilderness areas or
residents than a dead or dying one, however real. Evaluating endangered species must rest on the identification of human
the problem from the standpoint of cost-benefit analysis, wants and needs which would be jeopardized by a disputed
urban research planner Martin Krieger, in an article in Sci- development” (Tribe 1974:1326–1327).
ence, concluded that “the demand for rare environments is a A modern example of “plastic trees” can be seen in the
learned one” and “conscious public choice can manipulate “wilderness lodge” constructed by the Walt Disney Corpora-
this learning so the environments which people learn to use tion in its Walt Disney World theme park in Orlando, Florida
and want reflect environments that are likely to be available at (USA) (Fig. 10.1). Disney’s lodge web page encourages
low cost” (Krieger 1973:451). Ultimately, Krieger concludes prospective visitors to “Escape to the rustic majesty of
“It is possible that by manipulating memory through the America’s Great Northwest. Inspired by turn-of-the-century
rewriting of history, environments will come to have new National Park lodges, Disney’s Wilderness Lodge celebrates
meaning” and that “In order to create substitutes, we must American craftsmanship and honors the beauty of the
endow new objects with significance by means of advertising untamed wilderness. Soak in the splendor of the great out-
and social practice. . .Much more can be done with plastic doors, from nature trails through pine forests to rocking
trees and the like to give most people the feeling that they are chairs that overlook a murmuring creek. Inside. . .enjoy the
experiencing nature” (Krieger 1973:453). rustic elegance of the stone hearth and roaring fireplace”
Here, in the world’s most respected scientific publication, (Walt Disney World 2018). Enjoyable as these comforts
Krieger advocates (1) manipulating memory (influence with- and aesthetics may be, many find the idea, and the reality,
out consent); (2) rewriting history (deception); (3) substitu- ethically repulsive. Environmental philosopher Sahotra
tion and simulation of natural environments with man-made Sarkar explains, “What we value in our experiences seems
ones (devaluing nature); and (4) creating value and meaning obviously mediated by what knowledge we bring to them
through advertising and “social practice” (confusing “value” beyond the immediate input of our senses. . .what seems to
with “benefit” and “meaning” with “demand”). matter is the authenticity of an experience. In other words, we
In these words, Krieger, perhaps unintentionally, has value relational properties including the historical prove-
defined a fundamental question to be answered in all debates nance of experience” (Sarkar 2012:154–155).
about the value of natural objects and environments. Namely, Sarkar’s point is that we must decide whether our ethical
is the goal of conservation ethics to determine the value of a obligation is to satisfy our desires or preserve what is valu-
creature – such as a tree – or a natural object – like the Grand able, independent of those desires. “We can be truly free to
Canyon – or is the goal to teach people to use and want pursue our ends,” notes Tribe, “only if we act out of obliga-
environments that can be made available to them easily and tion, the seeming antithesis of freedom. To be free is not
cheaply? In other words, do we address and determine the simply to follow our ever-changing wants wherever they
true value of diversity and rarity, or do we teach people not to might lead. To be free is to choose what we shall want,
miss them when they disappear? At a deeper level, Krieger what we shall value, and therefore what we shall be” (Tribe
draws us toward an even more important question – is a tree 1974:1326–1327). Tribe insightfully tells us we must be
(or anything else in nature) valuable because we (humans) more intentional in choosing our values, but is the value of
value it, or is it be valuable for some other reason? nature and all it contains simply and only a matter of human
The legal ethicist Lawrence Tribe, responding to Krieger’s choice? Are we choosing to conserve nature and its biodiver-
rationale for using plastic trees noted, “Policy analysts typi- sity as a means to some other end, or as an end in itself that
cally operate within a social, political, and intellectual tradi- should be valued for itself? To succeed as a discipline, values
tion that regards the satisfaction of individual human wants as in conservation biology must answer this question and so
the only defensible measure of the good, a tradition that mature into ethics that govern behavior if those values are
perceives the only legitimate task of reason to be that of to contribute to achieving conservation goals.
consistently identifying and then serving individual appetite,
preference, or desire” (Tribe 1974:1325).
Although humans perceive intrinsic and aesthetic values
Conservation biologists have rejected Krieger’s vision
in other species and natural objects, human appetites and
of providing “environments that people use and want at
preferences cannot be the ultimate foundation of value.
low cost” as a fundamental objective of their work.
Only when values are linked to purposes greater than self-
Why was this vision of “environmental management”
interest do such values become more than expressions of
rejected? What alternative visions have replaced it?
personal preference. As Tribe puts it, “By treating human
What do the alternatives reveal about what conserva-
need and desire as the ultimate frame of reference, and by
tion biologists value?
assuming that human goals and ends must be taken as
Fig. 10.1 The “wilderness lodge” the Walt Disney Corporation trails through pine forests to rocking chairs that overlook a murmuring
constructed in its Walt Disney World theme park in Orlando, Florida creek. . . and enjoy the rustic elegance of the stone hearth and roaring
(USA). Placed in a theme park in a major urban environment, Disney’s fireplace” (Walt Disney World 2018). Is such an artifact as the Wilder-
designated lodge web page nevertheless encourages potential lodge ness Lodge a legitimate service to tourists or an ethical misrepresenta-
visitors to “Escape to the rustic majesty of America’s Great tion of the value of the natural world? (Photo courtesy of Lt Powers,
Northwest. . . Soak in the splendor of the great outdoors, from nature under CC-BY-SA-4.0)
spp.) in an area used by elk (Cervus elaphus) as winter range

10.2 The Necessity of Value Judgments (Fig. 10.2). Her intention is to enhance forage production and
in Conservation quality for elk during winter – a particularly stressful period
of the year. Ongoing studies reveal that forage production on
10.2.1 Recognizing Management Actions the winter range is low, and so is the population growth of the
as Value Judgments elk using it. The action thus meets all criteria of our definition
of “management.” The site of the burn is located in a man-
Reflecting on the failures of deer management in US national agement unit that policy-making supervisors have designated
parks to control population irruptions and promote processes as “elk habitat,” (i.e., an area in which management is to give
contributing to ecological integrity, biologists William Porter priority to the needs and welfare of elk) and, thus, the action
and Brian Underwood noted that, “Whether we define eco- is a specific and particular enforcement of the “rule”
logical integrity in terms of species or processes, we must established by the policy: increase the production of
inevitably make a decision as to where in the irruptive resources that benefit elk.
sequence we choose to intervene. That choice represents a As benign and ordinary as this management action may
value judgment” (Porter and Underwood 1999:6). appear, it is fraught with ethical implications. First, what is
To explore the claim that management actions reflect the purpose of the management action? That is, what value
value judgments, let us begin by defining “management” in will be produced as a consequence of completing the action,
a conservation context as human action taken to remedy a in this case, burning sagebrush? Second, what is the defi-
deficiency in a system as informed by scientific assessment. ciency of the system that we are trying to remedy? Here the
Consider a very typical case of “ordinary” conservation man- deficiency is low forage production for elk. Is that deficiency
agement on public lands. A range manager, informed by best important, and how reliable is our scientific assessment of it?
available scientific data, decides to burn sagebrush (Artemisia Third, which stakeholders have legitimate interests in the
10.2 The Necessity of Value Judgments in Conservation 415
Fig. 10.2 A prescribed fire in sagebrush habitat in the Hart Mountain when to intervene in an ecological system, and therefore is fraught with
Wildlife Refuge in Oregon (USA). Even a seemingly “ordinary” man- ethical choices and considerations. (Photo courtesy of US Geological
agement decision represents a manager’s value judgment about how and Survey, public domain)
state of this system, and how will they be affected by this ethical implications. The questions and their arrangement in
management action? Every manager has duties, obligations, logical order are appropriate to every management action in
and trusts to fulfill to various stakeholders who have a vested conservation biology. Therefore, every conservation biolo-
concern in the state of the system being managed. Sometimes gist needs to understand how to categorize and evaluate
these duties are implicit and assumed, as with a manager in a different categories of values, as well as underlying
government agency with social and civic obligations to the assumptions in which ethical categories are embedded.
public. Sometimes they are explicit or even contractual, as in
the case of a manager who works for the members of a private
conservation organization, or an environmental consultant 10.2.2 Values and Ethics – Foundational
who is paid by a client to offer an informed recommendation. Definitions
Fourth, what kinds of values does the management action
endorse or enhance? What kinds of values does it neglect or Value refers to a general basis for an estimation of worth.
diminish? In this example, prescribed fire is likely to generate Values represent judgments of relative worth, merit, useful-
more forage for elk, but it will reduce habitat for species that ness, importance, or degree of excellence. Values can justify
are sagebrush specialists, like Brewer’s sparrow (Spizella concrete objectives, such as conserving biodiversity, but they
breweri) (Fig. 10.3) and sage grouse (Centrocercus are not the same as the objective. Ethics are systematic
urophasianus). Finally, when the action is completed and organizations of values that establish principles for conduct
the manager gives an account of what she did, will she be and behavior. Ethics reflect “practical philosophy,” an
able to defend her action and explain, to any audience, why ability – refined by education, training, and experience – to
she choose one action over another? choose between right and wrong. No set of scientific facts,
We can evaluate the process analytically and visualize it even scientific facts about endangered species or degraded
sequentially (Van Dyke 2005) (Fig. 10.4) to better grasp a ecosystems, by themselves, entails any statement of moral
logical way to think about this management action and its evaluation or moral prescription. That is, one cannot argue
from a descriptive premise (about what something is) to an

imperative conclusion (what ought to be done about a partic-
ular condition) because one cannot argue logically that
because something is (that is, because something exists) that
the same something is good (that it ought to exist). It is
common for people, including conservation biologists, to
make this mistake. The mistake is so common that
philosophers have given the error its own name, the natural-
istic fallacy: the error of arguing for what ought to be based
on how things are.
Conservation scientists gain knowledge of what “is” from
empirical studies. But conservation demands choices. Every
choice requires a decision about what is “good” or “best” to
achieve a given conservation goal. Conservation derives its
purpose by understanding its outcomes as expressions of
value. To satisfy the inherent drive for application intrinsic
Fig. 10.3 Brewer’s sparrow (Spizella breweri), a specialist of big to conservation biology and express management
sagebrush (Artemisia tridentata) in western North America. Prescribed applications in terms of norms, conservationist biologists
fire eliminates sagebrush, creating more favorable foraging conditions
for big game species like elk (Cervus elaphus), but reduced habitat for
must possess a coherent system of values they can universally
the Brewer’s sparrow. (Photo courtesy of Tom Koerner, US Fish and and persuasively articulate to anyone. If they fail to recognize
Wildlife Service) the necessity of coherent expressions of value, and fail to
affirm that statements of value are statements about truth,
about how things really ought to be, they will be left with no
arguments to offer except those that express conservation as
ratios of human economic benefits and costs. Indeed, many
conservation laws such as the US Endangered Species Act
explicitly state that economic considerations may not be used
to decide whether to adopt a particular action or policy, such
as listing a species as endangered or threatened (more in
Chap. 12). Value systems that can guide applications of
conservation are diverse, but persistent efforts to understand
such systems reveal the need to explain applications and
goals of conservation as more than personal preferences.
How can we begin to study (and, eventually, solve) this
kind of problem?
When describing natural systems, we often use words like
complexity, integrity, health, diversity, balance, stability, or
flourishing. These words, and others like them, are “cross-
over” words between science and ethics, between conserva-
tion and value. They can refer to facts about natural systems
that can be more or less measured, but they can also easily
slide into latent value judgments. Yet such words seem
necessary, because the values seem to emerge simultaneously
with the facts. Environmental ethicist Holmes Rolston III
expresses this recurring dilemma of conservation biologists
well. “For some, at least, the sharp is/ought dichotomy is
gone; the values seem to be there as soon as the facts are fully
in and both alike seem properties of the system. . .What is
Fig. 10.4 A schematic illustration of the logical flow of questions to be ethically puzzling, and exciting, in the marriage and mutual
asked in the ethical analysis of a management action, exemplified in this
case through evaluation of considering the decision to execute a pre-
transformation of ecological description and evaluation is
scribed fire on a winter used by an important game species, elk (Cervus that here an ought is not so much derived from an is as
elaphus) in the US state of Montana. (Figure design by F. Van Dyke) discovered simultaneously with it” (Rolston 1988:232).
10.2 The Necessity of Value Judgments in Conservation 417
10.2.3 How Values Inform Management contained statistically clustered groups of questions about the
respondent’s perceptions and attitudes toward wildlife.
The late Stephen Kellert of Yale University was a pioneer in Kellert was eventually able to construct a categorization of
developing methods through which human values and such responses into nine value categories (Table 10.1). By
attitudes toward wildlife could be assessed. In his book, carefully controlling for different group-specific variables
The Value of Life: Biological Diversity and Human Society, including age, gender, income, education, race, and rural or
Kellert explained why he began these studies. “I was primar- urban background, Kellert emerged with a comprehensive
ily concerned with the problem of how the effective manage- picture of how people in the United States viewed wildlife
ment of wildlife often seemed less a problem of manipulating and how such perceptions changed with sociological and
animals and their habitats than managing our own species’ demographic differences. Men showed greater inclination to
often callous and destructive disregard for much of the natu- value wildlife for utilitarian and dominionistic purposes,
ral world” (Kellert 1996:3). Because of his expertise in women for humanistic and moralistic purposes, patterns con-
investigating sociological aspects of human perceptions of sistent with the fact that men have traditionally made up the
wildlife, Kellert was sought out by the US Fish and Wildlife majority of membership in hunting and fishing organizations,
Service (USFWS). The agency needed his expertise because, women in animal welfare and protection organizations. Peo-
having traditionally managed wildlife resources on a sup- ple from rural residences or backgrounds, especially those
posed “value-free” approach, it was experiencing increasing that derived a living directly from commercial use of wild or
problems and conflicts with changing attitudes toward wild- domestic animals, had stronger orientation to utilitarian
life in the American public. As Kellert explained, “[Manage- values, while urbanites showed greater tendency to affirm
ment for sport hunting and fishing] had also led to a strong moralistic values. Education, however, was the single stron-
financial, political, and ideological dependence on sport gest attitude-shaping force. Kellert and his colleagues found
hunters and fishers. The price of such reliance had become that, the higher the level of education a person attained, the
a narrow management focus – and exclusion of most of the more likely they were to endorse ecologistic, scientific, mor-
public from the wildlife profession’s inner circles” (Kellert alistic, and humanistic values of animals (i.e., the more likely
1996:4). Kellert’s charge from the agency was to determine they were to express concern toward, affection for, interest in,
how the overall American public perceived wildlife, allowing and knowledge of animals) (Kellert 1996).
the USFWS to achieve “more equitable allocation of Kellert’s work provided a foundation for understanding
resources among users, a better basis for mitigating conflicts the different ways that human attitudes toward animals – a
among wildlife interest groups, ascertaining support for critical component of conservation – varied among people,
protecting and restoring rare and endangered wildlife, and what influenced that variation. By themselves, however,
educating the public about the value of wildlife and its Kellert’s categories were only descriptive markers. They
conservation, and more fully understanding trends in Ameri- provided no “ethic” of conservation for animal populations
can perceptions and uses of animals in the natural environ- or anything else. To move toward an ethic of conservation,
ment” (Kellert 1996:5). we must begin to examine why animals and other elements of
Over many years of research, Kellert and his colleagues nature might be valued, and what supports these value
accomplished their mission through sociological surveys that judgments.
Table 10.1 A typology of wildlife values and attitudes employed in assessing public attitudes toward wildlife
Type of value or
attitude Definition
Naturalistic Values that relate to enjoyment from direct contact with wildlife
Ecologistic Values associated with the importance of a species to other flora and fauna and to the maintenance of ecosystem
processes
Moral Values associated with inherent rights or spiritual importance of species
Scientific Actual or potential value associated with a species’ contribution to enhancing human knowledge and understanding of
the natural world
Aesthetic Values associated with the species’ possession of beauty or other perceived qualities admired by humans
Utilitarian Values associated with species as sources of material benefit or use
Dominionistic Values associated with the mastery and control of animals, typically through sport
Negativistic Attitudes associated with the avoidance of animals because of dislike or fear
Neutralistic Attitudes associated with the passive avoidance of wildlife because of lack of interest
Theistic Values associated with the belief that a supernatural deity or force creates, sustains, and values wild species
Developed from concepts in Kellert, S.R., 1991. Japanese Perceptions of Wildlife. Conservation Biology 5, 297–308

A scientist wrote to us saying, “People choose to study
snakes because they love snakes.” He went on to argue
that such affection generated its own “mission,” and
such mission generated its own advocacy for snake
conservation. Put another way, this scientist was saying
that people choose to study snakes because they respect
snakes as having a good of their own. Do you think that
personal affection or respect is a sufficient basis for
conservation mission and advocacy or an adequate
frame of reference for discourse with other scientists
and the public? Fig. 10.5 Categories of value and their relations. (Diagram by M. J.
Bigelow)
acknowledges and requires recognition of both instrumental

and intrinsic values of species, and, with regard to species,
the relational aspects of their value to the world around them,
10.3 How Do We Categorize Conservation including human beings. Both instrumental and intrinsic
Values? values therefore merit further analysis.
10.3.1 An Overview of Value Categories

10.3.2 Instrumental Values
There is no single universally accepted method of
categorizing conservation values, but we will begin with a 10.3.2.1 General Considerations
dichotomy between instrumental and intrinsic values. All human cultures have been and continue to be sustained
Instrumental values measure the usefulness of a creature or through natural capital, including goods and services derived
object in meeting a need or providing a service to another, from living organisms, so instrumental values play a promi-
usually a human, and thus facilitating human welfare or nent role in any discussion of the value of non-human
happiness. Intrinsic values reside within an object itself. In creatures. The most common method of estimating the instru-
other words, something has intrinsic value if it is “valuable in mental value of goods is through economic evaluation. Biotic
and for itself – if its value is not derived from its utility, but is resources supply all food for humans and, directly or indi-
independent of any use or function it may have in relation to rectly, most of our fuel and medicines. Plants are important
something or someone else (Callicott 1986:140, emphasis sources of human clothing and structural materials, and are
his). Put another way, intrinsic value in nature, or “natural used to beautify personal property, enhance land values, and
intrinsic value,” is value that is present in “some natural reduce soil erosion. Animals provide services through their
occasions without contributory human reference” (Rolston labor in transportation or other work efforts, as sources of
1986:110). We can further subdivide intrinsic value into food, and as companions to human beings. In human enter-
“non-relational” versus “relational” forms of intrinsic value. tainment and recreation, plants and animals play significant
For example, if I choose to wear a diamond on my finger, roles in increasingly varied and economically important
store it in a safety deposit box in a bank, or carry it with me ways. The role of economics is so important in conservation
whenever I take a trip, the intrinsic value of the diamond that it will receive its own chapter in this book (Chap. 11), but
remains unchanged. In contrast, the intrinsic value of a living the role of economics in conservation decision making is
organism requires relationships for that value to be expressed significant and deserves some attention here.
and to continue. An endangered species like, for example, the Many biotic resources are scarce resources. In these cases,
mountain gorilla (Gorilla beringei beringei) may have a their use can only be increased at the cost of foregoing
unique good of its own, but it will need fruit and vegetable something else that is valued (Randall 1986). Biotic
matter to eat, forests to live in, and a mate to raise offspring resources are potentially renewable resources, yet also poten-
and pass on its genes for that value to be fully expressed and tially degradable and exhaustible. Even though the goals of
continued through time. Within the larger categories of economics and conservation can appear at odds with one
instrumental and intrinsic values, specific sub-categories another in specific cases, they have fundamental interests in
exist for natural objects in general, and for species in particu- common, especially regarding species preservation. As econ-
lar (Fig. 10.5). The practice of conservation biology omist Alan Randall stated, “Since species survival is a
10.3 How Do We Categorize Conservation Values? 419
precondition for the use of the species as a resource, the goods A, B, and C if the resource is being used to produce
preservation problem in principle precedes all other biotic good D.
resource issues” (Randall 1986:79). The other four categories describe different kinds of non-
Economists attempt to estimate the market value of use values. Option value refers to the value of a resource’s
resources, including biotic resources, through evaluation of expected future use (i.e., what a person would be willing to
resource scarcity (supply) and the amount of satisfaction the pay to guarantee that the resource would be available for
resource provides for humans (demand). Historically, eco- future use). Quasi-option value is the value of preserving
nomics has been used in making decisions in conservation, options, given an expectation of growth in knowledge that
even when such decisions involve judgments about what is might lead to a future, but as-yet undiscovered or unrealized
“right” and “wrong.” Originally a branch of moral philoso- use of the resource. Quasi-option value can be conceived as a
phy that arose in the nineteenth century, economics only later kind of “speculation value” for what an investor might pay to
emerged as an independent and, eventually, more mathemat- preserve a resource, such as a rare tropical plant, given the
ical, discipline (Kelman 1986). Although no longer consid- expectation of increasing growth in knowledge of medical
ered a realm of moral knowledge, contemporary economic applications of all plant species generally. Bequest value is
evaluations influence what society believes is “right” in the value of knowing that something is preserved for future
decisions on conservation. generations. Existence value is the value of knowing some-
The economic valuation of biotic resources often can be thing exists at the moment (Randall 1986). In other words,
done through normal market processes. In economics, mar- the object is not something you are going to use now or in the
ket goods can be exchanged through some form of standard future, but you derive pleasure and satisfaction simply from
currency in an economic market. Some biotic resources, such knowing that it is there.
as game animals, timber, or rangelands, can be treated and
valued as market goods, and their market value assigned with 10.3.2.2 Determining Attitudes Through
relative precision. This valuation is easiest when the value of Sociological Surveys
the resource is measured in utilitarian terms, usually as some One method researchers use to assess attitudes of people
type of commodity. In that case, biotic resources often can be toward conservation is through surveys, as Kellert did in
treated as private goods (i.e., goods that can be bought, sold, determining human attitudes toward wildlife. Surveys can
and enjoyed solely by the buyer and seller). Natural resources consist of “closed-ended” questions (in which the respondent
also can serve utilitarian values as public goods, goods that selects a particular answer or a numerical value that best
are accessible to all and provide benefit to all, usually with represents her answer) (Table 10.2) or in-depth interviews,
the cost shared among the beneficiaries, such as clean water. which permit more complex responses to open-ended
Economic evaluation is easiest when biotic resources can questions. Survey design is a professional discipline within
be valued as private goods, becomes more complex when sociology and psychology. Its details are beyond the scope of
they are valued as public goods, and still more complicated the chapter; but some foundational principles are important to
when future as well as present values are considered. But know and follow: (1) questions and potential responses in the
these considerations also are part of economic valuation. In survey must be carefully worded according to strict protocols
economic terms, five categories of value help to refine the to avoid confusion and clarify categories of response; (2) the
assessment. Use value, the value derived from the actual use sample of individuals surveyed must be representative of the
of a resource, is the easiest to measure and most amenable to population and sufficiently large to avoid problems
evaluation by market forces. For every use, there is a unique associated with random error; (3) if subgroups of the popula-
set of opportunity costs which represent costs, or losses tion differ in important characteristics, representation of the
associated with the inability to use the resource to produce subgroups must be comprehensive and sufficiently large to
Table 10.2 Some examples of questions and their “closed-ended” responses in a survey of human attitudes toward wildlife
Question
type Examples Response
Closed- If a conservation biologist aspires to be effective in relating scientific findings to conservation policy, it is necessary T/F
ended that the basis of conservation ethics be understood.
The number of animals is more important than the genetic diversity of the group. T/F
The value of an organism should be decided by its supply and demand. T/F
Open- To what extent should ethics play a role in the process of developing conservation policy? Variable
ended How important is the genetic diversity of a population?
What criteria should be used to assign value to an organism?
Table designed by M. J. Bigelow based on concepts from Kellert, S.R., 1991. Japanese Perceptions of Wildlife. Conservation Biology 5, 297–308
ensure accurate representation, and (4) if some individuals do who use them (Dustin and McAvoy 1982). Such knowledge
not respond to the initial survey there must be follow-up can assist conservation biologists in determining what public
procedures to reach them. This follow-up is intended to attitudes are toward wildlife, conservation, and natural areas.
solve the problem of “non-response bias,” because In addition, user satisfaction surveys may be valuable in
non-respondents often have different characteristics than ini- designing public relations campaigns to win support for
tial respondents. conservation goals. However, survey, interview and user
Coupled with in-depth interviews, survey data may reveal satisfaction data, by themselves, do not evaluate the validity
not only attitudes of people toward wildlife and other biotic of the attitudes that they identify, nor do they help conserva-
resources, but also the cultural basis and background of such tion biologists argue persuasively for alternative attitudes or
views, as was demonstrated in Kellert’s study. Throughout ethical perspectives.
the world, data from multiple studies reveal that humans are
most concerned for creatures that are large, aesthetically 10.3.2.3 Tools for Economic Valuation: The Role
attractive, phylogenetically similar to humans, and regarded of Cost-Benefit Analysis
as possessing capacities for feeling, thought, and pain The most common, and sometimes legally prescribed, tool
(Kellert 1986). Such species represent what have been called for determining the value of biotic and ecological resources
phenomenologically significant animals (Shepard 1978), or, in their natural state versus their value after detrimental acts
more commonly today, charismatic megafauna. Given such of development is cost-benefit analysis (CBA). Conceptually
human attitudes, it is no wonder that animals in this group are simple to imagine, but often practically difficult to achieve,
often chosen as emblems for major conservation cost-benefit analysis assesses benefits of a particular action
organizations such as the World Wildlife Fund (giant and compares them to the action’s environmental costs. The
panda, Ailuropoda melanoleuca) (Fig. 10.6), as preferred alternative is one in which benefits most outweigh
representatives for government agencies like the US Forest costs. Cost-benefit analyses are often used in environmental
Service (Smokey Bear), or as a country’s national symbol impact statements (EIS) required under the US National
(for the US, the bald eagle, Haliaeetus leucocephalus). Environmental Policy Act. “Costs” of the action described
Survey data also can be used to assess human relationships in the EIS typically include both short- and long-term envi-
with nature in terms of personal experience. One of the most ronmental consequences associated with it. “Benefits” to
common measures is “user satisfaction,” a metric which species, biodiversity, or ecosystem preservation must include
determines the level of personal satisfaction an outdoor rec- both instrumental values – such as game, water, range, and
reationist experiences in a particular recreational activity. The timber – and non-instrumental values – such as visual and
higher the satisfaction, the more the experience met or scenic appeal, religious significance, or historic importance.
exceeded the expectations and desires of the recreationist. Cost-benefit analyses also may be associated with the pro-
This measure has been used to evaluate the quality of natural posed listing of a species as threatened or endangered, with
environments as well as the preferences and values of those broad changes in environmental or conservation policy, or
with the effects associated with new conservation legislation.
Government bureaucrats, private consultants or corporations,
non-governmental conservation organizations, or other
individuals or groups that have a vested interest in the valua-
tion of biotic resources might perform the actual analysis.
CBA is appealing to decision makers because it attempts
to translate all values associated with a decision into a com-
mon denominator – like market value in dollars – so that
diverse entities (e.g., timber values and wildlife values) can
be directly compared. Cost-benefit analyses are also
appreciated for their thoroughness. They force decision
makers to identify and evaluate the value of all entities
affected by their decision. In a pro-development society,
cost-benefit analysis is attractive because it places the burden
of proof on those who value the preservation of biotic
resources rather than on proponents of development. In
Fig. 10.6 The giant panda (Ailuropoda melanoleuca), symbol of the CBA, defenders of biodiversity must show that species pres-
World Wildlife Fund and a classic example of a “phenomenologically
significant animal” (Shepard 1978) that evokes strong feelings of iden-
ervation is at least as economically valuable as a proposed
tification, affection, and concern in humans. (Photo courtesy of J. Patrick development that is designed from the outset for market
Fischer, under CC-BY-SA-3.0) valuation and consumption.
10.4 Moral Value: Assigning Intrinsic Values in Conservation 421
Criticisms of CBA are numerous. Some acts that might be from it in the future to you ought not to overfish this species
judged morally wrong might have high economic benefits so that its kind does not perish from the Earth.
and low costs, whereas some acts considered morally right These considerations bring us to another critical question.
can have low benefits and high costs. In addition, CBA uses Are values intellectual abstractions that people invent and
currencies most appropriate to private transactions of eco- manipulate, or are they external realities that people recog-
nomic goods as guides for public policy, thus equating pri- nize and respond to? Some ethicists assert that intrinsic value
vate preferences with cultural and social values. CBA is an illusion, that all values are anthropocentric, residing in
assumes that preserving a species should be valued for its human consciousness. Therefore all values are human
human benefits rather than as an act of moral obligation. preferences subject to economic evaluation. A thing is good
Thus, CBA fundamentally asserts that all environmental when it tends to increase utility and bad when it tends other-
values are what moral philosophers would call demand wise. But not all agree with this perspective.
values– the idea that an environmental entity has value if Most ethicists would admit that the locus of all value is
and only if a person has a felt preference for that entity human consciousness, and some would say that the percep-
(Sarkar 2012:42). This perception reveals why the distinction tion of value in nature is an observer-dependent, secondary
between human benefits and moral obligation is critical. We quality that arises solely out of the observer. But others would
will examine this distinction in more detail through examina- argue that, although the locus of all value is within the human
tion of an oft-used method, contingency valuation, for deter- consciousness, the source of all value is not (Callicott 1986).
mining market values of non-market goods, like endangered Ethicist Holmes Rolston III explains the concept by saying
species, in our next chapter on conservation economics. But that the human self “has a semipermeable membrane” of
to go further now, we examine questions related to a different value perception. A person, or “self” in Rolston’s words,
value category, that of intrinsic value. perceives something of the “natural” value (ENV) of an object
in nature and this produces an internal recognition of that
value, an “experiential value,” (EEV) in the observer
10.4 Moral Value: Assigning Intrinsic Values (Fig. 10.7). Rolston gives an example using a woodland
in Conservation wildflower, the trillium (Trillium spp.), and the experience
of enjoying its beauty.
10.4.1 Where Does Intrinsic Value Reside? . . .there is some ought-to-be beyond the is, and so the plant
grows, repairs itself, reproduces, and defends its kind. If, after
A fundamental question in environmental ethics is whether enjoying the Trillium in a remote woods, I step around it to let it
we are examining an ethic about the environment or an ethic live on, I agree with this defense, and judge that there is an
intrinsic objective value, valued by me, but for what it is in itself
of the environment, a distinction Holmes Rolston III stresses (Rolston 1986:111).
as the difference between secondary environmental ethics
and primary environmental ethics (Rolston 1988). The first
sees the environment as an important stage upon which the
drama of human choice is played out, but ethical decisions
are for and about people, and environmental entities are
means to achieve correct ethical decisions about humans.
What we would call secondary environmental ethics are
really classical forms of ethics, for example, utilitarianism,
applied to environmental limitations. For example, you
should not overfish this species if you want to continue to
derive food and income from it in the future. In this view, all
of nature’s goods, including all species, are human goods. In
contrast, a primary environmental ethic is one in which
environmental entities themselves, such as non-human spe-
cies, ecosystems, or even “the land,” are treated as moral
subjects. Such entities do not make moral choices and are not,
therefore, moral agents, but as moral subjects, humans can
treat them rightly or wrongly in an ethical sense. These Fig. 10.7 A schematic illustration of the relationship between the
natural entities possess a “good of their own” that should be perception of intrinsic value by a human observer and the locus of
acknowledged and protected, regardless of its costs or such value in a natural object. In the diagram, object E possesses a
natural or intrinsic value (Env) which is perceived by a human observer
benefits to humans. In a primary environmental ethic, our (“self”) and internalized as a person experience of apprehending the
example would be changed from you should not overfish this value of the object E, an “experienced value” (Eev) of E. Based on
species if you want to continue to derive food and income concepts from Rolston 1986. (Original drawing by M. J. Bigelow)
Every organism, notes Rolston, is a unique “genetic set,” that value, and what is its source? Moral extensionism, the
which the organism protects, “. . .the genetic set is a norma- process of transferring moral considerations traditionally
tive set. . .it distinguishes between what is and what ought to reserved exclusively for humans to other creatures or to all
be. . .The genome is a set of conservation molecules. A life is of nature itself, is one foundation for establishing intrinsic
spontaneously defended for what it is itself” (Rolston value for non-human life.
2012:97, emphasis his). In such defense, the organism
manifests the reality that it has a good of its own. As Rolston
explains, “. . .in the language of conservation biology, a plant 10.4.2 Establishing Intrinsic Value Through
is already engaged in the biological conservation of its iden- Moral Extensionism
tity and kind, long before conservation biologists come on the
scene. What conservation biologists ought to do is respect 10.4.2.1 Biophilia- Moral Extensionism Through
plants for what they are in themselves – projects in conserva- Genetic Heritage
tion biology. The point of such thinking about plant informa- Moral extensionism is the idea that moral standing ought to
tion, about genetic information, is that we should value life” be conveyed to things (animals, plants, species, the earth)
(Rolston 2012:106–107). And the organism values more than that traditionally are not thought of as having moral stand-
itself as an individual. It values its kind. “A female animal ing. One justification for moral extensionism is genetic pre-
does not have mammary glands,” notes Rolston, “nor a male disposition, formalized in the so-called Biophilia Hypothesis,
testicles, because the function of these is to preserve its own first articulated by biologist E. O. Wilson. Wilson defined
life: these organs are defending the line of life bigger than the “biophilia” as the innate tendency to focus on life and lifelike
somatic individual. The locus of the value that is defended processes. . .to the degree that we come to understand other
over generations is as much in the form of life, since the organisms, we will place greater value on them and on
individuals are genetically impelled to sacrifice themselves in ourselves” (Kellert and Wilson 1993:4–5). Biophilia is not,
the interests of reproducing their kind. . .A species is a living says Wilson, a single instinct, but a complex of “learning
historical form. . ., propagated individual organisms, that rules” that can be separated and evaluated analytically. Bio-
flows dynamically over generations” (Rolston 2012: philia (literally, “love of life”), asserts Wilson, is genetically
127–128). Thus, the attribute of valuing (i.e., the thing that programmed into us based on our evolutionary past, and
is valued) is present independent of human presence near it or holds high fitness value. As Stephen Kellert expressed it,
recognition of it, and that value persists through time as a “the [biophilia] hypothesis suggests that the widest valua-
species. This is the very definition of intrinsic value, But the tional affiliation with life and lifelike processes. . .has
attribution of valuing (the act of giving value) comes only conferred distinctive advantages in the human evolutionary
through humans, what Rolston calls “value ignition” struggle to adapt, persist, and thrive as individuals and as a
(Rolston 2012:114). species” (Kellert and Wilson 1993:42).
Our experience and perception “let us in on” the presence One line of evidence for the adaptive value of the biophilia
and qualities of value in a natural object. And our conscious- impulse can be seen in studies of interaction between children
ness is then able to “share” with the object the qualities of and nature, specifically children who suffer from Attention-
value that were always present in it, but which we now Deficit Hyperactive Disorders (ADHD) (Information Box
perceive and to which we can now respond. “All natural 10.1). Wilson argues that the biophilic response of the
science,” notes Rolston, “is built on the experience of nature, ADHD children is in fact common to all humanity, and
but this does not entail that its descriptions, its “facts,” just represents “an advance in moral reasoning. . .to create a
are those experiences. Valuing could be a further, non-neutral deeper and more enduring environmental ethic. . .The only
way of knowing about the world” (Rolston 1986:104). In way to make a conservation ethic work is to ground it in
fact, it may be that ability to perceive the intrinsic value of ultimately selfish reasoning – but the premises must be of a
other species is not only appropriate for humans, but an new and more potent kind. . .a deep conservation ethic [based
important distinction of being human. on] biophilia. . .The more the mind is fathomed in its own
Most conservation biologists act as if they appreciate an right, as an organ of survival, the greater will be the reverence
intrinsic value of non-human species, and demonstrate their for life for purely rational reasons”(Wilson 1984:126).
appreciation with a career of commitment to the welfare of Biophilia, then, is an expression of genetically
other creatures. However, if the value inherent in these programmed self-interest, not a means of making moral
perceptions is to be shared with others and transformed choices. Rolston explains why this is not a satisfactory
from personal preference to real “knowledge,” it must be ethic. “The theory is not revealing anything about values in
expressed and explained in ways that can be understood by nature; it is just confusing us. Selfishness is indeed real. . .but
others. If species possess intrinsic value, and if such value we do not want to speak as though animals and genes were
should change our behavior toward them, what is the basis of ethical agents in conditions of only superficial similarity”
(Rolston 1993:393). “To try to see all ethics as nothing but

extrapolated selfish genes,” Rolston argues, “might stunt Information Box 10.1 (continued)
humanity because it fails to realize the genuine human behavior. (2) When an ADHD child was given oppor-
transcendence – an overview caring for others. . . Humans tunity to interact with an animal instead of just
can get “let in on” more value than any other kind of life. watching it, changes in behavior were greater and
They can share the values of others and in this way become more persistent. (3) Positive speech and non-verbal
consummate altruists” (Rolston 1993:409). expression were facilitated by the presence of animals.
Biophilia may be a real genetic predisposition, but it (4) Rules explained to ADHD children relating to
cannot function as an operational conservation ethic. The animals were received as reasonable and carefully
strength of biophilic impulse is variable in individuals, and, followed, whereas rules of behavior in the classroom
we must ask, how rationally trustworthy is the impulse? If were rejected as arbitrary (Katcher and Wilkins 1993).
formed under environmental conditions of evolutionary past Katcher and Wilkins explain . . .hyperactive children
different from the human condition today, how do we know who were taught how to care for animals used their
that the biophilic impulse is morally “right”? Biophilia can experience to learn biology and to learn how to
offer a foundation to love life and all its processes and learn. . . . Nature was not merely subject matter; it
variety, but it cannot direct us in specific and difficult moral was the world common to teacher and student for
choices that are sure to come in real conservation effort. Can which both had to assume responsibility. With that
we make moral extensionism more specific? kind of teaching, the innate tendency to continue a
dialogue with other kinds of life can be joined to the
moral agenda of preserving that life (Katcher and
Information Box 10.1: ADHD Children
Wilkins 1993:193).
and Animal Care
Aaron Katcher and Gregory Wilkins studied children
with Attention-Deficit Hyperactive Disorders (ADHD) 10.4.2.2 The Rights of Nature – Moral
who were permitted to be part of a program of caring Extensionism to Individual Creatures
for zoo animals. To continue in the program, the chil- Moral extensionism in environmental and conservation ethics
dren had to be gentle with the animals, learn the proper becomes more focused when it moves from the status of
ways of holding and caring for them, spend time with genetic predisposition in human chromosomes to intentional
the animals caring for their needs, speak softly when moral consideration of non-human species. The most direct
around the animals, and avoid using speech that expression of such consideration is found in considerations of
devalued the animals or people around them. These “animal rights.” Ethics of animal rights emphasize that ethi-
rules demanded high levels of motor inhibition and cal extension should not be limited to only rational creatures
impulse control, things hard to achieve in ADHD chil- (Can it think what we think?), but to sentient creatures who
dren. Yet, compared to a control group of ADHD are capable of feeling pain and suffering (Can it feel what we
children who went to the zoo, saw the same animals, feel?). With this change of emphasis the operational criteria
but did not learn to care for them, differences were for ethical consideration is also altered, changing from “Can
dramatic. For ADHD students in the animal caring it reason?” to “Can it suffer?” Under this new criterion, the
program, the animals caught and held the children’s array of morally considerable species expands to include at
attention, reducing problems associated with their least birds and mammals, if not all vertebrates, and poten-
attention-deficit disorders. Some ADHD students who tially other forms of life. In its purest, or, some would say,
had made no progress in learning in a traditional class- most extreme form, an ethic of animal rights would forbid
room for 4 years were able to learn complex tasks in any human action that caused a sentient species to suffer or
animal care rapidly at the zoo. Even during free time at feel pain.
the zoo, the ADHD children spent most of that time in At first glance, ethics of animal rights might not seem
the company of the animal assigned to them. Many of relevant to conservation. Conservationists are concerned
the children in the zoo study had to be restrained or with populations, species, communities, habitats, and
medicated in a traditional classroom, but not when ecosystems, not individual animals. But a closer look brings
engaged with their animals. Katcher and Wilkins relevance into focus. The acts of conservation scientists affect
reached four conclusions: (1) Animals brought into individual animals, even in the most benignly intended
regular contact with children were powerful efforts. A conservationist attempting to find a cure for a
re-enforcers of sustained attention and controlled disease affecting thousands of individuals in a wild popula-
tion might, to understand the pathology of the disease, choose
(continued)
Fig. 10.8 The Kirtland’s warbler (Setophaga kirtlandii) (left), histori- found in areas where the Kirtland’s warbler was nesting. Such action did
cally one of North America’ rarest songbirds, was endangered, in part, not endanger the cowbird as a species, but thousands of cowbirds died to
because of nest parasitism by the brown-headed cowbird (Molothrus restore the warbler’s population. (Kirtland’s warbler photo courtesy of
ater) (right). The US Fish and Wildlife Service initiated a program of J. Trick, US Fish and Wildlife Service. Brown-headed cowbird photo
cowbird “control” which consisted of trapping and then killing cowbirds courtesy of the Missouri Department of Conservation)
to infect a small number of individuals in a laboratory to learn Some species, like the California condor (Gymnogyps
to cure it. Many, perhaps all, of the infected animals might californianus) and black-footed ferret (Mustela nigripes),
die in the effort to gain this knowledge. have been saved by capturing all remaining wild individuals
Other ethical issues affecting individual animals may be and breeding them in captivity. Since these “rescues,” new
less obvious, but even more pervasive. Sometimes, to save an populations have been returned to the wild. But some
endangered species, a more common competitor, predator, or individuals died in captivity, and all of the captured
parasite of the endangered population must be prevented individuals were deprived of the freedom they had formerly
from interacting with it, usually by killing the competitor, known. Recovery did not come without cost. Modern zoos
predator, or parasite. A formerly endangered North American are important centers of conservation efforts, not only in
songbird, the Kirtland’s warbler (Setophaga kirtlandii), was captive breeding and in, some cases, preserving the last
endangered, in part, by nest parasitism of the brown-headed individuals of extremely rare species, but as centers of con-
cowbird (Molothrus ater) (Fig. 10.8). To save the warbler, servation education intended to make their visitors more
the US Fish and Wildlife Service trapped and then killed sensitive to and supportive of the plight of wild species and
cowbirds in areas where the Kirtland’s warbler was nesting. the global preservation of biodiversity. But what of the indi-
Such action did not endanger the cowbird as a species, but vidual animals that provide the public “education,” or, to be
thousands of cowbirds died to restore the warbler’s popula- more honest, “entertainment?” Isak Dinesen, author of Out of
tion. In capturing and banding songbirds or waterfowl, mark- Africa, relates his experience of seeing two giraffes being
ing mammals with ear tags, placing radio transmitters on transported to a zoo on a ship.
vertebrates, or in-field sterilization of individuals of a com- Upon the deck, there stood a tall wooden case, and above the
mon species or hybrid to prevent it from breeding with an edge of the case rose the heads of two Giraffes. . . The Giraffes
endangered species (like the red wolf, Chap. 5), every expe- turned their delicate heads from one side to the other, as if they
rienced biologist knows that there is a risk of injury or were surprised. . .They could not know or imagine the degrada-
tion to which they were sailing . . .nor the terrible boredom in a
mortality to individual animals in the process of capturing, world in which nothing is ever happening. Crowds. . .will be
marking, or performing surgery, in addition to unexpected coming in from the wind and sleet of the streets to gaze on the
complications following its release, no matter how routine Giraffes, and to realize man’s superiority over the dumb
the procedure or practiced the biologist. And even “passive” world. . .In the long years before them, will the Giraffes some-
times dream of their lost country? (Dinesen 1989:287–288).
studies involving only observation can cause animals physi-
ologic stress, changes in behavior and habitat use, increased This story poignantly illustrates the ethical conflict of the
vulnerability to predators, or abandonment of nests or young. modern zoo which, in the words of Stephen Kellert, seems
Despite every precaution, studying animals sometimes caught “between the goals of mass entertainment . . .and, on
results in suffering or death to the animals studied. the other [hand], public education and conservation” (Kellert
1996:85). Modern zoos often refer to themselves as “arks” of for humans who safeguard their interests to speak for them, to
conservation, alluding to the biblical story of Noah (Genesis act “at their behest.” Such thinking has legal precedent. If a
6–9) who preserved every kind of creature with himself and child’s parents die and leave the child a large fortune, the
his family in the midst of a great flood. As Holmes Rolston III child will not be able to “issue commands” to its lawyers
has pointed out, the animals in the ark got off. In zoos, they about how the money should be used or invested. But if the
never will. So Rolston notes perceptively that, in this context parents have designated a guardian, someone who “speaks”
“ark is a euphemism for a prison” (Rolston 2012:84). for the child, the guardian may take legal action on the child’s
Whether we are concerned for the welfare of animals in behalf. So legal expert Christopher Stone argued in his clas-
zoos or in research, no conservation biologist should ever sic work, Should Trees Have Standing?, that conservation
say that concern for the welfare of individual animals does organizations can serve a similar legal role as “guardians” of
not matter. It was public concern over the harvesting of natural entities like forests, lakes or mountains (Stone and
whales and the mortality of dolphins killed in tuna fishing Hardin 1974). In fact, Stone’s argument for giving rights to
that led to legislation, technology, policy and practices that natural objects was provoked by an actual legal case, Sierra
have reduced injury and mortality from these activities Club versus Morton. In 1971, the Sierra Club sued the United
(Chap. 12). States Department of the Interior (Rogers C. B. Morton was
Concern for animal rights, welfare, and humane treatment then US Secretary of Interior) to halt the development of a ski
is now a matter of corporate and institutional concern in all resort by the Disney Corporation in the Mineral King Valley
forms of animal research, including conservation, through the of California (Fig. 10.9). The case went all the way to the US
work of oversight committees on animal care and use, which Supreme Court, where the Court ruled against the Sierra Club
now exist in all major universities, organizations and because the club could not show a vested interest in the
agencies involved in research using animal subjects. Such Mineral King Valley that would be harmed by development.
review of and intentional accountability for proposed studies The Sierra Club lost the case, but their lawsuit sparked the
is one of the most important means of considering the relative development of new legal and ethical theory regarding natu-
weight of risks and benefits to animals used in or affected by ral objects, beginning in the Supreme Court itself. Writing the
research in conservation. Ultimately, however, it is the indi- dissenting opinion for the justices who disagreed with the
vidual researcher, not the committee, who will play the most Court’s ruling, Justice William O. Douglas asserted,
important role in ethical decisions in conservation research Inanimate objects are sometimes parties in litigation. A ship has
and practice. At both professional and personal levels, con- a legal personality, a fiction found useful for maritime purposes.
servation biologists must weigh the risks of harm, suffering, The ordinary corporation is a “person” for purposes of the
or death their study might bring to an animal compared to the adjudicatory processes, whether it represents proprietary, spiri-
tual, aesthetic, or charitable causes. So it should be as respects
good it might do for its population or species. valleys, alpine meadows, rivers, lakes, estuaries, beaches,
ridges, groves of trees, swampland, or even air that feels the
10.4.2.3 Extending Legal Rights to Natural Objects destructive pressures of modern technology and modern life. . . .
Consideration of moral rights for individual animals leads The voice of the inanimate object, therefore, should not be stilled.
That does not mean that the judiciary takes over the managerial
naturally to consideration of legal rights for individual functions from the federal agency. It merely means that before
animals, as well as for non-living natural entities, including these priceless bits of Americana (such as a valley, an alpine
“land.” Historically, legal rights have been limited to human meadow, a river, or a lake) are forever lost or are so transformed
beings, but precedents began to arise in the nineteenth cen- as to be reduced to the eventual rubble of our urban environ-
ment, the voice of the existing beneficiaries of these environmen-
tury when corporations began being recognized under law as tal wonders should be heard.
the legal equivalent of “persons” to whom rights, as well as
obligations, could also be assigned. Over time, an evolution
of thinking in the legal profession eventually established that The opinion of Justice Douglas, the writings of Stone and
something had “rights” if it met three criteria. (1) Action other legal scholars, and further aggressive legal action by the
could be taken at its behest. (2) If the entity (in legal terms, Sierra Club and other environmental organizations were
the “plaintiff”) sues for its rights in court and such rights are important stimuli to the development of a more inclusive
upheld, the court must take its suffering into account. (3) In form of moral extensionism for nature that has become
granting relief, the relief must run to the benefit of the known as “deep ecology.”
plaintiff.
“Behest” is an unfamiliar word in conversation, but an 10.4.2.4 Intrinsic Value Through Identification
important one in law. It means “at or upon request or urgent with Nature – Deep Ecology
command.” Natural entities like forests, lakes, or entire The term deep ecology was first coined by the Norwegian
ecosystems cannot make requests or give commands, but philosopher, Arne Naess in his 1973 article, “The Shallow
some legal scholars have argued that it should be permissible and the Deep, Long-Range Ecology Movements.” Drawing
Fig. 10.9 The valley of Mineral King in the US state of California. against the Sierra Club because the club itself could not show a valid,
Here the Sierra Club sued the Walt Disney Corporation to stop their vested interest in the Mineral King Valley. (Photo courtesy of the
development of a ski resort. The US Supreme Court ultimately ruled Sequoia and Kings Canyon National Parks, US National Park Service)
on current and past developments in ecology, law, and ethics, they often describe the core beliefs of deep ecology in
like those exemplified in Sierra Club v. Morton, Naess statements of contrast:
argued that current approaches to ecological problems were
too narrow in their scope and too dependent on technology in • Harmony with nature v. dominance over nature
their prescriptions. Such an approach, argued, Naess, did not • Biocentric equality – all nature has intrinsic worth
reach the heart of human behavior toward the environment. v. nature as resource for humans
He proposed an ethical paradigm of “deep ecology” that • Simple material needs v. material and economic growth
would, in the words of Deep Ecology advocates Bill Duvall for growing human population
and George Sessions, go “beyond a limited piecemeal shal- • Earth “supplies” limited v. belief in ample resource
low approach to environmental problems” and “articulate a reserves
comprehensive religious and philosophical worldview. The • Appropriate technology and non-dominating science
foundations of deep ecology are the basic intuitions and v. high technological progress
experiencing of ourselves and Nature which comprise eco- • Doing with enough and recycling v. consumerism
logical consciousness” (Devall and Sessions 1985:65). • Minority tradition and bioregionalism v. national/
Deep Ecology arose in large part in protest over the centralized economy
perceived capitulation of ecological “experts” to and with
the very practices and systems that were destroying the These principles are not unique to Deep Ecology. In fact,
earth. As philosopher Theodore Roszak lamented about the because Deep Ecology is so eclectic in its sources of ideas
problem of using “expert testimony” to solve conservation and inspirations, it can be difficult to determine what truly
problems, “While undeniably well-intentioned and capable defines it as a philosophical or ethical position. As its own
of stopgap success on specific political issues, it leaves founder, Arne Naess admits, “It is a characteristic feature of
wholly untouched the great cultural questions of our times. deep ecological literature that it contains positive reference to
It does not challenge the universally presumed rightness of a formidable number of authors belonging to different
the urban-industrial order of life.” (Roszak 1972:26). Deep traditions and cultures” (quoted in Devall and Sessions
Ecologists see themselves as those who protest against what 1985:226). Despite these difficulties, Deep Ecology and its
they describe as a “dominant Western worldview” that proponents are a persistent voice in discussions of contempo-
creates a divide between humans and nature. Therefore, rary conservation ethics, law, and policy. Its adherents
express their ideas in many ways. For example, the environ- spots, their continued existence in a natural state” (Leopold
mental group, Earth First! has been described as a “deep 1966:240). Leopold expressed the essence of his ethic in a
ecology direct action group” (Devall and Sessions 1985: summary moral maxim: “A thing is right when it tends to
257), shown by its willingness to damage property or even preserve the integrity, stability, and beauty of the biotic
risk human life in defense of nature. The contributions of community. It is wrong when it tends otherwise” (Leopold
Deep Ecology borrow from and contribute to various ethical 1966:262).
traditions in environmental ethics today. We will see these Leopold saw value in every species because each species,
more clearly as we go further. in his view, contributed to ecological function. Contrasting
ecological value to economic value in his own state of
10.4.2.5 Intrinsic Value Through Ecocentrism Wisconsin, Leopold wrote, “One basic weakness in a conser-
Aldo Leopold is known for and credited with the formulation vation system based wholly on economic motives is that most
of “the land ethic,” an ethical paradigm explaining how members of the land community have no economic value.
humans ought to relate to their non-human environment. A Wildflowers and songbirds are examples. Of the 22,000
career forest and game manager with the US Forest Service higher plants and animals native to Wisconsin, it is doubtful
during the first three decades of the twentieth century, whether more than 5 percent can be sold, fed, eaten, or
Leopold did not arrive at the idea of treating “land” with otherwise put to economic use. Yet these creatures are
ethical consideration by training, but by experience. After members of the biotic community, and if (as I believe) its
years of managing forest ecosystems as separate stability depends on its integrity, they are entitled to continu-
compartments of “resources,” he came to believe that such ance” (Leopold 1966: 246–247). Leopold stressed three
systems were more than an independent collection of points that environmental ethicists would formalize later as
commodities. Rather, they represented a relationship of inter- ethical principles. First, the value of biodiversity should be
dependent processes. Although Leopold’s views were based on a species’ contribution to ecological stability. Sec-
informed by analysis of science and experience of conserva- ond, such value implied a “right to life” for creatures which
tion management, he realized that an ethical transformation contribute to that stability. Third, the loss of a species could
also had to take place in both natural resource managers and lead to dysfunction of the community, and was therefore
the public they served. Such a transformation must recognize ethically wrong.
that “the land” had intrinsic value and develop an ethic that These premises formed the foundations of an ethic today
upheld the land’s health and ecological integrity. known as ecocentrism, which asserts that the fundamental
“It is inconceivable to me,” wrote Leopold, “that an ethi- entity to which both values and rights apply is the biotic
cal relation to land can exist without love, respect, and community, not individual specimens or species. Leopold
admiration for land, and a high regard for its value. By argued that the value of a species does not reside in itself,
value, I of course mean something far broader than mere but in its value to the integrity of the community of which it is
economic value; I mean value in the philosophical sense” a part. Ethicist J. Baird Callicott rephrased Leopold’s original
(Leopold 1966:261). Leopold’s writings on the ethical moral maxim of the land ethic in more modern, ecocentric
aspects of land management were published after his death terms as, “A thing is right when it tends to disturb the biotic
as a collection of essays entitled A Sand County Almanac. community only at normal spatial and temporal scales. It is
Here Leopold articulated a “land ethic” that endeavored to wrong when it tends otherwise” (Callicott 2006:131). Many
make ethical treatment of land a central issue in conservation. conservation biologists today see themselves as heirs of
Leopold’s coupling of values to applied science began to Leopold’s legacy to restore ethics and value to the science
change fundamental assumptions not only about the best of conservation. As Fiedler et al. (1997:84) put it, “Today the
use of natural resources, but also about the purpose of eco- emergence of conservation biology, perceived as a distinct
logical studies. These changes opened the door for develop- discipline, is a direct result of the failure of resource manage-
ment of a value-driven approach to conservation, without ment fields. . .to fully embrace the values espoused by
which the field of conservation biology could not have Leopold. . .”.
emerged. Leopold viewed his conclusions as ethically self-evident
In his argument for a land ethic, Leopold began writing, and scientifically based. “The land ethic,” he wrote, “simply
many years before Christopher Stone, about the rights of enlarges the boundaries of the community to include soils,
natural objects, asserting that it was the land’s intrinsic waters, plants, and animals, or collectively: the land”
value that led to its “rights.” Speaking against those who (Leopold 1949:204). But it is not as simple as his statement
saw the land only as a repository of “natural resources,” suggests, and, in examining its details and complexities,
Leopold wrote, “A land ethic, of course, cannot prevent the Leopold’s Land Ethic is not without critics. If species derive
alteration, management, and use of these ‘resources,’ but it value from their usefulness to community function, that is an
does affirm their right to continued existence, and, at least in expression of the ecological usefulness, not of intrinsic value.
As conservation biologist David Ehrenfeld noted, conservation values, from religious traditions. Such traditions
“. . .Leopold provides no real justification for preserving are important in creating and refining a person’s individual
those animals, plants, and habitats that are almost certainly conservation ethic and, with that, society’s ethic. As the
not essential to the ‘healthy functioning’ of any large ecosys- noted animal ecologist Charles Elton put it, “The first [reason
tem” (Ehrenfeld 1976:651). Taking this accusation a step for conservation], which is not usually put first, is religious.
further, Leopold’s detractors have accused him of There are some millions of people in the world who think that
“ecofascism,” making every species subservient to the col- animals have a right to exist and be left alone, or at any rate
lective function of the community (Callicott 1999a). that they should not be persecuted or made extinct as spe-
Leopold’s concept of “community” invokes a now cies.” (Elton 1958:143).
discredited view of interdependent populations, rather than We would trivialize conservation ethics by ignoring
of loose associations of independent populations that occur Leopold’s advice, Elton’s perceptions, and the majority of
together because of similar environmental tolerances human experience if we avoided ultimate issues of value
(Callicott 1999b). Leopold asserted that community stability addressed by religion. Religious traditions engage questions
was derived from community diversity, but many studies do inherent in intrinsic value in profound ways and answer some
not support this view. Reflecting on such findings, David of the most problematic dilemmas of conservation valuation:
Ehrenfeld remarked, “The most diverse communities were does intrinsic value exist, what is its source, and how ought
those that had occupied the most stable environments for the we to respond?
longest period of time: they were dependent on stability – not
the reverse” (Ehrenfeld 1976:652).
Many argue that intrinsic value cannot exist apart from a 10.5 Conservation Value and Practice
higher moral authority. Leopold himself acknowledged that a in Religious Traditions
conservation ethic apart from such understanding was incom-
plete. “No important change in ethics,” he wrote, “was ever 10.5.1 Intrinsic Value in the Judeo-Christian
accomplished without an internal change in our intellectual Tradition
emphasis, loyalties, affections, and convictions. The proof
that conservation has not yet touched these foundations of Most of the early writers, activists and government leaders in
conduct lies in the fact that philosophy and religion have not the US conservation movement, including John Muir,
yet heard of it. In our attempt to make conservation easy, we Gifford Pinchot, and Stephen Mather were individuals
have made it trivial” (Leopold 1966:246). inspired by a biblically-based, Christian worldview as well
The majority of the world’s seven billion people are as profound religious experiences, and often expressed their
“touched by these foundations of conduct,” drawing their ideas in biblical language and imagery (Stoll 2015)
understanding and applications of values, including (Fig. 10.10). They were not re-inventing Judeo-Christian
Fig. 10.10 Gifford Pinchot (left), Stephen Mather, (center) and John responsible for its care. (Photos courtesy of Yellowstone National
Muir (right), three prominent late nineteenth and early twentieth century Park, the US Library of Congress, and the US National Park Service,
conservationists inspired in their conservation efforts by a Christian respectively)
world view of the Earth as God’s good creation and humankind as
10.5 Conservation Value and Practice in Religious Traditions 429
teaching, but following its precedents and practices which 10.5.3 Biblical Teaching and Application
had been established in Christian faith for many centuries. in Conservation
However, the current view of the Judeo-Christian tradition in
conservation is negatively influenced by a single publication 10.5.3.1 The Moral Value of Nature
which appeared in 1967, an essay entitled “The Historical Many ethicists would argue that the most important problem
Roots of Our Ecologic Crisis” by UCLA historian Lynn to be solved in conservation ethics is to provide a basis for the
White Jr. For many in conservation, the narrative of and intrinsic value of nature. The Hebrew scriptures address this
conversation about Judeo-Christian influence in conservation issue immediately. In the first chapter of Genesis God repeat-
starts here, and so that is where we also will begin. edly proclaims that both specific kinds of creatures and
natural objects are “good.” In this context, the goodness
acknowledged and perceived by God is not utilitarian but
10.5.2 Beginning in the Middle – The Historical intrinsic. He has no self-interests that could be used to deter-
Roots of Our Ecologic Crisis mine the value of the creatures in relation to his own interests,
nor does he have need of anything from creation (Callicott
Originally given as an address at the 1966 meeting of the 1986). The repeated statements of creation’s goodness are
American Association for the Advancement of Science made by God, not by humans or in relation to humans. Such
(AAAS) and published the following year (1967) in the understanding forms the foundation of the contemporary
AAAS’s most prestigious journal, Science, “The Historical ethical framework sometimes referred to as the Judeo-
Roots of Our Ecologic Crisis,” by historian Lynn White Christian Stewardship Environmental Ethic, and its
Jr. identified the Judeo-Christian tradition as the cause of implications for conservation are profound. Based on these
the environmental crisis, asserting that it taught that nature texts, as ethicist J. Baird Callicott asserted: “The Judeo-
had no reason to exist except to serve humans. Christianity, Christian Stewardship Environmental Ethic is especially ele-
said White, established a dualism of humanity and nature, gant and powerful. It also exquisitely matches the
teaching that “it is God’s will that man exploit nature for his requirements of conservation biology. . .and confers objec-
proper ends” (White 1967:1205). Out of this perspective, tive intrinsic value on nature in the clearest and most unam-
White asserted, arose the view that everything in the world biguous of ways – by divine decree” (Callicott 2006:120).
existed for the use of human beings, and without such use
there was no value in natural objects. Use came through 10.5.3.2 The Moral Imperative – Human Response
exploitation, and exploitation increased through technology. to Nature
“Both our present science and our present technology,” After addressing the issue of nature’s intrinsic value in Gen-
concluded White, “are so tinctured with orthodox Christian esis 1, the prescription of active care by humans for creation
arrogance toward nature that no solution for our ecologic is explicated in the next (second) chapter as a moral response
crisis can be expected from them alone” (White 1967:1207). to the goodness of creation. “The Lord took the man and put
White’s views were repeated in academic and popular him in the Garden of Eden to till it and to keep it” (Genesis
circles throughout the 1970s and became a staple in 2:15). The verbs rendered “cultivate” and “keep” are more
discussions of ecological ethics in textbooks on ecology literally translated “serve” (abad) and “protect” (shamar), the
published during that decade (Krebs 1972; Colinvaux 1973; latter usually signifying loving care (“The Lord bless you and
Hinckley 1976). Influenced by White’s essay, the Judeo- keep (shamar) you,”) (Numbers 6:24). Both verbs in Scrip-
Christian tradition was vilified in all things environmental, ture usually describing service to God, especially as vocation
from discussions of landscape architecture (McHarg 1969) to (Walton 2001:185).
pollution and species extinctions (Ehrlich 1971). White’s Although White imputed the injunction to “subdue the
thesis did not survive intellectual scrutiny among historical earth and rule over the creatures” in Genesis 1 as a license
scholars (O’Connor 1988; McKay and Jentoft 1998), but his for oppressive behavior by humans toward creation, most
ideas remained popular in environmental circles long after biblical scholars understand these words in light of specific
being discredited in academic ones. Jewish and Christian acts given to humans in Genesis 2:15. Such a view is more
scholars themselves also responded with rebuttals. From consistent with the biblical ideal of ruling as an act of service
these writings emerged the modern Judeo-Christian Stew- (Deuteronomy 17:14–20, Matthew 20:25–28, John 13:
ardship Environmental Ethic (Callicott 2006), now one of 3–15). The Judeo-Christian tradition asserts that humans,
the dominant worldviews in environmental ethics. made in the image of God, bear unique responsibilities in
the care of creation that they are empowered and authorized In this view of reconciliation, conservation effort is under-
to discharge. stood as consistent with God’s purpose of redemption and
reconciliation of all things to himself. Today many conserva-
10.5.3.3 The Moral Designation –Assignment tion biologists identify themselves as Christians, and cite the
of Legal Protection to Land Bible’s message of creation’s redemption as warrant for their
Rightly did Holmes Rolston III call Judaism a “landed faith,” hope that present efforts in conservation are significant to
(Rolston 2012:15), one in which God’s promises to his peo- such future redemption. In the pages of Conservation Biol-
ple are manifested in the gift of a “good land” (Deuteronomy ogy, 30 conservation biologists expressed this conviction.
8:7–9). The land of Canaan (modern day Israel, Jordan, and . . .Christians are committed by their biblical beliefs not only to
Palestine) is God’s gift to the people of Israel (in biblical the conviction that God himself cares for his universe in a daily
context) to be used with thankfulness, respect, and reverence. and ongoing way but also that he helps and guides people in
The Jews therefore viewed the land as a “promised” land, their conservation efforts. . .every time we celebrate a conserva-
tion success story such as the recovery of the white rhinoceros in
destined for abundant life (Rolston 2016). This understand- southern Africa, we are strengthened in this present hope that
ing of the land as a gift from God is further developed in the God is working with us to redeem his creation (Stuart et al.
books of Exodus and Leviticus, Hebrew scriptures that expli- 2005:1690–1691).
cate God’s law to his people, God institutes a rest for the land,
a “land Sabbath.” “When you enter the land I am going to
10.5.3.5 Historical Expression
give you, the land must observe a Sabbath to the Lord”
Biblical teachings summarized here have been expressed in
(Leviticus 25:2). Every seventh year the land shall not be
various ways over many centuries. In the monastic tradition
cultivated, but shall receive “a Sabbath rest.” The Bible treats
of medieval Europe, monks of the Order of Saint Benedict
land as a moral subject and legal rights are imputed to it, with
were instructed to make their living from the land as farmers.
penalties prescribed for its abuse (Leviticus 26:27–35).
Benedict instructed the monks not only to prevent misdeeds
Violation of the land Sabbath is named in the Hebrew
toward nature, but to participate in work fostering harmoni-
scriptures as one of the reasons foreign adversaries deported
ous relationship with nature through sustainable agriculture
the Jews from their homeland. Speaking of Nebuchadnezzar,
and natural landscaping (Dubos 1972). One monk, viewing
the Babylonian king who conquered Israel, the author of II
the land around the monastery at Clairvaux, France, spoke of
Chronicles states, “He carried into exile to Babylon the
how the nearby river nourished the fish and watered crops
remnant, who escaped from the sword. . .Then the land
and trees. Perhaps the first author to describe ecosystem
enjoyed its Sabbath rests, all the time of its desolation it
“services,” the monk endeavored to list all services provided
rested, until the seventy years were completed in fulfillment
by the stream – which he described as “friendly,” “faithful,”
of the word of the Lord spoken by Jeremiah” (II Chronicles
and “kindly” – so as to give “thanks due to it.”
36: 20–21). Recall Christopher Stone’s criteria for
Leaders of the Protestant Reformation wrote about
establishing legal “rights” for a natural object or entity –an
principles of conservation in specific terms and applications.
entity has rights if action can be taken at its behest, if the
John Calvin explained the text of Genesis 2:15 this way.
judgment accounts for injury done to it, and if relief granted
The custody of the garden was given in charge to Adam
runs to it benefit. The parallels to Stone’s criteria here are . . .on the condition that, being content with a frugal and
striking. When the land Sabbath was not observed, God acted
moderate use of them, we should take care of what shall
on the land’s behalf to restore its rest. Taking its injury into
remain. Let him who possesses a field, so partake of its yearly
account, the nation is deported, creating relief that “runs to fruits, that he may not suffer the ground to be injured by his
the benefit” of the land.
negligence: but let him endeavor to hand it down to posterity
as he received it, or even better cultivated. (Calvin 1645:46).
10.5.3.4 The Redemptive Hope – Non-human Such teaching formed the basis of eighteenth century conser-
Creation in God’s Redemptive Plan vation efforts in the United States, as Calvinist Christians
Hebrew scriptures see ecological abuse as rebellion against
(Puritans, then Congregationalists) saw conservation as a
God (for example Hosea 4:1–3) and prescribe, not better moral duty of improvement and stewardship of nature. As
conservation practices, but reconciliation between people
historian Mark Stoll notes, the Puritans succeeded “in
and their creator (Hosea 2:18–23, Isaiah 11:6–9). New Tes-
establishing an extraordinary and enduring intellectual
tament writers assert that the promise of reconciliation framework for the work of conservation” (Stoll
described in Hebrew writings are fulfilled in the incarnation,
2015:60–61) which formed the worldview of most US con-
as God becomes part of his creation in Jesus Christ, and
servation leaders through the mid-twentieth century (Stoll
through Christ’s death reconciles all created things to himself 2015).
(Romans 8:18–22, Colossians 1:15–20).
10.5.4 Islamic Teaching on Conservation and dependence that Allah imputed to natural systems. The
concept Judeo-Christian thought would describe as “steward-
Islam arose in the seventh century A.D. in the Middle East, ship” is referred to in Islam as “viceregency” (Zaidi 1991).
although, like Judaism, it traces its historical ethnic origins As the Qur’an’ states, “Behold, the Lord said to the angels: ‘I
through Abraham. Its adherents, Muslims, believe that Allah will create a viceregent on earth. . .” (Qur’an’ 2:30).
(God) communicated to humanity through his representative, The third pillar, akhirah, arises from the Islamic view of
the prophet Mohammed, whose teachings form the principle the physical world as a testing ground of human character.
distinctions of Islam. Like Judaism and Christianity, Islam Faithful stewardship is one criterion by which God
offers a monotheistic perspective with a theocentric basis, determines the faithfulness of humankind to him, and from
perceiving the natural world as a creation of God that reveals this, decides a person’s eternal destiny. Because there is no
his glory and attributes. “The seven heavens and the earth and division of sacred and secular, eternal destiny is enforced in
all therein declare His glory: there is not a thing but celebrates present circumstances. Under Islamic law, if one ceases to
His praise. . .” (Qur’an’ 17:44). Also like Judaism and Chris- manage land responsibly, one can lose ownership of the land.
tianity, Islam maintains that God is transcendent. Although Natural resources are considered common property, not
God values his creation, he is not the same as his creation and merely of humans but of all living creatures (Masri 1992).
is not to be equated with it. God is “totally other,” but fully Expression of these principles, in the context of conservation
encompasses his creation and lovingly cares for it. Here Islam can be studied in detail in the comprehensive summary and
shares with Judaism and Christianity the concept of imma- arrangement of Islamic teachings on stewardship, The
nence-- that God operates within the physical world, always Islamic Principles for the Conservation of the Natural Envi-
and everywhere intimately present with it, yet distinct from ronment, the collaborative effort of contemporary Saudi
it. Because God created the world and works within it, the scholars (Callicott 2006).
world is not profane, but holy. It is a place to worship God in
any circumstance. Therefore “the whole earth is a mosque”
(Manzoor 1984). 10.5.5 Conservation in Hinduism
The intellectual pillars of Islam are tawhid (unity), khilafa
(trusteeship), and akhirah (accountability, or, more literally, Hindus believe that there is one Supreme Being who created
the hereafter) and form the foundations of its conservation the universe but not a “God” in an Islamic, Jewish, or Chris-
ethic (Hope and Young 1994). From the concept of tawhid, tian sense of a transcendent, Supreme Being, existing inde-
Islam perceives religion and science, value and fact, as a pendently of other beings that are his own creations. Rather,
unity (Hope and Young 1994; Wersal 1995), making Islam all things that appear as individual entities are reflections and
explicit in its opposition to separate the world into divisions manifestations of the one essential being or Brahman. One of
of secular and sacred. From this perspective, Muslims hold a the sacred writings of Hinduism, the Bhagavadgītā, states
sacramental view of the physical universe with no distinction that the Supreme Being “resides in everywhere” (Chap. 13,
between religious and secular, nor any concept akin to “sep- verse 13). In this view the Hindu perception of nature is best
aration of church and state”. Thus, all conservation law must understood as prakrti, the matrix of the material creation.
be grounded in Islamic law. One purpose of Islamic law and Prakrti is seen as the expression of the supreme intelligence
religious teaching is to provide independent judgment, cor- and physical form of Brahman. Hindu scholars and teachers
rection, and regulation of scientific activity. From this view- often refer to Brahman metaphorically as a tree, with its roots
point, many Islamic scholars and scientists are critical of “above” (in the spiritual dimension) and its branches
western science because they see its separation from religious “below,” in the physical world. The branches of Brahman
tradition as the cause of its abuses. As Islamic scientist are conceived as five fundamental elements of prakrti; sky
Seyyed Hossein Nasr put it, “Western science has become (space), air, fire, water and earth. Before every Hindu wor-
illegitimate because scientists and the rest of society fail to ship service, these elements are purified within worshippers
see the need for a higher knowledge into which it could be and the external environment in which the service takes
integrated. The spiritual value of nature is destroyed. We place. Flowers are offered as purification for sky, incense
can’t save the natural world except by rediscovering the for air, light for fire, water for water, and fragrance for the
sacred in nature” (quoted in Hope and Young 1994). earth. Thus it is natural for Hindus to view nature as some-
Islam sees nature as teleological, harmonious, and depen- thing internal rather than external, neither alien nor hostile,
dent. Because such traits represent its original state and God’s but inseparable from human identity and existence (Rao
continued intention for it, Islamic belief supports an attitude 2000). In Hinduism, planet Earth itself is personified as the
of moral obligation in regard to human interaction with female deity Bhuma devi, and given praise for her glories and
nature. The human role is one of khalifa (trustee). The virtues in various sacred Hindu texts (Oxford Centre for
human duty is to maintain the appropriate purpose, harmony, Hindu Studies 2018).
To the Hindu, every act, willfully performed, leaves a 10.5.6 Conservation Teachings in Buddhism
consequence in its wake because human life and action are
inseparable from their environment. This is the concept of Buddhism is an agnostic religion, recognizing no deity. It
karma, from the Hindi root kr, “to do.” The concept of Karma focuses on mastery of self and integration of the self with
supports the belief that every human action creates its own one’s surroundings through direct knowledge, discriminating
chain of reactions and events that will always be with that awareness, and deep compassion (Kaza 1990). Some scholars
person, as well as inescapable consequences that must be have called Buddhism the world’s most eco-centric religion
faced (Dwivedi 2000). As stated in another sacred writing (Sponsel and Natadecha-Sponsel 1993) because, for the Bud-
of Hinduism, the Mahābhārata, “an action, which has been dhist, “an environmental ethic becomes a practice in
committed by a human being in this life, follows him again recognizing and supporting relationships with all beings”
and again (whether he wishes it or not)” (quoted in Dwivedi (Kaza 1990:24). Because Buddhism stresses the importance
2000:15). Attaining an ideal life depends on choosing right of self-mastery, it speaks directly to one of the problems
actions and living within an ethos, or set of duties (dharma) perceived by Deep Ecologists in motivating human beings
that produce good consequences (good karma), supported by to change their behavior toward nature. As Deep Ecologist
a purity and balance of the five basic elements within and George Sessions puts it, “[People] have mostly opinion about
around a person. The goal is a life harmonious with nature by causal sequences in Nature in that their perceptions and
creating an environment free of pollution, because polluted thoughts are colored by their ego desires. They are essentially
elements make the human body subject to disease and distor- helpless and passive, moved by emotions, fears, and desires
tion, no longer an expression of Brahman (Rao 2000). And based on ignorance and imagination, and living life largely by
because of Hinduism’s doctrine of birth and rebirth (reincar- reacting to external causes and situations,” (Sessions
nation), Hinduism requires not simply respect for other 1985:239). Buddhism addresses this problem directly, seeing
creatures, but reverence for them, for, in Hindu writings, one of its central goals to be that of sublimating and eventually
even the Supreme Being takes on various incarnations as a eliminating the problem of desire and, removing that, reliev-
fish, a tortoise, and a boar, among others (Dwivedi 2000). ing the suffering (dukka) that all life experiences in the world.
The duties warranted by these beliefs are stated in Hindu The fundamental axiom in Buddhism is the Law of
writings, such as Prthivī Sūkta, a hymn devoted to praise of Dependent Origination, also called the Law of Dependent
Mother Earth, as explicit duties. In this hymn, the worshipper Co-Arising, a formalization of the concept that all events and
makes this request, beings are interdependent and interrelated (Kalupahana
O, our Mother Earth! Sacred are thy hills, snowy mountains, and 1987). In the words of its own sacred writings,
deep forests. Be kind to us and bestow upon us happiness. May For one who truly sees the pure and simple arising of phenomena
you be fertile, arable, and nourisher of all (quoted in Dwivedi and the pure and simple continuity of conditioned things, there is
2000:10). no fear. When with wisdom one sees the world as just like grass
and wood, not finding any selfishness, one does not grieve with
Hinduism has had a long and significant influence on the idea, ‘this is not mine.’ (quoted in Batchelor 1992:10).
ecology and conservation. Two of the earliest and most
influential writers in US conservation, Ralph Waldo Emerson On the basis of this law, Buddhism stresses a unity of self
and Henry David Thoreau (Chap. 1), were influenced by and environment. The Buddhist ideal of nirvana, the awak-
Hinduism in their perception and understanding of nature. ening into a state of bliss, is reached when the boundary
Hinduism’s teaching that all beings are an expression of the separating the self from its surroundings and all mortal
one essential being leads to a sense of identity between cravings is extinguished (Smith 1958), a concept embraced
humans and other living things. Non-human species and by adherents of Deep Ecology. Buddhism does not empha-
non-living objects are seen as manifestations of one’s own size that resources are limited, but that people should limit
life, and are therefore to be protected and preserved. their use of resources, and will do so as they learn to subli-
Although Deep Ecology intentionally and self-consciously mate desire.
draws from many religious traditions, it is perhaps most In Buddhism, the foundation of knowledge is personal
indebted to Hinduism because of Hinduism’s strong identifi- experience. Experiential knowledge is especially emphasized
cation of humans with other species and natural objects in spiritual matters, so personal meditation is accorded high
(Naess 1989). More than any other religious tradition, Hin- value, as are natural environments in which to meditate,
duism explicitly supports identification with non-human life which are valued as “sacred space” for meditation
as a genuine perception of reality and a basis for the care of (Brockelman 1987; Buri 1989). Buddhism’s goal is increas-
non-human creatures. ing self-knowledge, with increasing self-mastery and self-
restraint. Its “Middle Way” of correct moral behavior

emphasizes detachment from material things and present
concerns. Buddhism takes a high view of personal responsi-
bility because it shares, with Hinduism, the doctrine of
karma. Future happiness results from appropriate present
conduct. Wrong actions of the past will produce bad effects
in the present, so Buddhism encourages environmental edu-
cation and appropriate environmental behavior, not simply
for present consequences, but for future ones.
Buddhism can present problems in conservation ethics
because, at some levels, its teachings seem at odds with
conservation objectives. Buddhists are concerned to relieve
suffering (dukkha) in life, and in such teaching find affinity
for principles of Animal Rights advocates like Peter Singer
who equate moral consideration of a creature with its capac-
ity to suffer. But “species,” which are a conceptual abstrac-
tion, cannot suffer, and would not be proper objects of moral
concern. Further, Buddhists, in being concerned to relieve all
suffering, would not count the suffering of an endangered
species as more significant than the suffering of a common
one. British philosopher of religion Simon James explains the
implications. “What then of an endangered species, such as
the Indochinese tiger? Well, individual tigers can suffer, and
so they are morally considerable, which is to say that Fig. 10.11 The Asian open-billed stork (Anastomus oscitans), a spe-
according to Buddhist ethics one ought to exercise compas- cies that would be extinct in the nation of Thailand if not for the fact that
sion, loving-kindness, and other moral virtues in one’s its last remaining breeding ground lies within the sanctuary of Wat Phai
dealings with them. But what of the species Panthera tigris Lom, a Buddhist temple and its surrounding grounds near the city of
Bangkok. (Photo courtesy of Charles J. Sharp, under CC-BY-SA-4.0)
corbetti? The species is an abstract entity. It cannot suffer any
more than it can chase deer or slink unnoticed through the
grass. To think of it as a sentient is, one might say, to make a the species, but they do care about the storks (James
category mistake” (James 2006:89). Further, explains, James, 2006:91).
“[Buddhist teaching is opposed to] the kind of moral discrim- Conservation biologist Barbara Paterson asserts that Bud-
ination that would justify ascribing a panda or condor a dhist teachings may be the key to solving one of the most
higher moral value than a rat, say, or a starling. . .By the fundamental problems in conservation. “The modern conser-
lights of Buddhist ethics, dukkha is dukkha, regardless of vation paradigm,” notes Paterson, “conservation for and with
whether it is the suffering of a member of an endangered people, requires that we overcome the dualism of human
species or a member or a more common one. . .” (James versus nature, which creates antagonism between conserva-
2006:89). tionists and other people. [Buddhist teaching] provides a
Buddhist teaching can also find itself at odds with conser- basis for a conservation philosophy that sees the conserva-
vation effort because it would oppose the idea that, to con- tionist not as a defender of the natural world against the
serve one species, one might have to harm another species to harmful impact of human actions but as one who realizes
reduce competition, predation, or parasitism. Yet, when Bud- the interdependencies both between people and between peo-
dhist teachings are faithfully practiced, the outcomes are ple and nature, and who strives to awaken such awareness in
often positive for conservation. Because of the emphasis on others in order to achieve a better future for all” (Paterson
contemplation and meditation, Buddhists have a long history 2006:149).
of conserving natural areas for these expressed purposes.
Sometimes the effect of such conservation of space is the
conservation of species. For example, the Asian open-billed 10.5.7 Indigenous Belief Systems
stork (Anastomus oscitans) (Fig. 10.11) would be extinct in in Conservation
the nation of Thailand if not for the fact that its last remaining
breeding ground lies within the sanctuary of Wat Phai Lom, a The religions, worldviews, and cosmologies of indigenous
Buddhist temple and its surrounding grounds near the city of peoples throughout the world are varied and unique, but all
Bangkok. In this case, perhaps the monks do not care about have growing effect on conservation policies and decisions at
local, national, and even international levels. Scholars who Place in indigenous religions is a concept that does not
have made comparative studies of the ecological effects of simply signify spatial location or dimension, but relation of
indigenous cultures compared to so called “western the individual to the world through right interaction with a
cultures” offer some explanation of why this is so. Many particular place. Regard for place matters in conservation. In
indigenous cultures have maintained lands and resources the west African nation of Ghana, for example, “sacred
better than western cultures, not only because of their lower groves,” or, in one case, a sacred lagoon, are protected by
population densities, but also because of cultural values, some tribal groups because the site itself is considered a god
beliefs, and practices of restraint that limit use of resources or the habitation of a god, or the place contains sacred species
(Torri and Herrmann 2011). Highlighting spiritual and eco- viewed as gods or as species favored by the god of the grove
logical connectedness within Haudenosaunee (an alliance of or lagoon (Ntiamoa-Baidu 2008). Such protections or
six Native American nations called the Iroquis Confederacy) “taboos” observed by indigenous people follow strategies
culture, Joe Sheridan and Roronhiakenwen “He Clears the of protecting the place, species within the place, or restric-
Sky” Longboat argue, “diminishing biodiversity augers tively regulating use and removal of natural resources from
against the continued capacity to know how to think with the place (Ntiamoa-Baidu 2008). Although such taboos have
everything. The principle that every being participates in no standing in federal law, they have consequential effects.
everything immunizes against anthropocentrism and through Four species of sea turtles, black heron (Egretta ardesiaca), a
a satisfaction with Creation, offers a conscious humility that mollusc (Tympanotonus fuscatus), black and white colobus
keeps humans in their place of thankfulness, respect, and (Colobus vellerosus) and the mona monkey (Cercopithecus
appreciation. This ensures Creation’s continuation. Humans mona) have higher densities in such protected sites that in
think at their best if they know they are the last beings non-protected (and non-sacred) sites (Ntiamoa-Baidu 2008)
created. Literally, after all, humans are totally dependent (Fig. 10.12).
on everything else.” (Sheridan and Longboat 2006). Ecological knowledge (or wisdom) is viewed as a source
Indigenous belief systems are not only diverse, but today of spiritual insight with potential for transformative power of
often hybridized with global religions like Christianity, both humans and nature, not simply a dispassionate under-
Islam, Hinduism, and Buddhism. Yet they retain common standing of facts. Some scholars have gone so far as to
characteristics amidst such change. These include (1) a strong describe the role of ecological knowledge in indigenous
sense of place integral to belief and practice of the religion; societies as knowledge-practice-belief complex in individual
(2) ecological knowledge as a form of religious wisdom and and community life in such societies (Torri and Herrmann
transformative power; (3) unity with and ethical regard for 2011) in which ecological knowledge is not simply a means
the natural world and its creatures; and (4) felt and of understanding nature but a form of oversight, control, and
intentionally cultivated, repeated, and ritualized experiences re-direction of natural processes. The Temiar people of
of interacting with nature. Malaya, for example, define their understanding of their
Fig. 10.12 The black heron (Egretta ardesiaca) (left), black and white non-protected (and non-sacred) sites. (Heron photo courtesy of
colobus (Colobus vellerosus) (center) and mona monkey D. Keats, under CC-BY-SA-2.0, colobus photo courtesy of Tuxyso,
(Cercopithecus mona) (right), all species with higher densities in sites under CC-BY-SA-3.0, and mona monkey photo courtesy of L. Coret)
protected sites as sacred in indigenous religions in Ghana that in
“home landscape” not only in terms of spatial location and (Fig. 10.14). Working with the Michigan Department of
features, but also through the presence of particular species of Natural Resources, the Tribe has funded and, with their
plants native to the landscape and the spirits of the plants that staff and volunteers, supported restoration of this species in
interact with the Temiar and medicines derived from the Michigan’s lower Manistee River (Fig. 10.15).
plants that heal or transform human lives (Grim 2018). This Felt and intentionally cultivated, repeated, and ritualized
role of ecological knowledge is also particularly evident in experiences of interacting with nature are also characteristics
indigenous cultures that depend on hunting as a means of of indigenous religions. Such ritual experiences are often, not
subsistence. In such cultures, it is common for the hunter to surprisingly, related to the aforementioned characteristic of
ask permission of his quarry to take its life, or make an the importance of place, especially specific sacred sites. For
apology for doing so. Success in hunting is not viewed as a example, the Lakota people of the US Great Plains treat Bear
cause and effect relationship among events associated with a Butte, in the Black Hills of South Dakota, as a sacred site to
single hunt, but the outcome of a pattern of long-term rela- be used for prayer and meditation. The contemporary Lakota
tionship with the natural world and the creatures in it. Such writer Charlotte Black Elk said of Bear Butte that it was “a
beliefs can also lead to the view that some species, identified place where no war party would attack someone who was
or associated with spirits, gods, or goddesses, are active there to pray. It was respected by all the Tribes. Other people
protectors of the environment. In India’s Sariska Tiger came down – people who were our enemies came down to
Reserve, indigenous people living in the Reserve believe
that the decline of the tiger and other predators has been
one of the causes of regional deforestation because the tiger
is seen as the protector of the forest and symbolizes protec-
tion of the forest by the Hindu warrior goddess Durga, often
depicted riding a tiger (Fig. 10.13).
Unity with and regard for the natural world and its
creatures is expressed in many forms, often in a belief in
direct kinship with animals or natural forces as ancestors. In
the US state of Michigan, the Little River Band of Ottawa
Indians is organized by clans, each identifying a different
animal species or natural force as its ancestor (for example,
Fig. 10.14 The lake sturgeon (Acipenser fulvescens), a species once
Thunder Clan, Bear Clan). The Sturgeon Clan has been an common in the US Great Lakes and their tributary streams but reduced
important contributor in efforts to restore the lake sturgeon in abundance by overharvesting and habitat degradation. (Photo cour-
(Acipenser fulvescens) in areas where it has been extirpated tesy of M. Holtgren)
Fig. 10.13 The Indochinese tiger (Panthera tigris corbetti) (left), (right), often depicted riding a tiger. (Photo of tiger by Lotse,
principal species of conservation concern in India’s Sariska Tiger under CC3-BY-SA-3.0. The image of the Hindu female deity is a
Reserve. Indigenous people living in the Reserve view the tiger as the photographic reproduction of a work of public domain art)
protector of the forest, personified as the Hindu warrior goddes Durga,
Fig. 10.15 Staff and volunteers of the Little River Band of Ottawa The Sturgeon Clan of the LRBOI identify the lake sturgeon as the
Indians (LRBOI), an indigenous tribal nation in the US state of ancestor of their clan, creating strong affection for the species and
Michigan, work to restore populations of the lake sturgeon (Acipenser commitment to its restoration in the Manistee River, the principal river
fulvescens) in the Manistee River, a major tributary of Lake Michigan, within their tribal land in the US state of Michigan. (Photo courtesy of
where the lake sturgeon had been extirpated through overharvesting. M. Holtgren)
pray. We did not attack them” (Carillo 1998:105). Bear Butte a “goal-rational” perspective in conservation that has histori-
remains a place where Lakotans come to fast, pray, undergo cally supported traditional environmental agencies and
purification rituals and perform ceremonial dances with NGOs, a view in which measurable ends, achieved by tech-
others, all of which are interactive with the place itself and nical means, are the index of success (Abuyuan 2006). How-
its natural environment. ever, such an approach, even if successful, may do nothing to
actually enhance the public’s dedication to conservation or its
respect for nature because it does not address public motiva-
10.6 Practical Applications: Faith-Based tion or private practice. As environmentalist David Orr noted,
Contributions to Conservation conservation biologists “lack both a deep explanation of what
ails us and a larger cosmology that resonates with the public”
10.6.1 “Goal Rational” Versus “Value Rational” (Orr 2005:290). Goal-rational approaches emphasize techni-
Conservation cal competency but fail to frame conservation as moral
endeavor. Without a moral framework, most individuals out-
In their classic paper, “The Death of Environmentalism,” side the scientific community fail to perceive value in the
authors Michael Shellenberger and Ted Nordhaus, assert conservation effort, even when the effort achieves “success.”
“What the environmental movement needs more than any- All major world religions possess teachings about creation
thing else right now is to take a collective step back to rethink and nature, with sophisticated insights into the relationship
everything. We will never be able to turn things around as between human beings and their environment. Such insights
long as we understand our failures as essentially tactical and can lead to motivation for environmental stewardship that is
make proposals that are essentially technical” (Shellenberger viewed as culturally legitimate (Abuyuan 2006). Today
and Nordhaus 2004). The problem Shellenberger and religiously-based insights about conservation are applied in
Nordhaus describe is considered by many to be the fruit of faith-based organizations (FBOs) throughout the world. An
10.6 Practical Applications: Faith-Based Contributions to Conservation 437
understanding of their growing impact in conservation is and Evangelicals for Social Action for a similar purpose. In
relevant and vital to an understanding of contemporary con- addition to these and many other organizations, individual
servation efforts. denominations have engaged in a variety of activities to
promote environmental policies based on a biblical under-
standing of the stewardship of creation (e.g., Guenthner
10.6.2 Jewish and Christian FBOs 1995).
The Christian FBO most actively engaged in global con-
In September of 1989, the Jewish Theological Seminary of servation policy, management, and research is A Rocha
America devoted its High Holiday message, published as a International (ARI). Initiated in Portugal in 1983, ARI now
full-page ad in the New York Times, to the environmental coordinates activities of 20 national chapters on six
crisis (Schorsch 1992). In 1991, Jews and Catholic Christians continents. It is the only faith-based conservation organiza-
collaborated with the Institute for Theological Encounter tion with member status in IUCN, and its membership is
with Science and Technology (ITEST) to sponsor the sym- merited by significant successes in conservation, even under
posium and workshop, “Some Christian and Jewish difficult circumstances. In the politically unstable climate of
Perspectives on the Creation” which included discussion Lebanon, for example, ARI was successful in preserving one
and subsequent publication of Jewish and Christian of the last major wetland areas in the Near East land bridge
expositions of biblical teaching on creation stewardship between Europe and Asia, the Aammiq Wetland (Fig. 10.16).
(Brungs and Postiglione 1991). The subsequent formation A major stopover for hundreds of species migrating between
of the Coalition on Environment and Jewish Life (COEJL) Eurasia and Africa, ARI’s efforts combined and now con-
in 1993 has led to an even more active role for the Jewish tinue scientific study, an environmental education program
Community in conservation and environmental stewardship. for local schools, a community arts program for women, and
As part of its efforts, COEJL formed an Environmental a summer science club for local students. A Rocha Kenya has
Leadership Institute, produced a comprehensive Environ- worked effectively with local residents and government
mental Policy Platform, a program for “Greening officials in the Arabuko-Sokoke Forest, the largest remnant
Synagogues,” developed an umbrella organization, the Jew- of dry coastal forest in East Africa, to developed the
ish Global Environmental Network, and organized an exten- Arabuko-Sokoke Schools and Eco-Tourism Scheme
sive array of environmental education programs for students (ASSETS). In this effort, A Rocha Kenya developed facilities
of all ages. In 2005, COEJL helped to form and initiate the and services for tourists who come to see the forest’s note-
Jewish Global Environmental Network (JGEN) which began worthy plant and animal life, then diverts the funds generated
efforts to develop partnerships and collaborative initiatives to provide scholarships for local school children (“eco-
through which Jewish environmental leaders around the bursaries”) and to nature conservation. In the UK, the once
world could work together for a sustainable future (COEJL derelict Minet Site in Middlesex is being transformed into a
2007). More recently COEJL organized resources for county park and conservation area by A Rocha’s UK chapter
People’s Climate Shabbat (Sabbath), creating a collection of (A Rocha 2018).
writings based on Hebrew scriptures calling for action to stop
climate change and conserve natural resources, and
manifested in support of the recent People’s Climate March 10.6.3 Laudato Si – Pope Francis’s Call to Global
(29 April 2017) (COEJL 2015). Conservation
COEJL is but one of many Jewish and Christian conser-
vation efforts which have become too numerous to name. In The publication of the Encyclical Letter of the leader of the
fact, umbrella organizations and networking groups have Catholic Church, Pope Francis, (Fig. 10.17), Laudato Si0 , On
become necessary simply to list, and attempt to coordinate, the Care for Our Common Home, in 2015 brought the Chris-
the array of Jewish and Christian FBOs now active in conser- tian community to the forefront of attention in concern for
vation. Among these coordinating groups is the National environmental and biodiversity conservation. Encyclicals are
Religious Partnership for the Environment (NRPE), which the most authoritative and important publications of the
includes leadership from the National Council of Churches, Catholic Church, and media coverage of the document was
the US Catholic Conference, the Coalition on the Environ- unprecedented (Tucker and Grim 2016). Opening his letter
ment and Jewish Life, and the Evangelical Environmental with the words of Saint Francis (Laudato Si0 , mi’ Signore,
Network (EEN). The NRPE’s goal is to increase engagement “Praise be to you, my Lord”) the Pope reminded his readers
in environmental stewardship by local congregations as well that Francis continued by saying, “through our Sister, Mother
as to demonstrate and address connections between environ- Earth, who sustains and governs us, and who produces vari-
mental concerns and social justice. Similarly, the EEN was ous fruit with coloured flowers and herbs.” In the same spirit,
formed in 1993 through a cooperative effort of World Vision Pope Francis called for a repentance of human attitude
Fig. 10.16 The Aammiq Wetland in Lebanon, a major stopover area of named a Ramsar International Wetland primarily through the conserva-
hundreds of species of birds migrating between Africa and Eurasia. tion education and management efforts of the Lebanon Chapter of the
Threatened with destruction because of excessive water removals for Christian conservation organization, A Rocha. (Photo courtesy of
irrigation and overhunting, the wetland was preserved and eventually M. Skafi)
toward our Sister, Mother Earth. “We have come to see

ourselves are her lords and masters, entitled to plunder her
at will. The violence present in our hearts, wounded by sin, is
also reflected in the symptoms of sickness evident in the soil,
in the water, in the air, and in all forms of life.”
Pope Francis began with Saint Francis to show that the
ethic of care and concern for nature is deeply rooted in
historic church doctrine and practice, as well as scripture,
and drew heavily on the work of past popes, bishops,
conferences, and the Catholic Catechism itself – all of this
aimed at leading readers to a repentant attitude toward the
environment. Repentance toward the environment, said the
Pope, requires repentance toward the dignity of people. Cen-
tral to his message was the concept of “integral ecology,” an
ecology that fosters human capacity for moral development.
“To fully possess, use and enjoy these human capacities,”
said Pope Francis, “a human life must be grounded in three
fundamental relationships: with God, with our neighbor, and
with the earth itself. To enter into the third relationship,
Fig. 10.17 Pope Francis, leader of the Catholic Church, whose Encyc- human beings must realize that “each creature possesses its
lical Letter, Laudato Si0 , On the Care for Our Common Home own particular goodness and perfection. . .Each of the various
(2015), gained unprecedented media coverage and response from creatures, willed in its own being, reflects in its own way a
conservationists and brought the Christian community to the forefront
of attention in concern for environmental and biodiversity conservation.
ray of God’s infinite wisdom and goodness. Man must there-
(Photo courtesy of Korean Culture and Information Service, under fore respect the particular goodness of every creature, to
CC-BY-SA-2.0) avoid any disordered use of things (Catechism of the Catholic
Church: 339). In the end, asserted Francis, the environment

we must protect is not something external to us, something Information Box 10.2 (continued)
‘out there.’ “When we speak of the ‘environment’, what we are considered the most influential documents of the
really mean is a relationship existing between nature and the Catholic Church, and for Pope Francis to issue an
society which lives in it” (Pope Francis 2015:104). Encyclical. . .on the subject of the global environment,
The Pope’s message was not simply a religious proclama- was a very significant event indeed” (Raven 2016:255).
tion of interest to the Catholic Church, but a major event in Calling the Pope’s Encyclical a “breath of fresh air and
the scientific and conservation committee, where it created hope in times of darkness,” (Ceballos 2016:293), biol-
significant impact and thoughtful response (Information Box ogist Gerardo Ceballos noted that “In his Encyclical,
10.2). Although it remains to be seen what effects Laudato Pope Francis offers a rare glimpse of clarity from a
Si’ will have on global environmental conservation, the inter- major world leader addressing some of the most press-
action it has generated between the religious and scientific ing global environmental issues affecting Mother
community has been unprecedented. Nature – our common home, and the titanic efforts
that will be required to solve them” (Ceballos
2016:285). “The publication and outreach of Laudato
Information Box 10.2: Response of Conservation
Si’,” said Ceballos, “played a role in many important
Scientists and Ethicists to Laudato Si’
global discussions, and very likely had some influence
The Pope’s message of Laudato Si’ was quickly
on the outcome of the historic Paris Climate agreement
endorsed by the Ecological Society of America (ESA)
of December 2015. . .” Ethicists Tucker and Grim per-
with a statement combining approval by their past,
ceptively noted that, given the moral stature of both
present, and incoming presidents, calling it “an elo-
Encyclicals generally and Pope Francis specifically,
quent plea for responsible Earth Stewardship . . .is
Laudato Si’ “has enormous transformative power
clearly informed by the science underpinning today’s
within education” (Tucker and Grim 2016:263).
environmental challenges” (ESA 2015).
Realizing that Pope Francis is not the first Christian
The Quarterly Review of Biology, the world’s pre-
leader or writer to address environmental concerns,
mier English-language biological review journal,
Tucker and Grim nevertheless recognized the unique
recognized the Pope’s message with a special section
place and power of the Encyclical. Comparing Laudato
on Laudato Si’ in its 91st volume, 2016, including
Si’ to both past and contemporary works, they
invited responses by Cal DeWitt, Professor of Environ-
concluded that “Despite the richness of these
mental Studies at the University of Wisconsin-Madison
statements and the numerous books that have already
(USA) and one of the founders of the contemporary
been published, there is nothing available that
creation care movement in the Christian community;
compares to the Papal Encyclical in terms of moral
internationally known plant conservation biologist
influence. For efficacy in awakening minds and hearts
Peter Raven of the Missouri (USA) Botanical Garden
and for long-term educational awareness regarding
and member of the Pontifical Academy of Sciences that
environmental issues, this is a watershed document.”
advises the Pope on scientific matters; renowned biolo-
(Tucker and Grim 2016:263).
gist and expert on contemporary extinctions Gerado
Ceballos of the Institute of Ecology, National Autono-
mous University of Mexico; and internationally 10.6.4 Conservation FBOs in Islam
respected ethicists Mary Evelyn Tucker and John
Grim, co-directors of the Yale University Forum on In Islam, concerns in conservation are increasingly well-
Religion and Ecology. Noting the Pope’s recurring represented and put to practical application by the Islamic
theme of developing an integral ecology that reclaims Foundation for Ecology and Environmental Sciences
a moral environment in which people can make just (IFEES). Established in the mid-1980s, IFEES has grown in
choices, “Pope Francis,” said DeWitt, “is suggesting membership and influence to become the most widely
that we repurchase the wholeness of life, thereby to recognized international Muslim conservation organization,
work thoughtfully within the constraints and not only articulating the Muslim understanding of conserva-
opportunities of our 8000 mile diameter planet to tion and environmental stewardship, but working out such
make needed contributions to the integrity of human understanding in practical applications. Their efforts in
and societal life and of the life of the biosphere (DeWitt research, teaching, and training have reached the Muslim
2016:277). For his part, Raven noted that “Encyclicals community throughout the world, with recent projects
including Islam and Biodiversity, Schools4Trees and the
(continued) Green Mosques Project (www.ifees.org.uk/projects), and
IFEES has been a major organizer of the Islamic Declaration villagers a permit to cut trees to make farm implements, but
on Climate Change published in 2015. In Yemen, for exam- had granted a request in the same forest for a foreign com-
ple, IFEES initiated the rehabilitation of traditional water pany to cut trees to make sporting goods), and concerns for
conservation practices through the application of Islamic local environmental quality (the increase of soil erosion to the
conservation principles (IFEES 2007). IFEES contributes as detriment of local agriculture and water quality) (James
a consultant to such international conservation agencies as 2000). But they were also inspired by the widespread beliefs
The World Wide Fund for Nature and the Earth Charter of ordinary Hindus that trees were sacred objects. The forest,
Consultative Committee. Another increasingly effective in local Hindu folk teaching, is seen as the highest expression
Islamic FBO, the African Muslim Environmental Network of the earth’s fertility and productivity, personified by the
(AMEN) was formed in 2005. Among other efforts, AMEN goddess Vāna Durgā, the tree goddess and earth mother
has been active in coordinating the revival of traditional and (Shiva 1989). Most of the protesters were, initially, women,
sustainable fishing practices in Muslim communities along and they gave the chipko movement its most dramatic con-
the East African coast, as well as in establishing a program of frontation and most memorable slogan. In a protest in 1977, a
forest protection based on guidelines of Islamic law forest officer of the Indian government went into the forests
(ARC 2018). to convince the women that the proposed logging was scien-
tifically sound and economically indispensable. He ended by
saying, “You foolish women! Do you know what the forests
10.6.5 Conservation Activism in Hinduism bear? Resin, timber, and foreign exchange!” But, not
intimidated, the women hurled the question back with a
Perhaps the most active conservation effort in Hinduism very different answer. “What do the forests bear?” they
today is the Bhumi Project, a worldwide Hindu response to cried. “Soil, water, and pure air! Soil, water, and pure air
environmental issues facilitated by the Oxford Centre for sustain the earth and all she bears!” (James 2000).
Hindu Studies and the Alliance of Religions and Conserva- The complexities of the chipko movement must not be
tion (OCHS 2018). Working with Hindu individuals, oversimplified. Hinduism has been criticized in environmen-
communities, temples and organizations throughout the tal circles because of its core belief that the physical world
world, the Bhumi Project states its core values as being and its diverse entities are derivatives of the undifferentiated
those of respect, compassion and service toward the environ- and unmanifest Brahman, the supreme reality, thus making
ment (OCHS 2018). Representatives of the Bhumi Project the world and its biodiversity “less real.” In truth, the chipko
have participated in the People’s Climate March in movement did include an element of negating the world.
Washington, D.C. (USA) in April 2017, and, in the same However, it was not the natural world that was negated in
year, participated in a symposium on climate change, also in the chipko protests, but the world of scientific and economic
the US capital, hosted consultations on solar energy in India, reductionism that made natural objects worth no more than
and sponsored meetings on climate change in the UK (Bhumi their value as market goods. In this case, it was Hindu beliefs
Project 2018). that contributed to recognizing a greater value for the forests
Although these and other recent actions are relevant to than secular market-based valuations could discern, and this
conservation, Hinduism has a much longer history of forceful recognition saved the forests.
and practical engagement in the conservation of biodiversity.
The most famous expression of such engagement is the
chipko movement of northern India. Chipko is derived from 10.6.6 Conservation FBOs in Buddhism
a Hindi word meaning “to hug,” or “to embrace.” The move-
ment is dated from a protest near the town of Gopeshwar in An expression of the conservation values of Buddhism is
the province of Uttar Pradesh in 1973. Villagers, protesting modeled in current conservation efforts led by many of
logging policies, went into the forests and physically Buddhism’s most well-known and influential authorities.
embraced trees to be cut by loggers. The loggers, unwilling The Dalai Lama of Tibet, perhaps the world’s foremost
to harm the villagers, did not cut the trees. Buddhist leader, is also one of the world religious
What followed this initial protest was a long and complex community’s foremost conservationists. With his support,
struggle over government forest policies and development the Buddhist Perception of Nature Project (begun in 1985),
practices which continues to this day. The motivations sponsored by the World Wildlife Fund, identified and
behind the chipko movement were not inspired exclusively integrated environmentally relevant passages from Buddhist
by religious conviction. They also were motivated by politi- scriptures and secondary literature. In Thailand, the Buddhist
cal issues of self-governance (the right of the villagers rather leader Chatsumarn Kabilsingh contributed to this project by
than the government to determine the fate of local forests), framing Buddhist doctrines into “teaching stories” that have
issues of social justice (the government had denied the been distributed and used in environmental and conservation
education curricula in Buddhist cultures throughout South- among their staff, volunteers, and constituency; significant,
east Asia (Kabilsingh 1990). Many of these stories emphasize meaningful connection with local communities; and pro-
Buddhist instruction to not cause harm to others in one’s found ethical motivation for their conservation work. Today
environment and to protect natural objects such as trees, the work and influence of faith-based conservation
rivers, and animals (Kaza 1993). Also in Thailand, the Bud- organizations is so extensive and culturally pervasive that is
dhist monk Pongsak Tejadhammo established the has generated its own global umbrella organization, the Alli-
Dhammanaat Foundation in 1985 to preserve forests and ance for Religions and Conservation (ARC). Although offi-
create greater environmental harmony and security for local cially a secular, UK-based organization, ARC helps
villagers. The foundation is not merely a preserve, but a site coordinate efforts by the world’s major faiths to develop
of ecological restoration (Sponsel and Natadecha-Sponsel environmental programs consistent with their own core
1993). Throughout Thailand, Buddhist monks have adopted teachings, beliefs and practices. Similarly recognizing the
a strategy of forest protection by ritually ordaining individual influence of faith communities in the global conservation
trees and wrapping them in sacred orange robes normally effort, the United Nations Environment launched its Faith
worn only by monks. A devout Buddhist would never kill a for Earth Initiative in August 2017. Lead Principal Advisor
monk, so the symbolism is obvious in its cultural context. on Engaging with Faith-Based Organizations at UN Environ-
The Buddhist Peace Fellowship, founded in 1978 to address ment, Iyad Abumoghli, stated, “I personally believe that
peace and environmental issues, is yet another example of leadership empowerment and partnership with spiritual
systematic attempts by Buddhists to contribute constructively leaders is important. The words of faith leaders often get to
to worldwide conservation efforts. Today Buddhist environ- the hearts and minds faster than any other word, because they
mental organizations include One Earth Sangha and Green are considered to be the words of sacred scripts and represent
Sangha, as well as one of the largest global Buddhist what God has asked us to do” (UN Environment 2018). An
organizations, Tzu Chi. Although a broad-based relief and example of how effective such work can be, as well as its
development organization with varied interests and advantages over strictly secular approaches, is offered in the
ministries, Tzu Chi is actively engaged in environmental following example.
protection as one of its core missions. In addition to its direct
efforts in conservation, Tzu Chi has produced a number of
critically acclaimed environmental films including, “Keep it 10.6.8 Saving the Cedars of Lebanon
Grand,” “Stung By Climate,” and “The Disappearing Fron-
tier” (Tzu Chi 2006). Many of Tzu Chi’s efforts have been During the last 40 years, the coastline of Lebanon has expe-
directed at environmental education, such as its “Everyday is rienced rapid development. Its shorelines and hills have been
Earth Day” school programs, and disaster relief, often in converted from natural areas to roads, homes, and urban
response to environmentally-related events. business centers. Yet it was these shorelines and hillsides
that were the home of the famed cedars of Lebanon, forests
of massive and ancient trees unique in the Mediterranean
10.6.7 Future Roles and Contributions of FBOs world. Most of these forests had been destroyed by the time
in Global Conservation the international conservation community realized the gravity
of the situation. The United Nations Environmental
Conservation FBOs have proven capable of effective, pro- Programme (UNEP) and other conservation NGOs identified
ductive partnerships with secular organizations and agencies, these Mediterranean forests as a conservation priority, rank-
and their influence is growing. Today many conservation and ing them among the 200 most important world ecosystems to
development organizations have a person, or entire be protected. But were any left? Upon investigation, the
departments and programs, assigned exclusively to working answer turned out to be “yes.” A sizable ancient forest cov-
with faith-based communities and FBOs. The late Tony ering three hills, locally known as the Forest of Harissa, was
Whitten, leader of the Faith and Environment Program for- located north of Beirut (Fig. 10.18). But who owned it? The
merly a part of conservation initiatives at The World Bank, owner turned out to be a church, the Maronite Church of
said regarding FBOs in conservation, “I ask them (skeptics) Lebanon, which had held title to the land for centuries. To the
why on Earth they wouldn’t (engage FBOs)? . . .When they Maronite Church, this forest was not the Forest of Harissa,
reach people so easily, when their agendas coincide – what but the Holy Forest of Our Lady of Lebanon. In the center of
on Earth could be the reason for not doing it?” (quoted in the forest was the Cathedral of Our Lady and an enormous
Abuyuan 2006:221). Because conservation FBOs provide outdoor statue of the Virgin Mary. UNEP and other conser-
opportunity for people to transform faith commitments into vation organizations, including the World Wildlife Fund
meaningful action, faith-based organizations engaged in con- (WWF) prepared a 48-page proposal and delivered it to the
servation inspire high levels of loyalty and dedication church. The proposal demanded a promise from the church to
sufficient reason for preserving it. The Maronite Church did

not. They had preserved the forest for centuries as a sacred
trust to God, long before UNEP existed. As Martin Palmer of
the World Bank, a participant in this conservation effort,
observed in retrospect, “By insisting that people adopt their
view of the world, many campaigning groups cut themselves
off from natural allies, who may see things differently, but no
less compassionately” (quoted in Palmer and Findlay 2003:9;
summarized in Van Dyke 2010). Subsequent to making a
formal declaration to protect the forest and its biodiversity,
the Maronite Church created an ecology center for young
people, formally protected two other woodland sites, and
developed a program of environmental education and activ-
ism in 77 Lebanese villages and towns, making the church
one of the largest and most effective environmental
Fig. 10.18 Environmentalist Firas Bou Ghanem standing in front of organizations in Lebanon and one of the most influential
the ecologically fragile Jisr-el-Qadi region of the Harissa Forest. This advocates in the country for environmental protection.
forest, one of the last remaining reserves of the cedars of Lebanon, is Regardless of the foundation of value systems that moti-
preserved by the Maronite Church. (Photo by Victoria Finlay, Alliance vate people to act to conserve nature and its biodiversity,
of Religions and Conservation (ARC))
motivation must be transformed into ethical rules that govern
behavior at personal, social and governmental levels if such
abide by national and international laws to ensure protection
values are to matter in saving species. We end this chapter
of the forest. As the authors of the proposal put it, “The area’s
with an example from Indonesia that illustrates the conflicts
custodians must have protection of biodiversity as a first-
of values that lead to species endangerment, and the way such
order management objective. If other objectives take prece-
conflicts have been addressed in solving a significant conser-
dence over biodiversity protection, then the area as a whole,
vation problem: the problem of orphan orangutans.
or those parts of the area where other objectives take prece-
dence, should not be classified as a protected area” (quoted in
Palmer and Findlay 2003:8). Synthesis
Presented with the idea that protecting biodiversity was an The Bornean orangutan (Pongo pygmaeus) and
ultimatum from outside interests who possessed neither sym- Sumatran orangutan (Pongo abelii) are arboreal,
pathy nor understanding of the church’s mission, the Maro- forest-dwelling great apes whose populations are now
nite Church declined to enroll the forest as an area of restricted to the islands of Borneo and Sumatra. IUCN
protected status. Given the failure of this effort, ARC and currently lists both species as Critically Endangered, as
WWF tried a different tact. Working with more locally-based well as the more recently described orangutan species
forest conservation groups, they met directly with the patri- from Sumatra, the Tapanuli orangutan (Pongo
arch of the Maronite Church, and almost immediately gained tapanuliensis). All three populations have been declin-
a commitment to protect the forest. The reasons for that ing rapidly, primarily due to over 60% forest habitat
protection were framed in a new statement, written mainly loss in the last 60 years, accelerated by logging, indus-
by the patriarch, who expressed the new proposal in different trial agriculture, industrial plantation forestry, and for-
words for a different purpose: est fires. Apart from habitat loss and degradation,
For centuries the Church has defended the natural beauty and
orangutans are also beset by the illegal wildlife trade.
Godliness of the forests and hills of Harissa, as well as many Infant orangutans bring high prices as pets, creating
other holy places in Lebanon. . . .In so doing, we observe that the motivation to capture them by killing their mothers and
land and the flora and fauna on it, do not ultimately belong to then taking the infants to market (Fig. 10.19). In
us. We are simply the guardians of what belongs to God. . . .In
protecting this area, the Church will continue to ensure that the
Indonesia such practices are illegal, but common,
diversity of plants, trees, animals and birds given by God, nur- because laws against such actions are not consistently
tured by the Church, will be maintained. . . (quoted in Palmer and enforced, and profits are high.
Findlay 2003:9). Enforcement officers arrest some traders, but that
The contrasts of the two approaches reveal some of the only solves half the problem. The infant orangutans
reasons why FBOs in conservation succeed where secular cannot simply be released back into the forest because
conservation NGOs fail. UNEP felt that the presence of
(continued)
unique biodiversity within the Forest of Harissa was
Fig. 10.19 A caged infant orangutan (Pongo pygmaeus) awaits sale in Fig. 10.20 Taymur the orangutan meeting a veterinarian and his secu-
a pet market in Eastern Asia. Although illegal, such trade in infant rity personnel, after his flight home. His rescue was due to the immediate
orangutans is common because profits from sales are high and laws action undertaken by the Borneo Orangutan Survival (BOS) Founda-
against such trade are not consistently enforced. (Photo reprinted by per- tion, the Ministry of Environment and Forestry, and the Ministry of
mission of P. Hilton, # Paul Hilton) Foreign Affairs via the Indonesian Embassy in Kuwait in cooperation
with the Kuwait Zoo. (Photo courtesy of the BOS Foundation)
they lack the skills to live alone without their mothers.

Despite some government support for rehabilitation Rehabilitation Center in Indonesia’s Central Kalimantan
efforts for infant orangutans, the Indonesian govern- province. Here infant orangutans rescued from the pet
ment lacks funds to fully “rehabilitate” all orphaned trade are nursed back to health with proper diet and
orangutans that are taken out of the pet trade, but medical attention, then socialized with other orangutans
several non-governmental conservation organizations and helped to acquire forest knowledge and skills
have taken responsibility for implementing and (Fig. 10.20). In “Baby Forest School” they begin to
funding this work. The largest implementation learn foraging, nesting, arboreal locomotion, predator
organizations currently rehabilitating orangutans in avoidance, and other skills needed for their natural life-
Indonesia include the Borneo Orangutan Survival style. Orangutans from two and a half to five years old
(BOS) Foundation, International Animal Rescue are moved to “Forest School” where, in a forest, they
(IAR), Orangutan Foundation International (OFI), progress to learning advanced arboreal and foraging
Sumatran Orangutan Conservation Programme abilities in a larger natural environment. Orangutans
(SOCP), and the Centre for Orangutan Protection over 5 years old that complete training at Forest School
(COP). Funding for such efforts is global in nature, are moved to forested “pre-release islands,” where they
and comes from members of the public, international begin to live semi-independently. The final step is to
aid, grant-giving institutions, and several dedicated, release rehabilitated orangutans to a mainland expanse
international fundraising organizations, including the of protected forest. There is constant pressure to release
Orangutan Conservancy (OC), Orangutan Outreach, rescued rehabilitated individuals due to overcrowding in
The Orangutan Project (TOP), Save the Orangutan the rehabilitation centers. In Nyaru Menteng alone, there
(STO), and BOS Foundation. are approximately 400 orangutans (as of October 2018)
Many of the current efforts are being led by Indone- awaiting release or improved sanctuary care.
sian conservationists with support from national and The problem of orphaned orangutans illustrates both
international partners. As the problem of orphaned the complex nature and practical role of ethics and
orangutans has become more widely known, conserva- values in the work of conservation biology, and the
tion scientists, veterinarians, local Indonesians and interaction of ethics and values with science, econom-
foreigners from all walks of life have collaborated to ics, law, and culture. In every step of this example,
establish and support rehabilitation centers whose goal opposing forces of motive and restraint interact to
and curricula are to return orphaned orangutans to wild either make progress toward a solution or further exac-
environments. Among these projects, the largest has erbate the problem (Fig. 10.21). Initially, economic
been BOS Foundation’s Nyaru Menteng Orangutan reward provided motive that created large numbers of
Fig. 10.21 Actions associated

with problems of and potential
solutions for orphaned orangutans
(Pongo pygmaeus) in Indonesia,
motives that drive them, and legal
and ethical restraints that
constrain such motives and their
effects. (Figure design by F. Van
Dyke)
orphaned orangutans. If economic considerations are as economic motives helped create the problem, eco-
used as a basis for normative decision making, killing nomic restraints perpetuate it. Many conservationists
mother orangutans and selling their infants in pet believe that even after release in a “protected” area in
markets makes sense. Even with the risk of arrest and Indonesia, economic motives for profit from such
fines or imprisonment, the financial reward is great reserves, especially timber and mining, could under-
enough that the cost-benefit ratio leans heavily in mine laws established to protect the reserves,
favor of continuing these practices. Such actions are jeopardizing orangutans and other elements of
restrained, if imperfectly, by conservation laws Indonesia’s natural heritage.
intended to affirm the intrinsic value of orangutans Regardless of what species conservation efforts try
and overall biodiversity. But economic profit, not to protect, every conservation biologist must realize
only from the sale of infant orangutans, but also that the understanding and expression of conservation
manifested as bribery and corruption within the ranks values are essential for the effort to succeed. Until local
of enforcement officers and government officials, often residents and the international business and trade com-
overwhelms social and legal affirmations of the munity incorporate a conservation ethic that sees intrin-
orangutan’s intrinsic value, supported and informed sic value in the orangutan, it will continue to make
by careful studies of orangutan diets, their social and sense to capture young orangutans and sell them as
reproductive behavior, and habitat needs. But the pets to realize their instrumental value as economic
actions taken to affirm the orangutan’s intrinsic value profit. Until more affluent people in Indonesia and
are themselves constrained by lack of financial and around the world see the importance of practicing
human resources. The government of Indonesia does conservation ethics that curb demand for tropical forest
not value orangutans enough to divert scarce resources products and their associated habitat destruction, the
from pressing human needs to build more rehabilitation endangerment of the orangutan will remain a signifi-
centers, and orangutan conservation organizations face cant and tragic conservation problem that can only be
an ongoing struggle to finance a complete solution. Just solved by ethics that changes human minds and hearts.
(continued)
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Conservation Economics and Sustainable
Development 11
Economics is about the broad world of understanding how the incentives we face affect the decision we
make. . .The causes of biodiversity loss are founded in economics. Land-or-resource use decisions that affect
the nature world are grounded in economics. And most importantly, the solutions and their effectiveness are
grounded in economics.
Fisher et al. (2015:2–3).
Keywords 11.1 The Role of Economics in Conservation

Ecosystem services · Ecosystem valuation · Stock-flow
and fund-service resources · Revealed preference 11.1.1 Thinking Like an Economist
methods · Stated preference methods · Market-based
mechanisms · Ecological economics · Conservation Economics is the study of “how people make choices under
easements · Integrated conservation and development · conditions of scarcity, and of the results of those choices for
Payments for ecosystem services society” (Frank and Bernanke 2003). Although some
conservationists view economics as a persistent impediment
to conservation goals, thinking like an economist can help us
better understand how to make tough conservation decisions
Overview in the face of resource scarcity and constraints (Fisher et al.
In this chapter you will learn about: 2015). In fact, conservation organizations are very familiar
with the challenges of achieving their goals amidst a lack of
1. the role of economics in conservation theory and resources. Often these organizations are pushed to identify
practice. the most strategic areas for investment with the recognition
2. the characteristics of traditional (neoclassical) eco- that the choice of one activity means they must forgo another.
nomic theory and alternative views associated with A popular term for this tension is conservation triage, where
ecological economics. continuous threats to biodiversity and inadequate funding
3. economic tools and incentives that can contribute to make it inevitable that conservation managers apply tradeoffs
environmental protection. in decision making. As Bottrill et al. 2009 write, “In practice,
4. government-market interactions in support of con- all conservation managers and agencies allocate limited
servation policy. budgets to specific actions in the knowledge that there will
5. intersections between economics and sustainable be habitats and species that receive no, or less, investment
development. and that these might degrade or become extinct owing to the
choices made” (183).
Thinking like an economist can also help motivate con-
servation practice, as economic behavior is arguably one of
the most revealing expressions of national, corporate, com-
munity, and individual values. No conservation effort can

450 11 Conservation Economics and Sustainable Development
long endure without intimate connection to value. And few each of the foundational frames outlined here will help us
expressions of conservation can endure with vitality without identify how the economic benefits of nature for human
expressing those values in part through economic behavior, welfare can motivate conservation as well as finance it.
either through what we spend to protect what is needed by
other species, or what we do without in order that what is
needed by other species is not consumed and destroyed. 11.1.2 Ecosystem Services
Taking an economic approach to conservation does not
imply that intrinsic values and related motivations (Chap. Economic value is manifested in tangible goods (material
10) are less important, as moral imperatives do not require resources) and services (functions of value to us performed
economic justification. However, utilizing an economic by some other entity). Ecosystem services refer to particular
worldview can help us better navigate human biases, ecosystem conditions and functions that have value to
emotions, irrationalities, and the social context of economic humans – that sustain and fulfill human life (Daily 1997,
decision-making (Kahneman et al. 1991). Costanza et al. 1997). Although there is disagreement
Any discussion of economics in the context of conserva- among ecologists and economists about different categories
tion can be approached from two primary branches of study, and definitions of ecosystem services (e.g., Fisher et al. 2009,
microeconomics or macroeconomics. Microeconomics Boyd and Banzhaf 2007, Bateman et al. 2011), such services
centers on incentives that influence individual behavior are classified within the Millennium Ecosystem Assessment
within the economy and their intersections with private and (MEA) (2005) as provisioning (e.g., food and freshwater),
community property rights. This branch of economics is supporting (e.g., soil formation and nutrient cycling),
focused on markets, what factors influence the goods and regulating (e.g., flood and disease regulation) and cultural
services individuals consume, and the less-than-optimal (e.g., education and recreational). Subsequent work under
outcomes that result due to externalities (true costs and The Economics of Ecosystems and Biodiversity (TEEB)
benefits left outside of the market). We can also consider study, supported by Germany and the European Commission,
several larger challenges for conservation within the econ- has sought to clarify these concepts in order to prevent double
omy as a whole (i.e., macroeconomics) by determining the counting in ecosystem service audits and to ensure a broad
appropriate level of economic activity within society or view of ecosystem value (TEEB 2010, Fig. 11.1).
deciding how the government ought to think about its spend- Consider the services of a single ecosystem, a forest
ing or revenue streams. Both micro- and macroeconomics (Table 11.1). As you read this list, reflect on how many of
provide important considerations for conservation, but it is these services you actually think about paying for on a daily
often microeconomics that is reflected in the day-to-day basis. If your honest answer is few or none, you are not alone.
mechanics of conservation programs. Throughout history, humans have depended on these
Neoclassical economic theory forms the basis of much of services, but routinely take them for granted. These services
the world’s current economic practices and is consequently were simply expressions of “the way nature works.”
the focus of many of this chapter’s case studies. Widely In an effort to clarify ecosystem terms for economic anal-
accepted in the nineteenth century, particularly across ysis, Fisher et al. 2015 describe them this way:
Europe, neoclassical economics rose alongside the industrial
revolution as the cornerstone explanation for human eco- • Ecosystems are combinations of interacting species and
nomic behavior. However, there have also been critiques of their environment. They do many things, not all of which
neoclassical economics, and we will consider those levied by are ecosystem services (e.g., a wetland).
ecological economists in larger discussions around “sustain- • Ecosystem services are the subset of these natural process
able” economics and international development. that actually generate benefits to people (e.g., water puri-
Finally, there are three foundational economic concepts fication). Sometimes this is the production of a good.
embedded in many conservation efforts: ecosystem services, • Benefits are the actual things that increase human welfare
stock-flow and fund-service resources, and nonexcludable (e.g., clean water).
and nonrival goods. Over the past few decades, efforts to • Value is the importance of a given benefit to a person or
capture the value of nature through a market price have led to group, measured in dollars or other metrics (e.g., $30/gal/
many creative programs and projects around the globe, but year). This is often based on culture, market, norms, or
there is disagreement among conservationists about the other factors.
implications of hinging motivations for conservation on eco-
nomics. Some research shows that when social interactions The strength of connection between these services and
become financial ones, economic incentives “crowd out” human well-being varies across ecosystems and regions, but
altruistic motivations, causing individuals to become even global implications of their depletion or degradation are
less likely to conserve nature for the common good (Kerr serious. The 1360 scientists who compiled the World
et al. 2012, Fisher et al. 2015). When more closely examined, Resources Institute’s MEA at the behest of the United
11.1 The Role of Economics in Conservation 451
Fig. 11.1 Changes in drivers that indirectly affect biodiversity, such as can take place across different time scales. (Reprinted by permission
population, technology, and lifestyle (upper right corner), can lead to from the World Resources Institute, from Millennium Ecosystem
changes in drivers directly affecting biodiversity. These result in Assessment. 2005. Living beyond our means: natural assets and
changes to ecosystems and the services they provide (lower left corner), human well-being. Island Press, Washington, DC. # 2005 World
thereby affecting human well-being. Such interactions take place at Resources Institute)
multiple geographic scales and across scales. Similarly, the interactions
Nations found that, of 24 ecosystem services examined 11.1.3 Stock-Flow Resources and Fund-Service
worldwide, 15 (60%) were being degraded or used Resources
unsustainably. As ecosystems, and the services they provide,
are degraded, global biodiversity is diminished. Results of The material goods we use and derive from ecosystems and
the MEA demonstrated that failing to include the value of their associated biodiversity are diverse. Stock-flow
ecosystems has left critical dimensions of human well-being resources are goods produced through a transformation pro-
out of economic decision-making. As ecosystem services cess, usually self-renewing, that occurs within the ecosystem
become scarce resources, we will have to pay higher prices itself. Goods produced in this category can then be extracted
to use them. Thus, efforts to more accurately value the for use at a certain rate or “flow” from the standing crop or
benefits of ecosystem services are increasingly important “stock” of a resource. For example, timber from a forest
within current economic analysis and policy development. ecosystem is a stock-flow resource in which a standing crop
Table 11.1 Examples of ecosystem services provided by a forest

Ecosystem service Examples from forests
Gas regulation Trees store CO2 and growing trees create O2; forests can clean SO2 from the atmosphere.
Climate regulation Greenhouse gas regulation; evapotranspiration and subsequent transport of stored heat energy to other regions by
wind; evapotranspiration, cloud formation, and local rainfall; effects of shade and insulation on local humidity
and temperature extremes.
Disturbance regulation Storm protection, flood control (see water regulation), drought recovery, and other aspects of habitat response to
environmental variability controlled mainly by vegetation structure.
Water regulation Tree roots aerate soil, allowing it to absorb water during rains and release it during dry periods, reducing risk and
severity of both droughts and floods.
Water supply Evapotranspiration can increase local rainfall; forests can reduce erosion and hold stream banks in place,
preventing siltation of in-stream springs and increasing water flow.
Waste absorption capacity Forests can absorb large amounts of organic waste and filter pollutants from runoff; some plants absorb heavy
metals.
Erosion control and sediment Trees hold soil in place, forest canopies diminish impact of torrential rainstorms on soils, diminish wind erosion.
retention
Soil formation Tree roots grind rocks; decaying vegetation adds organic matter.
Nutrient cycling Tropical forests are characterized by rapid assimilation of decayed material, allowing little time for nutrients to
run off into streams and be flushed from the system.
Pollination Forests harbor insects necessary for fertilizing wild and domestic species.
Biological control Insect species harbored by forests prey on insect pests.
Refugia or habitat Forests provide habitat for migratory and resident species, creating conditions essential for reproduction of many
of the species they contain.
Genetic resources Forests are sources for unique biological materials and products, such as medicines, genes for resistance to plant
pathogens and crop pests, and ornamental species.
Recreation Ecotourism, hiking, biking.
Cultural Aesthetic, artistic, educational, spiritual, and scientific values of forest ecosystems.
Reprinted by permission from Island Press, from Daly, H.E., and J. Farley. 2011. Ecological economics: Principles and applications. (2nd edition).
Washington, DC: Island Press. # 2004 & 2011 The Authors
or stock of mature trees transforms water, sunlight, and activity and land-use change (Keenan and Williams 2018).
nutrients from the soil into new biomass (the trees get bigger) By removing this carbon from the atmosphere, such
and new individuals (seeds that germinate into new trees). If a ecosystems provide a collective service of climate regulation
stock-flow resource (like timber) is harvested or extracted at a and moderation. Although we can expand this capacity via
rate less than or equal to the rate of its renewal process, it is afforestation, such a service cannot be stockpiled (we cannot
self-perpetuating. We can manage such a resource for “store up” capacities for carbon removal this year to be used
“sustained yield” much as we could “harvest” interest income next year). The service must be delivered at a fixed rate.
from a bank account as long as we leave the principal intact. Thus, although the service cannot be “used up,” it can be
All stock-flow resources share common attributes. First, “worn out” if such ecosystems are degraded to the point that
stock-flow resources are those that have been materially their ability to remove atmospheric carbon is diminished. By
transformed by ecosystem processes into usable goods. Sec- clearly distinguishing between these two resource types, we
ond, they can be used at any rate desired, but, in many cases, are better able to define the parameters of so-called “sustain-
some rates of use are too great to be sustained indefinitely. able” use.
Third, stock-flow resources can be stockpiled or “stored up”
for future use. They do not have to be used all at once, even if
taken out of their ecosystem context. Finally, stock-flow 11.1.4 Nonexcludable and Nonrival Goods
resources can be used up, but not worn out.
In contrast, ecosystems also provide fund-service As essential as such goods and services are, they have proved
resources. A fund-service resource can suffer wear and tear repeatedly problematic for traditional economic theory,
from a production process, but it does not become a part of which seeks to classify all goods across a continuum of
the thing it produces. Instead, the fund provides the service at excludability and rivalness (Fig. 11.2). An excludable good
a fixed rate, so the service is best measured in some metric or resource is one in which ownership of the resource permits
that describes output over time. For example, from 2007 to the owner exclusive use of the resource and provides the
2016, terrestrial ecosystems globally removed an estimated owner with the ability to exclude others from such use. A
3.61 Pg C per year from the atmosphere, which amounts to rival good or service is such that the enjoyment or use of the
33.7% of total anthropogenic emissions from industrial good or service infringes on someone else’s ability to enjoy
11.1 The Role of Economics in Conservation 453
Fig. 11.2 Simplified matrix of

the rivalry and excludability of
economic goods. (Graphic design
by R. Lamb)
the same good or service. If a pizza is set before us, every freshwater by one person or group in a discrete location
piece that I eat is a piece that you cannot eat, and vice versa. may limit the availability of freshwater to other users at that
Thus, the pizza is a rival good. Because of their dependence same location (thus making it a common pool resource,
on market mechanisms to allocate and set prices for rather than a public good, for stakeholders affected at that
resources, neoclassical economics has proven adept at the site) (Kretsch et al. 2016). Common pool resources provide
valuation and distribution of rival and excludable goods and particular challenges in that the ongoing and increasing use of
services (also known as private goods), and those goods a resource by those who are not excluded from it, or whose
which, although non-rival, can still allow for excludability use is otherwise unregulated, can result in the decline of
with appropriate pricing (toll or club goods). overall system health. Fisheries in the ocean is a common
Many ecosystem goods and services are non-excludable. pool resource under pressure as many countries harvest as
How would one, for example, exclude others from receiving many fish as possible to maximize their own profits. The
the oxygen produced by the photosynthesis of trees in a result is a stock-flow resource tapped beyond its capacity to
forest? Even if one owned the forest, the breathable air it self-renew (Chap. 8).
produces flows all over the planet, and all receive its benefits. Economists Herman Daly and Joshua Farley summarize
Some ecosystem goods and services are also non-rival, in the excludability problem facing many ecosystem services
that one person’s use or enjoyment of the good does not eloquently: “If a good is not excludable, someone can use it
affect or infringe upon the ability of others to use or enjoy whether or not any producer of the good allows it. If people
the good. If an upstream wetland absorbs most of the water can use a good regardless of whether or not they have to pay
from a sudden downpour, and thus protects my downstream for it, they are considerably less likely to pay for it. If people
riverfront home from being flooded, it also protects my next- are unwilling to pay for a good, there will be no profit in its
door, streamside neighbor just as much. My benefit of pro- production, and in a market economy no one will invest in
tection from the wetland does not in any way infringe on my producing it, or at least not to the extent that the marginal
neighbor’s benefit of protection. We are both high and dry. benefit to society of producing another unit is equal to the
Consequently, many ecosystem services such as fresh water marginal cost of production (Daly and Farley 2011).” In
or a stable climate are seen as public goods (non-rival and short, Daly and Farley see public goods and common goods
non-excludable) and fall outside the marketplace. (or common pool resources) as challenging to both protect
Although public goods may be viewed by economists as and to restore once degraded if the benefits of its use are not
providing immutable (i.e., unchanging and everlasting) connected to individual or corporate profit.
benefits to society, the reality is much more complicated. In the following section, we will consider how the tools of
The need for some form of governance for public goods microeconomics may be used to reduce or resolve some of
(to avoid overuse or under-provision) usually arises when the challenges associated with the maintenance and protec-
there is some degree of rivalry or when changes in land tion of ecosystem goods and services, and evaluate the
management or ownership threatens non-excludability respective roles of institutions in incentivizing such behav-
(Kretsch et al. 2016). For example, over-exploitation of ioral change.
11.2 Microeconomic Approaches

to Conservation Dilemmas
11.2.1 Fundamental Assumptions of Supply

and Demand
In neoclassical economic theory, market price for any good

or service stems from supply and demand – the buying,
selling, trading of these resources based on the behavior of
those that supply the good and those that are the consumers,
or buyers, of it. Higher demand for a particular resource is
generated because the price gets lower while an increase in
supply to the marketplace occurs because there is a potential
to make more profit – buyers are willing to pay more for the
good or service (Fig. 11.3). Although each individual has
their own supply and demand curves for a particular good or
service – showcasing how much one is willing to pay under
which conditions – most of the curves analyzed by
economists are aggregates – they attempt to explain the
cumulative factors and motivations affecting supply and Fig. 11.3 Supply and demand curves, where the market clearing price
demand across the entire marketplace. (or equilibrium price) is given at point (q, p). As the demand or supply
Applying these concepts to conservation, Zhao et al. con- curves shift, the market clears at a different price and quantity. For
example, the demand curve might shift to the right, as shown here,
sidered how quantifying the supply and demand factors in due to changes in seasonal demand for certain products. (Graph design
soil conservation, and the feedbacks between them, could by R. Lamb)
lead to improved ecosystem management (2018, Fig. 11.4).
Here, soil conservation was considered the ecosystem ser- of ecosystem restoration locally and the transmutation of
vice, with the corresponding capacity to control erosion and these effects across time and space. By taking all of these
facilitate sedimentation depending on ecosystem structure things into account via several “on the ground” case studies,
and land management (Zhao et al. 2018). Although many the authors were able to make targeted recommendations for
studies focus on maintaining the “supply” of this service managing both the ecosystem service and human demand for
(by protecting ecosystem function), the authors insisted that certain benefits. As a next step from this study, a formal
the quantification of both supply and demand (in this case the economic market could be created to more accurately reflect
human benefits derived from soil conservation) would the “price” of using this service based on the quantification
improve decisions made about the tradeoffs between them. and intersection of these supply and demand curves.
Or, using the terms defined by Fisher and colleagues in Sect. Economists assume that consumers purchase the combi-
11.1.2, by quantifying the benefits of soil conservation, based nation and quantity of goods that maximize their own well-
on the health of the ecosystem service, they could help clarify being, or utility, while producers simultaneously make ratio-
the economic value of that service to people. nal decisions to provide such goods based on potential for
Zhao et al. focused on both the maximum allowable profitability. Further, economic theory assumes that these
erosion rate (the maximum rate at which this fund service market agents account for all potential costs, the full oppor-
resource could be used without being worn out) and the tunity cost, given the consumer or producer’s own resource
current erosion rate. If the current erosion rate exceeded the restraints. From here, economic analysis can illuminate the
maximum, there were negative consequences for ecosystem maximum price (P) an individual with (Q) units of a good or
supply. However, if the current erosion rate was below the service would be willing to pay for an additional unit (mar-
maximum, the implications for the ecosystem supply were ginal benefit). Such analysis can also identify the cost of
positive. In quantifying the demand, the authors identified the producing the additional unit (known as the marginal cost).
individual beneficiaries and their locations (the benefits may The quantity and price where marginal cost equals marginal
change across scales – upstream, midstream, downstream) benefit is known as the market equilibrium.
and various indicators of socio-economic benefit (increase in The maximization of consumer and producer surplus, and
food production, extended reservoir operation period, and the creation of economic efficiency, is expected to occur
improvement in water quality). They also considered the automatically without further intervention from the govern-
feedbacks between supply and demand, such as the effect ment or other institution. An efficient market then is one
11.2 Microeconomic Approaches to Conservation Dilemmas 455
Fig. 11.4 A general framework to couple human–nature systems by potential land-use change influencing such characteristics. (Reprinted
soil conservation service assessment; where supply is characterized by by permission from Elsevier, from Zhao, W., Liu, Yue, Daryanto, S., Fu,
ecosystem processes and functions, and demand is quantified as the total B., Wang, S., Liu, Yanxu, 2018. Metacoupling supply and demand for
human benefits derived. Taking a broader view, demand may be soil conservation service. Current Opinion in Environmental
influenced by policy goals such as the UN’s Sustainable Development Sustainability 33, 136–141. # 2018 Elsevier)
Goals, while supply is influenced by both ecosystem characteristics and
assumed to maximize social welfare, such that current levels

of production allow society to derive the largest possible net
benefit from the market (EPA 2010). However, it does not
guarantee an “equitable” or “fair” distribution of good pro-
duction or consumption across all those participating in the
market. Unincorporated ecosystem goods and services are
examples of market failure, where the market does not
fully represent social values due to externalities, and the
true costs and benefits of an activity or good are not linked
to consumer or producer decision-making.
11.2.2 The Challenge of Externalities
One of the challenges of economics is to create markets for

environmental entities for which a market did not previously
exist. Properly developed, environmental markets can stimu-
late market efficiency and improve overall economic effi- Fig. 11.5 The impact of a negative externality on a general supply and
ciency, while supporting higher levels of biodiversity demand curve where, D is the market demand (marginal benefit) curve
for the product; MPC is the firm’s marginal private real-resource cost of
conservation. To do this, economists must address production, excluding the cost of the firm’s pollution on households;
externalities, the “costs or benefits that are not explicitly MSD is the marginal social damage of pollution (or the marginal exter-
captured in a decision or action” (Fisher et al. 2015). A nal cost) that the firm is not considering; and MSC is society’s marginal
classic example of a negative externality (Fig. 11.5) is air social cost associated with production, including the cost of pollution
(MSC ¼ MPC + MSD). For example, a firm producing some product
pollution, where some economic costs (i.e., damage to prop- might also be generating pollution as a by-product. The pollution may
erty and infrastructure and loss of productivity of people and impose significant costs – in the form of adverse health effects, for
crops), social costs (i.e., physiological impacts on human example – on households living downwind or downstream of the firm.
beings, mostly to the cardiovascular and respiratory systems) Because those costs are not borne by the firm, the firm typically does not
consider them in its production decisions. Society considers the pollu-
and environmental costs (i.e., damage done to ecosystems tion a cost of production, but the firm typically will not. (Graph from
through the atmosphere) are borne by many other people EPA 2010)
beyond, and often not including, the powerplant’s owners simplify complex decisions and compare policies, projects or
(Rodrigue et al. 2017). Consequently, these costs are not programs, but it also carries risks. Environmental benefits can
factored directly into the producer’s decision-making (or in be hard to measure, even when costs of regulation are known
reality, the consumer’s economic decision-making, when (i.e., increased cost of scrubbers on smokestacks vs. lives
they purchase the artificially inexpensive products created saved with cleaner air). Benefits also tend to accrue over time
as a result of this production processes). In contrast, ecosys- while costs are immediate. And, even though the common
tem services are generally considered positive externalities, metric of the dollar can be viewed as an advantage, it is
where the benefits from ecosystem conservation accrue to difficult to convert all types of costs and benefits to a single
others beyond the landowner (recall the earlier example of a unit without some degree of uncertainty or subjectivity. Such
non-excludable public good like oxygen provided by trees an approach also may obscure exactly what is being
planted in an individual’s yard). measured (such as the potential number of live lost) (EPA
The traditional solution to externalities is to internalize, or 2010).
include them in the market itself – giving a private incentive The US Environmental Protection Agency, well known
to provide a common good (Fisher et al. 2015). One actor that for its use of CBA to justify its environmental policies, has
corrects market failures is the government, as it makes laws, been the subject of increasing scrutiny over what it chooses to
enforces standards, levies taxes, or creates subsidies on include or exclude from its analyses. In May 2019, the Trump
behalf of the public good (Sect. 11.4). Businesses and Administration sent a memo to EPA staff notifying them of a
individuals can also influence the market towards inclusion. review of existing CBA process. The purpose was to make
Although the primary goal of these actors is to maximize their changes in how costs and benefits are currently calculated –
own utility, the ecosystem services that may be currently namely how human and ecological benefits are calculated
unincorporated could be realized (or better protected) to (Green 2019). The Trump administration had long argued
their benefit. A common tool that actors use to justify their that the EPA, under the previous Obama Administration,
decisions and potential policy interventions is cost benefit overestimated health risks of various environmental
analysis (CBA), otherwise known more positively as benefit regulations to the determent of industry. Therefore, a change
cost analysis. If the value can be quantified, ecosystem goods in calculation strategy might showcase (preferentially for
and services can be compared alongside other known costs industry) a different economic result. The potential environ-
and benefits. mental and social implications of such a methodological
change are many. For example, Trump’s EPA had initially
calculated that repealing and replacing Obama’s Clean Power
11.2.3 Cost Benefit Analysis Plan, a climate policy focused on reducing emissions from
coal fired powerplants, with its proposed Affordable Clean
CBA is a tool we have explored earlier in regard to its role in Energy (ACE) rule, would result in an additional 1400 pre-
ethic and value analysis, especially its use in evaluating mature deaths per year. However, with the new proposed
instrumental value (Chap. 10, Sect. 10.3.2). From an eco- CBA model, the number of deaths would be significantly
nomic perspective, CBA helps economic actors decide if a reduced (Green 2019). This example of politically-driven
decision (be it management or policy oriented) is warranted economic analysis emphasizes the importance of transparent
or not – in other words, it helps answer questions such as: CBA processes open for public comment and review, and the
Will we gain more than we lose in economic terms? What are need for improved methods and incorporation of ecosystem
our options? What if we do nothing? Remember that neoclas- service valuation.
sical economics assumes all actors in the market are con-
stantly making tradeoffs, and that with perfect information,
the resulting decision(s) will be the most efficient. You might 11.3 Methods for Valuing Ecosystem Goods
immediately recognize, given our previous discussion about and Services
externalities, that the CBA is only “efficient” if all relevant
factors are included in the analysis. Economic cost-benefit 11.3.1 Should We Price Nature?
analysis must consider a range of views, including perhaps
what happens to non-human species. Each actor using this There are many who view putting a price on nature to be a
tool must first define who they will include (perspective), for vital tool for nature conservation today. As long as the world
how long (time), and which scenarios they are willing to continues to use neoclassical economics to frame its CBA,
consider (alternatives) relative to their goals. there is a strong rationale for including more precise
Because CBA represents these considerations in economic (in amount and quality) information on ecosystem values.
terms, a common dollar sign lets you compare different units. Investing in scientific assessment of such ecosystem services
Such analysis is praised for its transparency and ability to is a basic starting point that could be further motivated by
11.3 Methods for Valuing Ecosystem Goods and Services 457
policy demand. For example, one of us (Lamb) has worked and John Loomis estimated the economic value of various
on NASA Carbon Monitoring System (US) projects that seek measures of stream restoration to private property values
to improve our estimations of existing forest carbon stocks using a hedonic property model that treats property value
and fluxes as well carbon sequestration potential as forests as a function of its structural characteristics (S), neighbor-
mature (Hurtt et al. 2019). This work is done in large part hood (N ), and environmental quality (Q), expressed in the
because this science is requested by state governments inter- identity
ested in improving their carbon budgets relative to climate
mitigation goals. Given its spatial precision, this same sci- Pi ¼ f ðSi , N i , Qi Þ
ence could also be used to develop an afforestation incentive
program with a price on carbon. Ignoring these ecosystem where the subscript i refers to each value for an individual
values has shown to result in market failure and socially (ith) property (Streiner and Loomis (1996). Because
undesirable outcomes. Further, as Fisher et al. 2015 argue, attributes of property value are typically grouped, Streiner
ecosystem valuation may also help call attention to the value and Loomis created “restoration packages” that could be
of nature in our lives and aid us in determining the size of applied to different properties. Restoration package A
incentives necessary to encourage provision of these benefits included improving fish habitat and the acquisition of addi-
for society. tional land along the stream by the California Department of
In microeconomics, there are two general classes of valu- Water Resources Urban Stream Restoration Program for a
ation techniques for ecosystem services: revealed preference streamside education trail. Restoration package B featured
and stated preference methods. Revealed preference properties where streams were restored in ways that reduced
methods “use observed behavior such as market purchases flood damage, cleaned up, revegetated and stabilized the
to make inferences on values of goods and services” (Fisher stream bank, cleared obstructions from the stream channel,
et al. 2015). Such approaches work with either actual markets and added aesthetic elements such as check walls, rock or
or proxy markets, such as the market for property which stone walls or wood plank walls along the stream.
reflects the various environmental attributes of the property Streiner and Loomis determined that individual elements
itself. Stated preference methods “ask people how much they of package A added US$15,000 to US$19,000 in property
are willing to pay/willing to accept for the gain or loss of a value. In package B, only stabilization and reduced flood
non-market good or service” (Fisher et al. 2015). Stated damage added value, but these still increased worth by up
preference approaches create hypothetical markets for envi- to US$7800. A joint model incorporating dimensions of both
ronmental benefits where there are currently no observable packages added over US$19,000 in value to an average value
market prices. Both revealed preference and stated preference to individual properties (Table 11.2, Streiner and Loomis
methods attempt to incorporate ecosystem values into the 1996). This analysis demonstrated that not every type of
marketplace. restoration added significantly to private property value, but
many did. Restorations that added value included both efforts
that restored inherent functions of the stream (e.g., improved
11.3.2 Revealed Preference Methods fish habitat) as well as those that more directly benefited the
property owner (e.g., reduced flood damage).
11.3.2.1 Hedonic Valuation Models for Private There are many global examples where proximity to or
Property inclusion of nature increases property values (i.e., Thorsnes
One of the long-term goals of conservation in socio- 2002; Morancho 2003; Radeloff et al. 2010). Although eco-
economic context is to move local residents from a reactive nomic literature still uses the term hedonic pricing to describe
posture that denies responsibility for the current state of a site this economic technique, the nascent literature on ecosystem
to a proactive posture that not only accepts, but anticipates, services has instead begun to use the term production func-
responsibility for the site and does even more than is required tion as the method relates directly to the how the “value of
in its restoration (Clarkson 1995). One way to illustrate these any particular good or service is ‘produced’ by a variety of
principles is to see how environmental quality can be treated factors” (Fisher et al. 2015).
as a “value added” dimension of private property exchange in Ecological economist Taylor Ricketts and colleagues
a residential setting. To understand how this works, we look adopted this production function approach when considering
to an application of this method in the US state of the economic value of forest proximity to coffee production
California. (2004). In this study, the authors focused on a single large
The California counties of Contra Costa, Santa Cruz and (1065 ha) coffee farm in Costa Rica and considered data on
Solano surrounding the San Francisco Bay area include resi- pollination and farm yields across a gradient of distance from
dential property valued among the highest in the United two adjacent forest patches (Fig. 11.6). Only feral
States. In these counties, resource economists Carol Streiner Africanized honeybees were present in the area, which
Table 11.2 Values of alternative “conservation packages” associated with stream restoration efforts adjoining private residential property in Contra
Costa, Santa Cruz, and Solano Counties, California. Values in US$
Restoration measure Absolute value of restoration Value of restoration relative to property value (%)
Restoration package Aa
Fish habitat improvement $15,571 11
Land acquisition for education trail $19,123 13
Education trail established $17,560 12
Restoration package Ba
Streambank stabilization $4488 3
Reduced flood damage $7804 5
Joint model
Education trail established with streambank restoration $19,078 13
Adapted from Streiner, C.F., and J.B. Loomis. 1996. Estimating the benefits of urban stream restoration using the hedonic price method. Rivers 5:
267–278. # S.E.L. & Associates
a
Note: The individual measures cannot be added together because they are simply alternative measures of the joint effect of all these variables in the
package
“pollination by wild bees increased coffee yields near forest

patches. Ambient pollination services were adequate in near
and intermediate sites; neither seed mass nor fruit set
increased with hand-pollination” (Ricketts et al. 2004). The
authors then combined the results of these experiments with
market prices to estimate the income contributed to the coffee
farm by the two major neighboring forest patches (Ricketts
et al. 2004). They found that the surrounding forest
“contributed an average of US $62,000 per year (i.e., 7% of
total farm income) in 2000–2003” which was “an order of
magnitude greater than that recognized by Costa Rica for
other forest ecosystem services, of at least the same order as
major competing land uses, and infinitely greater than that
recognized by most governments (i.e., zero)” (Ricketts et al.
2004). In other words, given the authors’ production function
(relating coffee yield to forest productivity), if the “two
specific forest patches were destroyed, much of the farm
would wind up more isolated from tropical forest than it
had been and the owners would lose coffee production
worth $60,000 in net revenue” (Fisher et al. 2015).
11.3.2.2 The Travel Cost Method – A

Behavior-Based Technique
The Hedonic Travel Cost Method (TCM) works on a simple
Fig. 11.6 Map of study area and sites in the Valle General, Costa Rica. but reasonable assumption: the more valuable an environ-
The focal coffee farm, Finca Santa Fe (1065 ha), is in white; stippled mental resource or amenity, the farther people are willing to
area is a mix of coffee, pasture, and sugar cane; black areas are forests. travel to experience it, the more they are willing to spend per
The three focal forest patches are labeled A (46 ha), B (111 ha), and C
trip, and the more trips they are willing to make. Such
(34 ha). Study sites are labeled n, i, and f for near, intermediate, and far
distance classes. (Reprinted by permission from the National Academy variables can be integrated, at least in part, through a TCM
of Sciences, from Ricketts, T.H., Daily, G.C., Ehrlich, P.R., Michener, demand curve such as that shown in Fig. 11.7. Here the cost
C.D., 2004. Economic value of tropical forest to coffee production. per trip (a reflection of distance, y axis) is related to the
Proceedings of the National Academy of Sciences
number of trips a person makes to the site that contains the
101, 12,579–12,582. https://doi.org/10.1073/pnas.0405147101. #
2004 National Acdemyof Sciences) amenity or resource (x axis). Note that as the cost of the trip
rises, the number of trips decreases. Another variant of the
allowed the researches to have both hand-pollinated and TCM treats the decision to visit the site or not as a function of
naturally (“ambient”) pollinated treatments. The authors the observable characteristics of the site; it can be effective if
found, by keeping all other variables constant, that it is used in a “before” and “after” approach to a site, such as a
11.3 Methods for Valuing Ecosystem Goods and Services 459
maximum product of x (number of visitor days) and

y (entrance fee). For example, if the managers charge a
$10 daily fee at Manuel Antonio Park, they can expect
4000 visitor days, or a revenue of $40,000. If, on the other
hand, they charge an entrance fee of $22.50, use days are
cut in half (2000 visitor days) but revenue increases to
$45,000.
Objective Two: Minimize financial cost to the parks. Perhaps
the National Park Service has a limited budget and insists
that parks be self-supporting through the collection of
daily fees. If we assume that costs increase with increasing
numbers of visitors, then it should be possible to deter-
mine a “supply curve” that relates visitor days to park
Fig. 11.7 A travel cost method (TCM) demand curve that estimates the
costs. Where park costs per day intersect the demand
value of an environmental amenity, such as a national park. Unlike a
neoclassical supply and demand curve, the TCM demand curve makes curve of the daily entrance fee, supply equals demand. If
travel costs (y axis) serve as the analog of price, and number of trips the entrance fee is set below this point, visitor days
(x axis) the analog of demand. Willingness to pay for the environmental increase as costs increase, but revenue decreases, creating
amenity or service is represented by the area ABCD and its value is the
a deficit. If the fee is set at a higher level, revenues may
product of travel costs and number of trips. In this example, if it costs
US$30 for a trip to the park, three trips will be taken, so a value of exceed cost, but the park is “underused” in terms of
US$90 represents the value of the park to the individual. (Graph services that could be provided. Of course, park costs
designed by M. J. Bigelow and F. Van Dyke) may be a constant; that is, perhaps it costs just as much
to manage the park whether any visitors come or not. If
comparison of visitation before and after an environmental this is the case, then the “supply” curve is simply a
improvement (e.g., creating a lake for fishing). horizontal line. But it is still valuable to determine the
As discussed earlier, economics uses supply and demand point at which it intersects the demand curve of the
curves to relate price to quantity demanded; in a TCM curve, entrance fee-visitor day relationship because that point
travel costs are the analog of price, and number of trips the represents the minimum entrance fee that must be charged
analog of demand. If a person is willing to make six trips, to recover costs of operation.
each costing $20, then the person’s willingness to pay is 6 x Objective Three: Minimize environmental cost to the parks.
$20 or $120. This willingness to pay is also referred to as Without considerable constraints, increased visitation
consumer surplus. Aggregate consumer surplus is calculated almost certainly will increase environmental degradation
as the area under the TCM demand curve (Loomis 2000). If to a park. If the Park Service can determine a maximum
we want to compare two different areas, and we suspect that acceptable threshold of such damage, beyond which fur-
one area is of higher environmental quality, we could calcu- ther damage would degrade or destroy the park’s value
late the TCM demand curves for both areas. If our hypothesis and purpose, they can theoretically determine the thresh-
is correct, the TCM demand curve from the better environ- old number of visitor days allowable and set revenues to
mental area should lie to the right of the other. create this level of demand. Using this strategy, price is
In addition to its usefulness in estimating values of con- actually used as a tool to control or limit demand, and
sumer surplus, values of environmental improvements, and through such control to limit degradation to the environ-
relative values of different areas, TCM also can be useful to ment. By limiting degradation, park managers may
managers in setting prices for entry fees. The Costa Rican enhance persistence and diversity of populations that can
National Park Service learned, through TCM analysis of continue to reside within park boundaries and decrease
three of its parks, that a common fee was not the most operational costs of park management and maintenance.
economically efficient method of raising revenue because Objective Four: Set an entrance fee that is appropriate to the
tourist demand differed among parks (Chase et al. 1998). actual incomes of most native Costa Ricans to ensure that
Not only did the demand curves show different y intercepts nationals are not “priced out” of their own parks by
and slopes, but they also had different shapes. The Park wealthier foreign tourists. Here the demand curve can be
Service could potentially use such data in four alternative used to address the issue of access and social equity. An
ways, depending upon their objective: entrance fee of $17 per day may increase revenues com-
pared with one of $5, and European or North American
Objective One: Maximize revenue generated from entrance tourists may be willing to pay it. Average-income Costa
fees and estimate total maximum revenue from the three Ricans, however, may not be able to visit their own park!
parks. This objective can be achieved by determining the If the Park Service considers that such a condition
represents an injustice to its own citizens, it would choose different alternatives (for example, how much would you
a fee appropriate to average national income. Using its pay to preserve an endangered bird, snake, plant, butterfly,
demand curve, it would then be able to predict expected or beetle) that allows the person to assign different valuations
visitor days and make its management plans accordingly. to different kinds or categories of species. Such contingent
In fact, many countries do take such income discrepancies valuations are then used to determine preferences.
into account, and charge a lower entrance fee for their own The WTP approach also can be used to ask more directly
citizens than for foreign visitors. how much the person would pay to keep the habitat itself in
an undeveloped state, how much to have it turned into a golf
Often humans do not know how much value they derive course, a housing development, a water treatment plant, or a
from ecosystem services until they deplete them. A more window factory. To give the question a greater sense of
proactive approach to conservation might include minimizing legitimacy and plausibility, the question may be phrased in
environmental costs to the park, as described in Objective the form of a potential bond issue, such as a state tax to
Three, as well as calculating the economic replacement cost preserve the habitat of an endangered species. In this form,
of similar goods and services. This approach could help the researcher can determine: (1) how many people would
determine the cost of restoring an ecosystem or artificially vote for the bond issue at a given level of taxation, (2) how
replacing lost ecosystem services. Another similar method many people would be unwilling to pay anything at all,
might be to assess the damage costs avoided, in which case (3) the average cost valuation of those against the bond issue
the physical relationship between a change in environmental but willing to pay some lesser amount, and (4) the maximum
quantity and quality and some physical measure of damage is taxation rate of the bond issue likely to pass in an open
quantified, and then per-unit values of the damage are used to election (Hunter 1996). Results could be used to infer the
calculate the cost that would be avoided by the environmental value of a non-economic good (an endangered species) or
conditions or change (Fisher et al. 2015). state (an undisturbed habitat). The proposed mechanism of
payment (taxes, government bonds, recreation fees, or direct
cash contribution) can have a significant effect on the answer.
11.3.3 Stated Preference Methods Some conservationists support WTP approaches because
they permit normally non-economic goods, such as
11.3.3.1 Contingent Valuation endangered species, to stand on equal footing with hydro-
Various techniques of contingent valuation (CV), can be electric dams, power plants, or subdivisions. Even if the
used to create “shadow markets” for nonmarket goods average US citizen would pay only two cents to know that
associated with ecosystem services by attempting to deter- the Hungerford’s crawling water beetle (Brychius
mine consumer and user preferences. The most effective use hungerfordi) is alive and well, the valuation of this sentiment
of these methods is when the question is relatively straight in the US population would run into millions of dollars. WTP
forward, and the responses are easy to analyze relative to allows economists to determine the value of non-economic
simple regression variables such as payment amount, goods, such as endangered species, that can lead to
respondent’s attitudes and socio-economic characteristics assessments of high monetary valuation, permitting
such as income, age, education, etc. (WERF 2008). conservationists to argue that the existence value of a species
The Willingness to Pay (WTP) approach is one form of is worth more than the gains to be realized from its
CV that attempts to assign monetary worth to a non-use value destruction.
(Chap. 10), such as a species existence value, by asking, An alternative but related CV approach is the Willingness
usually through surveys, what a person would be willing to to Accept [Compensation] (WTA) method. Unlike WTP,
pay in exchange for the preservation of a given entity, such as which attempts to determine what a respondent would pay
a rare species, under specific circumstances. To an econo- for an environmental amenity, WTA attempts to determine
mist, “benefits” associated with a resource are those things what the respondent would accept as compensation for losses
that give it value, and something has value if someone is suffered as a result of gaining or maintaining such an ame-
willing to pay for it, no matter what their reasons. To assess nity. Like its counterpart, WTA typically uses survey
benefits of this kind, a typical WTP survey item might pro- methods to determine the average payment affected
pose, “Suppose an undeveloped tract of tallgrass prairie at the individuals would accept for losses they incur as a result of
edge of your town was found to contain a population of a rare conservation practices. WTA can be an effective and neces-
species of butterfly. How much would you be willing to pay sary method in cases where the achievement of a conserva-
to keep this area from being developed and preserve the tion goal or satisfaction of an environmental amenity come
butterfly population?” The WTP approach is often made with a definite and tangible cost to local residents. In fact,
more sophisticated by giving the respondent a range of determining an acceptable and just level of compensation is
11.4 The Role of Moderating Institutions 461
Fig. 11.8 Compensation

payments for three types of
domestic or farm animals injured
or killed in verified wolf attacks in
WI from 1985 to 2006. Deer
(Odocoileus virginianus) were
farmed in forested enclosures.
Springer Nature, from Treves, A.,
Jurewicz, R.L., Naughton-Treves,
L., Wilcove, D.S., 2009. The price
of tolerance: wolf damage
payments after recovery.
Biodiversity and Conservation 18,
4003–4021. # 2009 Springer
Science+Business Media B.V.)
often the only way to break otherwise irresolvable value attributes, the levels of which may vary from one alternative
conflicts that may arise in conservation efforts at regional or to another (WERF 2008). The survey respondent is then
landscape scales. asked to choose among alternatives, indicating the trade-
The re-introduction of the gray wolf (Canis lupus) in offs the respondent is willing to make given an estimated
Yellowstone National Park in the 1990s (more in Chap. 12) monetary value for each alternative (Fig. 11.9). Thus, choice
could not have proceeded without a compensation program experiments can also be designed to determine a person or
established by Defenders of Wildlife, the Bailey Wildlife group’s WTP and WTA. This approach is particularly well
Foundation Wolf Compensation Trust, to reimburse ranchers suited to situations where there may be numerous competing
outside the park for livestock losses caused by wolves options or priorities for stakeholder consideration, particu-
(Treves et al. 2009, Fig. 11.8). Likewise, fair compensation, larly those cases where stakeholders may be asked to inform
determined by WTA, is increasingly viewed as essential in management of ecosystem services which differ in the type
establishing national parks in developing nations where many and nature of benefits they provided to such stakeholders. For
people obtain a living from natural resources through hunt- example, one of the attributes within a choice set could be the
ing, gathering, and pastoral agriculture. For example, the cost that respondents would have to pay for an enhancement
establishment of Andasibe-Mantadia National Park in in the state of the ecosystem in question or the compensation
Madagascar, an area with one of the world’s highest densities people would receive for the deterioration from that state
of endemic species, could not have succeeded without a (Lienhoop and Schröter-Schlaack 2018).
program to compensate local residents for losses associated
with changes in land use in and around the park
(Shyamsundar and Kramer 1996). As the demand for forest 11.4 The Role of Moderating Institutions
products increases among the rural poor, Madagascar is faced
with additional challenges in appropriately compensating 11.4.1 Institutions from Economic Perspective
individuals such that the economic motivation to deforest
the park itself is mitigated (Neudert et al. 2017). As noted earlier, markets fail due to incomplete information,
externalities, and the existence of public goods (Pindyck and
11.3.3.2 Choice Experiments Rubinfeld 2017). Market failure can result in serious envi-
Choice modeling, sometimes referred to as “choice ronmental consequences. For example, the market invisibility
experiments,” is another kind of stated preference method. of many ecosystem services has spurred the steady loss of
The questionnaires used in such experiments are similar to forests, soils, wetlands, and coral reef around the globe
those in CV, but the form of questions is different in that (Russi et al. 2013). The methods discussed in Sect. 11.3
respondents are presented with a series of choice sets. The represent efforts to assign monetary prices to ecosystem
choice alternatives are defined using a common set of goods and services in order to incorporate this value directly
Fig. 11.9 An example of a choice set presented to participants in a study (availability of resources 10 years into the future, measured as mussels
focused on the potential implementation of an Ecuadorian governmental collected per day per user), and monetary incentive (the individual pay-
program designed to preserve coastal mangrove forests known as Socio- ment in case of accepting the agreement, measured in US $ per user per
Manglar (“Mangrove Partners”). Because coastal mangrove forests are year). (Reprinted by permission from Elsevier, from Maldonado, J.H.,
state owned, the choice set focuses on different sets of concessions users Moreno-Sanchez, R., Henao-Henao, J.P., Bruner, A., 2019. Does exclu-
might make. The attributes are obligations (time invested in care and sion matter in conservation agreements? A case of mangrove users in the
management, measured in days per week), exclusion (range of access Ecuadorian coast using participatory choice experiments. World Devel-
and use of mangrove areas by third parties), future resource availability opment 123, 104619. # 2019 Elsevier Ltd)
into decision-making, particularly through economic cost- costs of enforcement and tend to create more favorable
benefit analysis. In economics, institutions play a role in relations between government regulators and private enter-
correcting market failure by offering a set of rules (formal prise. The disadvantage of voluntary programs is that they
or informal) to govern behavior (Fisher et al. 2015). may require heavier tax burdens to operate and thus result in
Institutions can take many different forms in the effort to what economists call deadweight social losses or distortions
better manage our natural world and are developed and in economic efficiency and market function caused by the
enforced by a diverse set of actors. diversion of earned income to the government through taxes.
One prominent economic actor is the government, which For example, increased taxes on income usually lead to
can provide environmental protection and promote conserva- reduced demand for consumer goods and reduced production
tion through creation of policy. One way to think of policy is of such goods. In this case, the reduction in demand is not
as “a definite course or method of action selected from among real, but is rather a “deadweight loss” imposed on economic
alternatives and in light of given conditions to guide and activity by the government through taxation. Nevertheless,
determine present and future decisions” (Merriam-Webster voluntary programs are more economically efficient than
2003). Although a variety of actors can create policy, specific mandatory ones if the tax revenues needed to support the
actions of government are considered examples of public voluntary program are low and the costs of government
policy, or actions taken on behalf of the public in service of services relative to the private cost of the same services are
a specific goal. Public policy generally takes one of three also low, or if the voluntary program costs less than the
forms: regulation, voluntary programs, and market-based mandatory program, or both (Wu and Babcock 1999). In
approaches. To economists, these three forms could also be general, voluntary programs are most cost-efficient when
considered different types of institutions. the number of individuals or businesses involved is large
Regulation has historical precedent. Most of the early relative to the total population, and government services
environmental legislation in the US, and later in other provide nonrival public goods such as information or techni-
countries, initially took the form of mandatory regulations cal assistance.
imposed by government agencies. As will be discussed in Market-based approaches to environmental policy use
Chap. 12, the Clean Water Act, Clean Air Act, and the markets, price and other economic variables to incentivize
Endangered Species Act all derive their authority from the actors to reduce negative environmental externalities or
ability to regulate individual and corporate behavior enhance their contribution to social welfare. Market-based
according to objective, pre-set standards, and to impose strategies are designed to engage the private sector, as these
fines or imprisonment for failure to comply. These acts, inducements (monetary or near-monetary) encourage firms
however, owe more of their success to their requirements (e.g., companies or industries) to reduce environmental harm
for information than coercive force. All require full disclo- to the point that it is still financially valuable for them to do
sure of facts regarding individuals or businesses engaged in so. Often an industry’s cost-savings also translates to cost-
activities covered by legislation, so that the public, the press, savings for customers (EPA 2019). Although market-based
the courts, and legislatures can evaluate them. This public approaches might be least costly in terms of abatement and
access to information that has done more to create compli- enforcement, they do not always address equity concerns
ance with environmental laws than the threat of penalty surrounding environmental issues. As a result, market-based
(Quinn and Quinn 2000). approaches can produce “pollution hotspots,” in which pol-
The impetus of regulation has also spurred innovation in lution becomes concentrated in economically-disadvantaged
pollution reduction technologies, effectively creating a “mar- areas. Further, although regulation has the advantage of
ket” for pollution abatement devices where none previously setting firm limits on environmental damage, market
existed. Regulations imposed on the US auto industry by the incentives do not always achieve these same results as
Clean Air Act led to rapid technological developments in quickly (or any results at all) if the incentive is not propor-
automobile technology, such as catalytic converters, tional to the desired outcome (EPA 2019).
unleaded gasoline, more fuel-efficient engines, hybrid
autos, and, eventually, electric and hydrogen-powered cars
(Gerard and Lave 2005). But coercion has its place. Through 11.4.2 Government-Market Interactions
mandatory regulations, government can require individuals
or businesses to meet environmental standards they would 11.4.2.1 Market-Based Mechanisms –Taxes
otherwise disregard as uneconomical. Taxation imposed on undesirable activities follows the
Governments may also adopt and promote regulation “polluter pays principle” and can be an effective mechanism
through voluntary programs that provide information, tech- for correcting market failure. When implemented, economic
nical assistance, and cash and material subsidies to encourage strategies following the “polluter pays principle” force
compliance. Voluntary programs reduce or eliminate the polluters (rather than society) to pay for the pollution they
create and prevent them from externalizing pollution costs. climate change mitigation activities and further incentivize
Taxation is especially appropriate and effective in controlling the production of green technologies and energy (Information
pollutants that are widely dispersed, because the government Box 11.1). Although such a tax targets industry, policies
is the actor best equipped to address the large areas and large should be designed to ensure the potential financial impact
numbers of people affected. If the government taxes a power on consumers (with industry passing down the tax to
plant for hydrogen sulfide emissions produced in generating consumers) does not exacerbate existing economic
electricity on a per unit basis, it accomplishes two things at inequalities.
once. First, the government is reimbursing itself for the social
costs of air pollution, including increased costs of healthcare
Information Box 11.1: Putting a Price on Carbon
(due to respiratory diseases caused or aggravated by the
Pollution
pollution), increased costs of property damage to the
Carbon dioxide emissions represent perhaps the single
government’s own public buildings, structures, and lands
greatest threat to global climate stability. A tax on
(from the effects of acid rain) and increased costs of preserv-
carbon emissions would internalize the cost of such
ing species, habitats, and ecosystems that may decline due to
emissions, which are currently external to the produc-
pollution. Second, the government creates an incentive for
tion process, but quite real in the form of increased
the pollution producer to reduce pollution on its own, since
worldwide temperatures and their unprofitable effects
every unit reduction in pollution lowers cost and increases the
on crop yields, increased incidence of violent and
margin of profit. Two indirect benefits often result from the
unstable weather events, rising sea levels, and
second effect -- pollution control becomes part of the intrinsic
increased risk to biodiversity worldwide (Malcolm
“organizational culture,” and, if the pollutants are material in
et al. 2006). As industries seek to lower production
nature and removed by the producer prior to emission, taxa-
costs, one of their first priorities, under a carbon tax,
tion may open up new markets for the pollutants to be used in
would be to invest in technologies that reduce carbon
beneficial ways. For example, “scrubbers” in smokestacks
emissions and increase efficiency of energy use,
remove pollutants created by coal burning and thereby accu-
slowing the negative effects associated with global
mulate “sludge.” Although harmful as an air pollutant, the
climate change. Such a tax would make reduction in
sludge is high in sulfur and can, if appropriately applied, be
carbon emissions part of “corporate culture” by making
used as a fertilizer to supply an important plant nutrient for
the reward for reduced emissions intrinsic to a
crops. The Tennessee Valley Authority, one of the nation’s
company’s profit and loss, or to a nation’s GNP.
largest producers of electrical power, now makes from
As early as the 1990s, economists suggested that a
US$6–10 million annually by selling the sludge gathered
tax on carbon would be one of the most aggressive
from its scrubbers. And the Indianapolis Power and Light
options in the fight to mitigate and curb greenhouse gas
Company is planning to adjust its operating conditions to
emissions (i.e., Pearce 1991). Because the tax would be
produce higher-quality sludge (Hoffman 2000).
revenue generating, governments could consider a
Taking a more comprehensive approach, governments
range of policy options for redistributing such revenue,
could create a tax structure based on environmentally detri-
potentially adopting a fiscally neutral stance and using
mental activities. The more an individual or business
revenues to finance reductions in incentive. In 2008,
participated in these activities or purchased products
British Columbia, Canada introduced a revenue-neutral
associated with them, the greater their tax burden. For
carbon tax covering 70% of all provincial emissions
individuals, such taxes might take the form of higher sales
(UNFCCC 2019). All revenue generated is subse-
taxes on less fuel-efficient cars, taxes on non-recyclable or
quently returned in the form of personal and business
non-biodegradable products, or taxes on home heating or
tax measures, such as reductions in personal income
cooling practices that produced high levels of pollution. An
tax rates and corporate income tax reductions. One
example of this in the area of public health is a tax on sugary
ancillary result of such a tax has been steady growth
drinks, where a one cent per ounce tax on soda is estimated to
in the clean tech sector -- “the ‘West Coast Clean
decrease consumption in the US by more than 10% (TPC
Economy: 2010–2014 Jobs Update’ notes that British
2019).
Columbia has 68,165 clean economy jobs, a 12.5%
For business and industry, there might be an expansion of
increase since 2010. The Update also found that B.
existing taxes and fees on pollution and waste on a per unit
C.’s clean economy GDP rose to $C 6.31 billion by
basis. As long as the marginal cost of reducing emissions is
2014, a 19.3% increase from 2010” (UNFCCC 2019,
less than the tax, this tax gives a profit-maximizing company
Information Box Fig. 11.1).
an incentive to figure out ways to reduce its emissions. Most
recently, governments around the world have proposed and (continued)
implemented a tax on carbon pollution, in order to hasten
water in intensively farmed areas indicate that conventional

Information Box 11.1 (continued) corn and soybean farming is not environmentally sustainable
(USC 2017). Loans and subsidies absorb farmers’ losses and
encourage such conventional practices to continue, even if
done with the intention of supporting a farmer’ livelihood or
cultural practice.
11.4.2.3 Market-Based Mechanisms –Tradable

Permits
Tradable permits force regulators to identify a maximum
acceptable level of pollution or depletion that is ecologically
sustainable. Once determined, regulators must distribute the
rights to pollute in some fair manner so that the market can
attain efficient allocation of permits through trading. Pollu-
Information Box Fig. 11.1 Changes in aggregate employment tion rights can operate in a free market, but only after ecolog-
for British Columbia based on the implementation of a carbon tax ical and political boundaries are established (Daly 1999).
(solid line). The alternative annual trend in aggregate employ- Allocation of pollution rights is again a manifestation of the
ment without the carbon tax is shown as the solid dotted line,
with the light grey lines representing the confidence interval of
“polluter pays principle” through which government sells
the projection. The vertical solid line indicates the implementa- such rights on a per unit basis rather than taxing each unit
tion year for the carbon tax. (Reprinted by permission from produced. In effect, tradeable permit approaches tacitly assert
Elsevier, from Yamazaki, A., 2017. Jobs and climate policy: that an ecological system, such as the atmosphere, has value
Evidence from British Columbia’s revenue-neutral carbon tax.
Journal of Environmental Economics and Management 83, 197–
through the services it delivers. To degrade environmental
216. # 2017 Elsevier Inc) services through pollution represents a social cost, for which
the polluter must pay. However, as an incentive, pollution
rights are made transferable. If a polluter does not “use up” all
11.4.2.2 Market-Based Mechanisms – Subsidies his pollution rights (because of increased efficiency and
Just as taxes can curb undesirable behavior, governmental cleaner production), he can sell them to another polluter,
subsides act as a reward to incentivize good behavior. offering a cost-effective way to achieve overall reductions.
Subsidies are transfers of payment to either “encourage Cleaner producers gain an economic advantage, and “dirty”
uptake of a policy, program or institution, or aid in polluters must pay more up front or buy more rights to
overcoming any cost burdens a policy, program or institution pollute. Over time, the pollution standard, or “pollution
may introduce” (Fisher et al. 2015). For example, the North cap,” can be tightened to increase pressure on market
American Wetlands Conservation Act (NAWCA) provides participants, increase the value of the permits, and further
funding for the protection, restoration and enhancement of reduce overall emissions (GWP 2017).
waterfowl habitat in the US, Canada, and Mexico. Since Tradeable permits have a growing use in protecting eco-
1989, NAWCA has granted US$1.48 billion and generated system services. For example, the US has a No Net Loss rule
US$4.34 billion from partner contributions to fund habitat for wetlands, designed to ensure that developers invest in a
projects across 33.4 million acres (13.5 million ha) (Ronis level of mitigation that ensures that their projects will result
2018). NAWCA, and its corresponding North American in “no net loss-or net benefit-to ecosystems” (Hayes and
Waterfowl Management Plan, have promoted economic Gentile 2016). Although this is in some sense a regulatory
investment in wetland protection at international scales. policy, it has spurred private investment in land and wildlife
Some governments have programs that provide cash conservation by creating the opportunity for “wetland banks”
payments or other forms of remuneration for activities that that function like market-based tools (Fisher et al. 2015), as
cannot be environmentally sustained in the long term. Price well as making wetland assessment and restoration a major
supports in agriculture, for example, require cash payments to “growth industry” among private environmental consultants.
farmers when market prices fall below profitable levels. In Industries that will cause the loss of wetland area can fund the
the United States, major federal loans and subsidies for restoration of another “equivalent” wetland somewhere else
farmers have been approved in every year in which large in the state or region. Wetland banks help facilitate this kind
harvests result in falling prices for corn and soybeans (Urry of credit transfer. Functioning like a cap-and-trade approach,
2015). Although not every farming system has the same level where the net loss is zero and the trade is in wetland credits,
of environmental impact, soil erosion rates, pesticide and the balance of buying, selling, and trading wetland areas is
fertilizer hazards, declines in native populations of plants determined by the market (Fisher et al. 2015). Some
and animals, and degradation of groundwater and surface economists have proposed applying tradable permits to
biodiversity conservation at-large to better reconcile eco- large treble hooks on weighted, heavy lines, heaved into the
nomic development with conservation (Wissel and Wätzold river with surf rods 8–12 feet long.
2010). As land-use change is a key driver of biodiversity loss, Paddlefish roe (eggs) are used as caviar, comparable in
one benefit of such a system would be to help landowners quality to the more famous caviar of sturgeons (Family
clearly calculate the opportunity costs of development. For Acipenseridae). In the southern US, paddlefish populations
example, Wissel and Wätzold suggest that “permit markets in the Mississippi River and its larger tributaries have been
provide an incentive for landowners to use their land in such a over-exploited, resulting in the closure of many state fisheries
way that a cost-effective allocation of land-use types emerges. in this region (Anderson and Leal 1997). Local entrepreneurs
Because habitat can only be destroyed if habitat of equivalent in Glendive, however, recognized that the fishers harvesting
value is restored, the overall ecological value in the designated paddlefish from the Yellowstone, and the nearby Fort Peck
area remains the same” (2010). Of the course, one of the key Reservoir and Lake Sakakawea, had no interest in the roe. In
challenges presented in this system mirrors that of wetland fact, fishers had historically dumped several tons of roe on the
banking – how does one define “identical” or “equivalent” banks of the Yellowstone near Glendive each spring,
habitats? Assessing habitats characteristics across type, space attracting large concentrations of flies and rats, creating a
and time would be a necessary first step. Despite such public nuisance and health hazard. Glendive business
challenges, a tradable permit system informed by expert planners conceived a plan to open a caviar processing plant
knowledge and precise estimation of ecosystem services, and market the caviar internationally. Approval of the project
might lead to creative opportunities for mitigating the impacts rested with state officials of the Montana Department of Fish,
of economic development (Wissel and Wätzold 2010). Wildlife, and Parks. Biologists in the department determined
that local populations of paddlefish appeared stable and
11.4.2.4 Government-Market Coordination – secure. The paddlefish habitat in the river, reservoir, and
Conservation and Paddlefish Caviar lake was protected and not faced with any serious threats of
In eastern Montana (USA), the town of Glendive is home to pollution. The caviar plan did not increase mortality to the
approximately 8000 residents. Its traditional economic base population or decrease recruitment, since the roe would come
was agriculture, particularly cattle ranching. Located along only from fish harvested by private anglers. However, to
the lower reaches of the Yellowstone River, Glendive is also ensure that profit motives did not lead to overharvest, state
famous for its paddlefish (Polyodon spathula, Fig. 11.10), officials stipulated that fishers had to donate their roe, not
which spawn over gravel bars in swifter sections of this part receive payment for it. To make this attractive, the Glendive
of the Yellowstone. Paddlefish are highly valued as food, and community agreed to employ individuals who would clean
even one individual can provide a lot of it. Typical adult an angler’s paddlefish without charge. In return for the
paddlefish may be 5–7 feet long (including their paddle-like cleaning service, the cleaner would keep the roe and give it
snout) and weigh 60–120 pounds. Each year, the paddlefish to the caviar factory. Finally, the state stipulated that, in
season (~1 May–15 June) attracts about 3000 anglers to addition to a regular fishing licenses, paddlefish anglers
Glendive. Because paddlefish feed mostly on plankton and would be required to purchase special paddlefish tags that
other microscopic organisms, they cannot be caught with must be affixed to each fish caught (Anderson and Leal
conventional bait and lures, but must be snagged, using 1997), but the number of tags that could be sold was fixed.
In this way, state officials could control the maximum num-
ber of paddlefish harvested in a season. Finally, the Glendive
business community agreed to devote half of the net revenues
from paddlefish caviar to the state of Montana for paddlefish
research and management (Anderson and Leal 1997). In its
first seven years of operation, the project grossed over one
million dollars in revenue.
There are dangers in tying conservation to market-driven
demand, but the Glendive-paddlefish example highlights six
elements of program design that can produce effective inter-
action between governmental authority and free market
systems.
Fig. 11.10 Paddlefish (Polyodon spathula) roe provides the basis for a
growing caviar industry in Glendive, Montana. Paddlefish can live up to
55 years (though average lifespan is 20–30), growing to be over seven
Element One: The resource was derived from stable
feet long and up to 200 pounds. They are easy to identify, with long, flat populations whose habitats were protected and managed
blade-like extended upper jaws that are almost one-third of their entire for the population’s benefit. The state, not the market,
body length. (Photo courtesy of USFWS, public domain) made biological determinations of the status of the
population and its habitat. The sustainability of both were were directed to improve the productivity of the resource,
mandated as given conditions required a priori by state thus enhancing its sustainability.
regulatory authority as prerequisites for any use of the
paddlefish as a human resource. Not all forms of resource use lead to resource depletion or
Element Two: Persistence of the resource in perpetuity was habitat degradation. When this is the case, conservationists
mandated, regardless of market comparisons of present can work with the private sector to create markets for
versus future values or considerations of opportunity resources that assign specific economic values to benefits
costs. The market was not consulted about whether pad- derived from such resources. The lesson from Glendive is
dlefish habitat could be used more profitably for other straightforward when coupled with lessons from conserva-
purposes, or whether the current value of paddlefish at tion history. The market is a bad master for conservation
higher rates of exploitation had more value than the future values, but can be a useful servant to achieve conservation
or option values of paddlefish populations in future outcomes, if its mechanisms are channeled to achieve profit
generations. The persistence of paddlefish in Montana according to predetermined, community-based conservation
was non-negotiable, set above the reach of the market by values enforced by government regulation at levels deter-
the state’s statutory authority. mined by scientific assessment. Administered wisely, the
Element Three: The rate of exploitation was determined by coupling of conservation values and market incentives can
biological criteria, not economic criteria. Optimal exploi- produce and efficiently distribute benefits that build broad-
tation rates of paddlefish were not determined according to based, community-level support for conservation more effi-
the criteria of profit maximization, supply and demand, or ciently and effectively that legislative mandates alone. Such
human welfare. The exploitation rate was determined by efforts move the concept of sustainability beyond the aca-
the biological productivity of the local paddlefish popula- demic and professional culture of conservation biology into
tion as determined by biologists, and exploitation was set the world of private business and economics. In this case, the
at a rate below maximum sustainable yield in order to citizens of Glendive came to understand that a healthy, stable
ensure continuance of a stable harvest. population of paddlefish was an index of their community’s
Element Four: The harvest was administered by the state in a well-being. Decline in the population would have been
manner that ensured that maximum sustainable yield was viewed by them as a symptom of distress.
not exceeded, and in a way that removed profit incentives
to violate yield restrictions. By requiring every paddlefish
to be tagged and having game wardens arrest violators, the 11.4.3 The Role of Property Rights
state enforced the conditions described above without in Conservation
regard to market values. By requiring roe to be donated
rather than sold, the state removed profit incentives to 11.4.3.1 Intersections Between Property Rights
harvest more paddlefish than allowed by law. and Economics
Element Five: Regulated private enterprise was allowed to Some economists would argue that all conflicts between eco-
create a market for a natural resource, to allocate the nomics and conservation reflect a failure to clearly define and
distribution of the resource according to supply and properly allocate property rights. If property rights are clearly
demand, and to permit the private sector to receive an defined and enforced, formerly non-rival, non-excludable
economic incentive. With restrictions that prevented the goods and services (public goods), or rival and
market from determining the amount of the resource that non-excludable goods and services (common pool resources),
could be harvested, the market was permitted to determine including those provided by ecosystems and biodiversity, can
the value of the resource that was harvested. A portion of be made rival and excludable, subject to market mechanisms
the profits went to those who marketed the resource, for valuation and allocation. Property rights, therefore, are
giving them incentive to continue, and placing an eco- another example of an institution, which mediates linkages
nomic value on the resource itself. between human societies and ecosystem services (Schlager
Element Six: Profits were tied to resource sustainability by and Ostrom 1992). These rights establish rules that govern the
mandating that a portion of the profits be re-invested in range of activities that individuals or groups can undertake
the productivity of the resource, not in expanding the with regards to a specific or a group of services. As outlined by
production or harvest capacities of the market. Histori- Schlager and Ostrom (1992), common examples of such
cally, profits from commodity resources have been used to natural resource rights might include:
increase the capacity of exploiters to take more resources
at faster rates. This pattern increases short-term profits but • Defining Access (right to enter a defined physical area and
destroys long-term sustainability. In Glendive, profits enjoy non-subtractive benefits)
• Withdrawal (right to obtain resource units or products of spring rains, and a heavy demand for water for crop irrigation
resource systems) reduced the Ruby River in Montana (USA) to historically
• Management (right to regulate internal use patterns and low levels, with many stretches characterized by isolated
transform resources by making improvements) pools with dry riverbed between. Hundreds of trout became
• Exclusion (right to determine who will have access right stranded in such isolated and (subsequently) overheated
and how that right may be transferred), and pools where they could not survive. However, the diverted
• Alienation (right to sell or lease exclusion, management or water turned out to be of low value to private landowners
withdrawal rights). who had used it for crop irrigation and, because they diverted
too much, ended up with excess water standing in their fields.
Over the past few decades, there has been an “explosion” of A US conservation NGO, Trout Unlimited, wanted to tem-
work on common property arrangements and common-pool porarily purchase (“lease”) water rights from the farmers and
resources to identify regimes that “allocate benefits equitably, divert it back into the river to save the trout. Trout Unlimited
over long time periods and with only limited efficiency losses” had the money to do so, as the amount of water needed would
(Agrawal 2001). Although diverse, many of these approaches have cost only US$4000. Unfortunately, state and federal law
build on neoclassical economic theory while recognizing that prohibited transfer of water rights for “nonbeneficial” uses,
complex environmental and conservation challenges do not traditionally interpreted to mean any use that did not benefit
reflect a dichotomous world in which the only actors of impor- humans. Because saving the trout was not a “beneficial use”
tance are “the market” and “the state” (Ostrom 2010). For and water rights were not tradable, water remained standing
example, citizens, local public entrepreneurs, and public officials in crop fields while thousands of trout died (Anderson and
have been able to engage in new ways, to establish creative and Leal 1991).
pol-centric (or decentralized) management partnerships. Some economists believe that making property rights
Drawing directly on Hardin’s Tragedy of the Commons tradable, especially rights associated with ecosystem
motif, economist Bruce Yandle describes a non-tragic ending services, could avert tragedies like Ruby River. But
to Hardin’s story of the herders (whom Yandle calls preventing such calamities is not merely a matter of
“shepherds,” making their livestock “sheep”) and their com- “tradability.” In a recent review of common design principles
mon pasture. Rather than simply let the pasture deteriorate that affect the probability of long term survival of a given
through unregulated grazing, the shepherds form a property rights system, Michael Cox and colleagues (2010)
“shepherd’s club” in which they share information and act offer the following list of best practices:
cooperatively (Yandle 1997). Like all good economists,
Yandle supports his thesis with a chart (Fig. 11.11). Sheep 1A. User Boundaries: Clear and locally understood boundaries
enter the pasture as long as average weight gain is at least as between legitimate users and nonusers are present.
good as the next best opportunity (any alternative pasture). 1B. Resource Boundaries: Boundaries that separate a specific
Herd expansion stops when the marginal gain is negative. common-pool resource from a larger social-ecological
Using this relationship, Yandle notes that “the club would system are defined.
allow sheep in the pasture until opportunity cost was equal to 2A. Congruence with Local Conditions: Appropriation and
the marginal product of the pasture, read at OA on the provision rules match local social and environmental
horizontal axis of the figure. Operating at that point, the conditions.
shepherd club would produce the largest amount of weight 2B. Appropriation and Provision: Appropriation rules match
gain possible: each member could conceivably be wealthier provision rules, so that distribution of costs is proportional
than before, depending on the rule for output sharing” to distribution of benefits.
(Yandle 1997:15). The shepherd’s club provides needed 3. Collective Choice Arrangements: Most individuals
safeguards against overuse of the common-access pasture affected by a resource regime are authorized to participate
through shared information. However, the shepherd’s club in making and modifying its rules.
does not solve the fundamental problem of property rights 4A. Monitoring Users: Users or individuals accountable to
that pertain to the use of the pasture. users monitor appropriation and provision levels of the
In one sense, property rights must be transferable or users.
tradable, just as environmental pollution rights were made 4B. Monitoring the Resource: Users or individuals account-
tradable in our earlier example of cap-and-trade policy. To able to users monitor the condition of the resource.
understand how such tradable rights could serve conservation 5. Graduated Sanctions: Sanctions for rule violations start
ends, consider an historical example. In 1987, the combina- very low but become stronger if a user repeatedly violates
tion of a mild winter with little snowfall, lower than normal a rule.
Fig. 11.11 Hardin’s (1968) tragedy of the commons expressed as a was equal to the marginal product of the pasture, read at OA on the
problem in marginal gains. The number of sheep entering the pasture horizontal axis. Operating at that point, the shepherds would produce the
(X axis) should be increased as long as average weight gain (Y axis) is at largest amount of weight gain possible. (Reprinted by permission from
least as good as the next best opportunity (any alternative pasture, Rowman and Littlefield Publishing Group, from Yandle, Bruce., 1997.
represented by the line labeled “opportunity cost”). Herd expansion Common sense and common law for the environment: creating wealth
stops when the marginal gain becomes negative. Logically, a commu- in hummingbird economies, Political economy forum. Rowman &
nity of shepherds would allow sheep in the pasture until opportunity cost Littlefield, Lanham, Md. # 1997 Rowman and Littlefield)
6. Conflict Resolution Mechanisms: Rapid, low cost, local council, has determined that those sorts of enterprises must be
arenas exist for resolving conflicts among users or with placed in zones designated as “commercial” or “industrial.”
officials. In effect, zoning is a form of land use regulation which asserts
7. Recognition of Rights: Rights of local users to make their that the public can limit future development to protect public
own rules are recognized by the government. interests, and that compensation is not necessarily provided. I
8. Nested Enterprises: When a common-pool resource is could petition the City Council for an exception to the zoning
closely connected to a larger social-ecological system, restriction if I can show that such an exemption would be
governance activities are organized in multiple nested warranted. But my petition will be made public and debated
layers to support sustainable resource use. in an open hearing that any of my fellow residents might
attend. The decision will be made by community delibera-
Advocates of property rights solutions to biodiversity tion, not market forces or prices. If my request is denied, I
conservation often assume that the best results come from have no basis for asking the city or my neighbors for potential
greater levels of private ownership and higher allocations of lost income that I could have received from making windows
property rights to private individuals, but these are not the in my factory or from guests using my hotel.
only property rights-based solutions available. Solutions to Zoning represents the “police power” of the public to
common pool resource dilemmas are diverse, including gov- regulate “external” costs associated with individual decisions
ernment management and community management through of private landowners. The zoning ordinance in this example
collective action, among others (Heikkila et al. 2011). is a preemptive strike by the community that prevents some-
one from placing undue levels of noise, traffic congestion, or
11.4.3.2 Transfer of Private Property Rights: potential environmental hazards in the proximity of where
Zoning and Conservation Easements people live. Thus, zoning can be a critical tool for private land
Zoning is an arrangement in which some of the property conservation. If a community zones an area around the town
rights normally associated with the individual owner of a as “open space” or “native prairie,” they are specifying a land
property are transferred to or held by the community in use that no private landowner can violate. Thus, zoning
which the property is located, effectively restricting what regulations can act as an incentive for conservation by limit-
individual property owners can do on their property based ing the size and intensity of development, or even banning it
on where the property is located. For example, if my home in altogether.
my town is in a section that has been zoned “Residential,” I Zoning has implications for conservation because zoning
am not at liberty to tear down my home and build a five-star regulations can reduce the cost of purchasing development
hotel or a glass factory, even if I have sufficient land and rights. If a potential land buyer is a conservation NGO such
money to do so. The community, more specifically the city as The Nature Conservancy and knows that the land it wants
is in an area zoned as “open space,” they know in advance crucial role as buffers for land dedicated to forestry, farming
that the owner has no basis to raise her price to what the land and ranching, even if not utilized for core habitat protection.
would be worth if converted to an industrial park or a new A recent survey of all easements purchased by The Nature
housing development, and no hope of getting such a price Conservancy between 1985 and 2004 showcased a wide
from any buyer. Thus, zoning designed for conservation can range of named purposes for such easements as well as a
lower the “ceiling” of expected costs for land acquisition and range of biodiversity threats they attempted to abate
use, and gives individuals or organizations intending to use (Table 11.3 and Fig. 11.12). Nearly all sampled easements
the land for conservation purposes a significant advantage as were intended to reduce habitat loss and fragmentation, a
buyers. Conservation programs can take advantage of the central threat to biodiversity (Rissman et al. 2007). However,
restrictions that zoning imposes because the developable the study also showed that 85% of sampled easements
land value is lower than it would be without regulation. allowed for a residence, new permitted structures, commer-
Conservation easements are a special case of land-use cial use or some subdivision of the property. As reiterated by
zoning that have been developed to make the value of con- author Adena Rissman and colleagues, “early easements
servation on private land more explicit and profitable to provided land trusts with an important lesson: even without
landowners. In an easement, the landowner agrees, usually additional development, subdivision of the property can cre-
with a government entity or a private conservation organiza- ate problems because it requires a land trust to maintain
tion, to restrict some activities or forms of development on relationships with multiple landowners and may fragment
his or her land to achieve specific conservation goals, such as land management” (Rissman et al. 2007). Therefore, in addi-
habitat or species protection. Such restrictions lower the tion to reassessing restrictions around property subdivision,
assessed value of the land, generating a reduction in property there is need for easement monitoring over time. Rissman
taxes for the owner and a reduction in inheritance taxes for et al. emphasize that only 20% of easements in their sample
the owner’s heirs. The owner, however, retains the rights to had quantitative monitoring programs, and that while quali-
possession, residence, non-prohibited activities, and legal tative information suggested the status of conservation targets
title to the land. Conservation easements work because they was steady or improving on nearly all (95%) easements, this
provide incentives for conservation by private citizens on information was subjective and might have been unreliable
their own land, and enable management action that supports due to the lack of monitoring (2009). The tension between
biological and community-based integrity rather than focus- private economic incentives for land-use change and the
ing solely on individual species. Easements can also play a overall conservation goal of the easement is one example of
Table 11.3 The number and purpose of conservation easements (n ¼ 119) identified by The Nature Conservancy staff based on easement
documents and supplemental project documentation. Survey respondents identified 24 types of conservation purposes stated in the easements
themselves or supplied from supplemental documents. Each easement had multiple purposes. Purposes common to 10% or more of the easements
were primarily ecological in nature
Easement Additional project documents
Purpose document only Total
Retain property and habitat undisturbed in a natural condition 119 0 119
Prevent uses that would impair, degrade, or interfere with conservation 94 0 94
values
Protect aquatic, marine, or wetland habitat and communities 47 14 61
Contribute to connectivity of surrounding protected areas 16 11 27
Accommodate educational or scientific activities 21 3 24
Provide public access, services, or scenic enjoyment 21 3 24
Protect habitat for animal migration routes 18 6 24
Prohibit certain further development activities, fragmentation 17 6 23
Protect endangered species 10 13 23
Buffer habitat or natural landscape feature 10 11 21
Provide for restoration activities 13 3 16
Promote compatible or heritage grazing 12 2 14
Protect historic value (i.e., land uses, structures) 5 7 12
Reprinted by permission from Wiley, from Rissman, A.R., Lozier, L., Comendant, T., Kareiva, P., Kiesecker, J.M., Shaw, M.R., Merenlender, A.
M., 2007. Conservation Easements: Biodiversity Protection and Private Use. Conservation Biology 21, 709–718. # 2007 Society for Conservation
Biology
extracted. These restrictions increased the company’s extrac-

tion costs, thereby reducing the royalty that Audubon
received compared to what it could have gotten without
regulations. That, as economists Terry Anderson and Donald
Leal note “is the price they pay for caring for the environ-
ment” (Anderson and Leal 1991:91).
Is Audubon’s action an example of practical and produc-
tive conservation policy, or a disturbing case of compromise
and rationalization counter to their own conservation ideals?
As recently as 2010, Audubon considered reopening drilling
in Rainy citing new technology that has made drilling less
invasive, and careful monitoring that could minimize
damage – all in the name of securing funds for much needed
coastal restoration (DeGregorio 2010). This tension is not
confined to Audubon. Commodities like oil and natural gas
Fig. 11.12 Conservation threats that the 119 sampled conservation have established markets that determine their prices. Such
easements aim to abate, as indicated by survey respondents from The commodities may generate revenues that can be re-invested
Nature Conservancy. Respondents could indicate multiple threats for in conservation, but the commodities are not what NGOs like
each easement. (Reprinted by permission from Wiley, from Rissman, A. Audubon are trying to conserve. Free-market environmental
R., Lozier, L., Comendant, T., Kareiva, P., Kiesecker, J.M., Shaw, M.R.,
Merenlender, A.M., 2007. Conservation Easements: Biodiversity Pro- economists like Anderson and Leal advocate the competitive,
tection and Private Use. Conservation Biology 21, 709–718. # 2007 private auction of lands with high values for wilderness or
Society for Conservation Biology) wildlife conservation to those who are willing to pay the most
for them. By some measures, this would certainly represent
what free-market economists would define as “efficient” allo-
the wider debate on the compatibility of human uses and cation, but it is hard to imagine even the most well-financed
biodiversity in protected areas (Naughton-Treves et al. 2005). conservation NGOs, like The Nature Conservancy, being
able to “outbid” ExxonMobil for a wilderness area or natural
11.4.3.3 Drilling Rights in Protected Areas: The sanctuary that both desired. Such a strategy would circum-
Case History of the Rainey Wildlife vent the role of public, community discussion and delibera-
Sanctuary tion about what might be the “best” use of such lands. But
The Audubon Society, a private conservation NGO, has an Leal and Anderson respond by proposing the creation of
admirable record of biodiversity conservation. As part of its “Conservation Endowment Boards.” Under this arrangement,
conservation effort, the Society purchases and manages lands national conservation boards, with members approved by a
of high conservation value as wildlife sanctuaries. One of country’s national Congress, would each have a narrowly
these is the Rainey Wildlife Sanctuary in Louisiana (USA). defined mission of protecting and enhancing a specific con-
Rainey is home to mammal species such as white-tailed deer, servation value. In their own words, Leal and Anderson
armadillo, muskrat, otter, and mink, as well as hundreds of describe the arrangement as one where “Each board would
species of birds. As an organization, Audubon opposes dril- have a fiduciary [“of or relating to holding something in trust
ling for oil and natural gas in wilderness areas in the US (i.e., for another”] responsibility under common law to carry out a
NAS 2017). However, it has historically taken a different single mission, and it would have the option of allowing
view of its own property rights, including the rights to fossil alternative uses in the area as long as those uses enhance
fuel resources. From the 1940s through 1999, Audubon per- the board’s overall mission.” Radical as this idea may seem,
mitted the development of oil wells within Rainey, even the Rainey Refuge provides an example of this principle in
though such wells were a potential source of pollution and action. In this case, the “endowment board” was the Audubon
other threats to wildlife. Yet, carefully monitored and Society. It held in trust, for its members, a mission to protect
constrained, Audubon argued that oil could be extracted biodiversity, but was willing to allow other activities on its
without measurable damage to the marsh. In return for refuges if they contributed to that mission. Of course, decades
allowing an oil and gas production company, Consolidated later Audubon decided that such activities were not congru-
Oil and Gas, to remove oil and natural gas on the sanctuary, ent with its mission, suggesting that the jury is still out on the
Audubon received royalties, but imposed contractual degree of compromise the organization is willing to accept or
restrictions on the company regarding how the oil could be defend.
subsistence, and only “misery and vice” would keep numbers

Point of Engagement Question in check (Piel 1995). Despite his mathematical logic,
For decades, Audubon made resource decisions on Malthus’ conclusions were generally disregarded because
their own private land that provided economic they appeared to be refuted by experience. Human ingenuity
resources to support their conservation goals. How- and technology have, in fact, shown far more than the arith-
ever, such activities arguably stood in contrast to their metic increase Malthus predicted. World human population is
public-facing fight against fossil fuel extraction in wil- now in its sixth doubling since Malthus published his Essay
derness areas. Was their action at the Rainey Refuge and has been sustained by more than five doublings of “the
justifiable given their broader mission or a case of means of subsistence” (World per capita GNP). Even since
unethical compromise to increase organization 1950, industrial technology (overriding any arithmetic con-
revenue? straint) has twice doubled the output of material goods (Piel
1995). With this record of achievement and growth in both
population and affluence, many neoclassical economists felt
11.5 Ecological Economics they could safely ignore Malthus’ ideas and treat his essay
simply as a historical footnote.
11.5.1 Historical Challenges to Neoclassical Not all economists, however, were prepared to dismiss
Economics Malthus’ views on growth and its limits. One of the most
influential modern economists to address the problem of
In his landmark book, The Environmentalism of the Poor, environmental constraints was Nicolas Georgescu-Roegen.
economist and economic historian Joan Martinez-Alier of the In his classic work, The Entropy Law and the Economic
University of Barcelona (Spain) takes a position regarding Problem, Georgescu-Roegen argued that, “What goes into
economics and the environment very different from free- the economic process represents valuable natural resources
market economists like Anderson and Leal. He writes, “In and what is thrown out of it is valueless waste” (Georgescu-
modern industrialized and industrializing societies there has Roegen 1993:76, emphasis his). “. . .Matter-energy,”
been a strongly argued view that enlarging the economic pie Georgescu-Roegen continued, “enters the economic process
(GNP growth) represents the best way of alleviating eco- in a state of low entropy and comes out if it in a state of high
nomic distribution conflicts between social groups. The envi- entropy.” Entropy is a measure of the amount of unusable
ronment came in, if at all, as an afterthought, as a energy in a system; as entropy increases, the amount of
preoccupation arising out of deeply held values on the sacred- energy available for work decreases. To illustrate,
ness of Nature, or as a luxury (environmental ‘amenities’ Georgescu-Roegen asserted, “. . .a piece of coal can only be
rather than necessities). The poor were ‘too poor to be used once. And, in fact, the entropy law is the reason why an
green.’ They must ‘develop’ to get out of poverty and, as a engine (even a biological organism) ultimately wears out and
by-product, they could then acquire the taste and the means to must be replaced by a new one, which means an additional
improve the environment” (Martinez-Alier 2002:16). tapping of environmental low entropy” (Georgescu-Roegen
Martinez-Alier offers a different view. He argues that conser- 1993:80).
vation is not to be treated as an amenity, but as a necessity, Georgescu-Roegen compared the Earth’s resources to an
especially for the poor. And the affluent of wealthy nations hourglass full of sand (Fig. 11.13), in which the sand in the
will not enhance environmental conservation and biodiver- upper part of the hourglass represented the store of
sity protection by increasing their technological efficiency in low-entropy resources. Its rate of movement into the bottom
waste disposal and fuel consumption, but by increasing their of the hourglass was its “flow,” controlled by rates of solar
frugality through limiting how much they use and consume. inputs. As the amount of low-entropy resources diminished,
Although an important modern marker in the fight for global the amount of high entropy waste (in the bottom of the
environmental justice, Martinez-Alier’s insistence on hourglass) increased and accumulated. Some of the upper
recasting of the relationship between environment and econ- sand coalesced into clumps and might move through the
omy has historical roots. neck of the hourglass all at once, analogous to fossil fuels
In his 1798 Essay on Population, economist Thomas that might accumulate large quantities of solar energy (when
Malthus wrote, “Population, when unchecked, increases in originally formed) and then can be tapped at higher rates of
a geometric ratio. Subsistence increases only in an arithmetic flow. But regardless of the rate of flow, the sand in the upper
ratio. A slight acquaintance with numbers will shew the half is destined to run out. And, unlike a real hourglass, this
immensity of the first power in comparison with the one cannot be turned over! Although humans have almost
second. . . In two centuries and a quarter, the population complete command of the energy stocks, for all practical
would be to the means of subsistence as 512 to 10.” Malthus purposes, they have no control over the flow of solar radia-
predicted that populations would grow beyond their means of tion. To Georgescu-Roegen, the implications of this
11.5 Ecological Economics 473
Let S denote the present stock of terrestrial low entropy and let r
be some average annual amount of depletion. If we abstract
(as we can safely do here) from the slow degradation of S, the
theoretical maximum number of years until the complete exhaus-
tion of that stock is S/r. This is also the number of years until the
industrial phase in the evolution of mankind will forcibly come to
its end.
Given the fantastic disproportion between S and the flow of
solar energy that reaches the globe annually, it is beyond ques-
tion that, even with a very parsimonious use of S, the industrial
phase of man’s evolution will end long before the sun will cease
to shine. (Georgescu-Roegen 1993:84–85).
11.5.2 Characteristics of Ecological Economics
The views of Georgescu-Roegen and others led to new ways

of thinking about interactions between environmental and
Fig. 11.13 The “hourglass analogy” of economist Nicolas Georgescu- economic processes and to the growth of ecological econom-
Roegen illustrates the relationship between entropy and economics.
ics as a distinct economic paradigm and discipline. Unlike
First, the hourglass is an isolated system, no sand enters or exits. The
sand in the upper part of the hourglass represents Earth’s low-entropy traditional growth, or neoclassical economics, ecological
solar resources – solar energy arrives to earth as a flow governed by the economics asserts that human-made capital cannot, in the
constricted middle of the hourglass. As humans consume terrestrial long run, substitute for natural capital in providing raw
stock resources, which ultimately depend on solar energy, high-entropy
materials and energy, stock (nonrenewable) resources, flow
wastes are produced. There is an important asymmetry here between the
two-sources of low entropy – the solar source is stock-abundant, but (renewable) resources, a sink for wastes, and key life support
flow limited, while the terrestrial source is stock-limited, but flow- systems including water, air, climate regulation, food, and
abundant (temporarily). Humans have increasingly become dependent biodiversity. Thus, ecological economics sees the human
on limited terrestrial stocks. (Reprinted by permission from Beacon
economy not as a self-sufficient system that could draw
Press, from Daly, H.E., 1996. Beyond growth: the economics of sustain-
able development. Beacon Press, Boston) material from or dump material into the environment without
restraint, but as an environmentally dependent subsystem of
human activity that would cease to function without environ-
understanding were profound: “There is an important asym-
mental goods and services (Fig. 11.14). In the past, the sense
metry,” he noted, “between our two sources of low entropy.
of independence was created because the human economy
The solar source is stock abundant, but flow limited. The
was small relative to the planetary biosphere and sources of
terrestrial source is stock limited but flow abundant (tempo-
raw materials and sinks for wastes were relatively large
rarily, over the short term). Peasant societies lived off the
(Fig. 11.14a). As the human economy has grown, the source
solar flow; industrial societies have come to depend on enor-
and sink regions of the biosphere have diminished due to use
mous supplements from the unsustainable terrestrial stocks.”
and degradation, and so have their capacities to provide
Thus, the principal question was not, “How many people can
resources and absorb waste (Fig. 11.14b).
the Earth support?” but “How long can a population of any
Traditional economists have often committed the error of
given size be maintained?” Thus, Georgescu-Roegen wrote,
thinking of “nature” or “the environment” as a subset of the
“Every time we produce a Cadillac, we irrevocably destroy
human economy, a place from which to withdraw resources
an amount of low entropy that could be used for producing a
during production and a place to dump wastes after produc-
plow or a spade. In other words, every time we produce a
tion was completed. In fact, the human economy, regardless
Cadillac, we do it at the cost of decreasing the number of
of its past or present size and impact, will always be smaller
human lives in the future” (Georgescu-Roegen 1993:85).
than and contained within the physical environment, and is
Geoergescu-Roegen’s view of the economic process leads
ultimately dependent on it in order to function. And, because
to radically different conclusions than those of traditional free
the human economy must always be correctly perceived as a
market economics. First, capital and resources are not
subset of the greater economy of the environment, the human
substitutes, but complements. Agents of transformation can-
economy must make environmental constraints a more
not create the materials they transform or the materials out of
explicit consideration in producing goods and services and
which they are made. Further, Georgescu-Roegen questioned
disposing of the waste that such production creates (Costanza
how long economic growth, fueled by energy from nonre-
et al. 1997). Specifically, these constraints are manifested
newable resources, could last. Analytically, Georgescu-
economically as:
Roegen expressed it this way:
Fig. 11.14 Schematic depiction of the relationship between the econ- economy has grown, source and sink regions of the biosphere have
omy and the environment. The environment provides raw materials and diminished due to use and degradation, and so have their capacities to
energy, stock (nonrenewable) resources, flow (renewable) resources, a provide resources and absorb waste (b). (Reprinted by permission from
sink for wastes, and key life support systems. In the past, the human The McGraw-Hill Companies, based on concepts from Goodland, R.J.
economy was small relative to the biosphere and sources of raw A., Daly, H.E., El Serafy, S., 1992. Population, technology, and life-
materials and sinks for wastes were relatively large (a). As the human style: the transition to sustainability. Island Press, Washington, DC)
1. increasing capital costs of obtaining raw materials and and services, combined with methodologies for valuing nat-
energy when depletion occurs; ural capital, and (5) alternative measures of human well-
2. increasing inputs required to produce each output from the being.
same capital due to diminishing economic returns (e.g., Years ago, when asked whether a newly-independent
more fertilizer and pesticides may be needed for each yield India would follow the British pattern of economic growth,
unit in agriculture); and Mahatma Ghandi replied, “It took Britain half the resources
3. increasing demand for more effective and expensive pol- of the planet to achieve this prosperity. How many planets
lution prevention and clean-up (expressed as higher input would a country like India require?” (quoted in Kainer et al.
costs or higher government or household expenditures). 2006:4). Armed with such insight, Ghandi would have under-
stood why ecological economists Herman Daly and Kenneth
Although alternatives to growth economics are diverse, Townsend called the term “sustainable growth” an “impossi-
common threads among ecological economists are that eco- bility theorem” because “When something grows, it gets
nomic activity should: bigger. When something develops it gets different. The
earth ecosystem develops (evolves), but it does not grow.
1. be practiced on a sustainable scale, Its subsystem, the economy, must stop growing, but it can
2. use methods and practices of fair distribution of economic continue to develop. The term ‘sustainable development’
goods and services, and therefore makes sense for the economy, but only if it is
3. provide for efficient allocation of resources. understood as ‘development without growth’ – i.e., qualita-
tive improvement of a physical economic base that is
In addition, systems of ecological economics require: (1) a maintained in a steady state by a throughput of matter-energy
redefinition of “growth” and a differentiation among types of that is within the regenerative and assimilative capacities of
growth, (2) an explicit determination of environmental the ecosystem” (Daly and Townsend 1993:267).
constraints on economic growth, (3) definition of the Because confusion over “growth” and “development” is
functions of the environment in economic systems, (4) crea- rampant when discussing the economy, ecological
tion and organization of markets for environmental goods economists distinguish three types of economic “growth”:
Fig. 11.15 As an example, a (poverty-adjusted) Index of Sustainable absolute terms. (Reprinted by permission from Elsevier, Gigliarano, C.,
Economic Welfare (ISEW) plotted against traditional ISEW and Gross Balducci, F., Ciommi, M., Chelli, F., 2014. Going regional: An index of
Domestic Product (GDP) for Italy, on per capita basis. The ISEW per sustainable economic welfare for Italy. Computers, Environment and
capita was remarkably lower than the GDP, and the gap has widened in Urban Systems 45, 63–77. # 2014 Elsevier Ltd)
(1) growth of biophysical throughput; (2) growth in produc- income lost to future generations by the current depletion of
tion or income; and (3) growth of human welfare. “Through- exhaustible resources such as fossil fuels and other minerals,
put” refers to flows of matter and energy from the first stage as well as by the loss of biological resources, and adding
of production through consumption. Growth in throughput is estimates of environmental damage in the form of possible
probably the most environmentally harmful, as it is changes in climate, air and water pollution, and noise pollu-
characterized by high rates of consumption of matter and tion); and (4) unpaid household labor. The ISEW also adds
energy, relatively low efficiency of energy use, low durability the value of expenditures on maintenance and construction of
and rapid replacement of material artifacts, and increasing streets and highways, public health, and education, and it
rates of waste production. subtracts defensive expenditures on health and education
Growth in production or income has the potential to be (e.g., trauma care and remedial reading programs), spending
more environmentally benign. If increased production is on national advertising (aimed at increasing demand), and
characterized by more durable goods generated with less costs of urbanization, commuting, and auto accidents.
energy per effort, it may put less stress on environmental The ISEW is a dollar value that can be compared with per
systems in the long run. “Income” is technically defined as capita GNP, the traditional measure of economic well-being.
the flow of service through a period of time that is yielded by Whereas the per capita GNP and GDP have continually
capital (Daly 1991) but increases in income do not necessar- increased in the United States and other industrialized
ily represent more material consumption and accumulation. countries, the ISEW has decreased. Comparing ISEW to
Rather, they represent increases in “services” delivered and GDP also shows the difference between estimates even if
satisfaction with those services. Thus, it is theoretically pos- the trend across time is similar (Fig. 11.15). Other such
sible for income to increase while environmental degradation alternative metrics of human welfare include the Genuine
decreases. Progress Indicator, first adopted by the State of Maryland
Growth in human welfare is at once the most important (USA) in 2010, the Gross National Happiness Index, utilized
and most difficult to measure of the growth indices. Tradi- by the Kingdom of Bhutan since 1972, and the Happy Planet
tional free market economics has used per capita GNP (Gross Index which explicitly considers a country’s ecological
National Product) or per capita GDP (Gross Domestic Prod- footprint.
uct) as its usual index, but ecological economists question
whether either statistic measures human welfare. One alter-
native measure to per capita GNP is the Index of Sustainable 11.5.3 Implications of Challenging
Economic Welfare (ISEW), which integrates (1) income dis- Business-as-Usual
tribution (difference between the richest one-fifth of the pop-
ulation and the lower four-fifths); (2) net capital growth 11.5.3.1 Emphasizing a Broader View of Value
(measured as total net capital growth by adding increases to for Ecosystem Services
manufactured capital and subtracting the amount required to After decades of debate regarding whether or not we should
maintain the same per-capital level); (3) natural resource put a price on nature, this impasse has slowly given way to a
depletion and environmental damage (measured as the depre- more targeted discussion about the “specific conditions under
ciation of natural capital by subtracting an estimate of the which monetary valuation may or may not be appropriate”
Fig. 11.16 Methodological

toolbox for integrated valuation of
ecological, socio-cultural and
monetary values of ecosystem
services. (Reprinted by
permission from Edward Elgar
Publishing, from Gomez-
Baggethun, E., Martin-Lopez, B.,
2015. Ecological economics
perspectives on ecosystem
services valuation, in: Martinez-
Alier, J., Muradian, R. (Eds.),
Handbook of Ecological
Economics. Edward Elgar
Publishing, pp. 260–282)
including whether such valuations are “scientifically sound, neoclassical economists have emphasized that the fundamen-
socially just or ethically fair” (Gómez-Baggethun and Mar- tal value is pleasure, or utility, and can be measured and
tín-López 2015). There has been further emphasis by leading expressed as preferences in the market, many ecological
global organizations such as, the Intergovernmental Platform economists believe that monetizing value, however
on Biodiversity and Ecosystem Services (IPBES), to consider quantified, can only partly capture the comprehensive picture
the broad range of ways “human societies understand the of nature’s importance for informing policy decisions
importance of ecosystems and biodiversity” beyond econom- (Gómez-Baggethun and Martín-López 2015). The idea of
ically focused measures of value (Turnhout et al. 2014). Even value pluralism, therefore, stems from the “assumption that
with wariness about monetizing nature, Gómez-Baggethun valuation of ecosystem services involves dealing with multi-
and Martín-López offer two key reasons why ecological ple and often conflicting valuation languages, whereby differ-
economists should seriously engage the debate and insist on ent values deserve distinct attention in decisions and yet may
a fuller view of value than might otherwise be communicated not be reduced to a single metric” (Gómez-Baggethun and de
by a “market environmentalism that economizes ecology” Groot 2011; Chan et al. 2012; Gómez-Baggethun and Martín-
(2015). First, because a core premise in ecological economics López 2015). In Fig. 11.16, Gómez-Baggethun and Martín-
is that ecosystems and biodiversity are the material López highlight a methodological toolbox for an integrated
foundations of human societies, metaphors that highlight valuation of ecological, socio-cultural and monetary values of
societal dependence on ecological life support systems may ecosystem services based on method groupings in the litera-
be more necessary than ever in a world where people are ture. In practice, these categories have leaky boundaries and
increasingly alienated from nature. Second, despite the his- can utilize both quantitative and qualitative methods across
torical leadership of ecological economics towards the devel- scales (Table 11.4). The methods highlighted as monetary
opment of an ecosystem services approach, the mainstream values have been covered previously (Sect. 11.3).
adoption of this concept has resulted in directions that The ecological methods highlighted by Gómez-
diverge significantly from the original intent (Gómez- Baggethun and Martín-López relate to the ecosystem
Baggethun and Martín-López 2015). functions, processes and components on which service
The term ‘value’ has been defined by the Oxford Dictio- depends (de Groot et al. 2002). The metric of ‘value’ here
nary as ‘the regard that something is held to deserve; the is the “integrity of the biotic and abiotic components that
importance, worth, or usefulness of something’ or as ‘one’s contribute to ecosystem service provisions, including
judgment of what is important in life’ (2019). While populations, communities, functional groups, functional
Table 11.4 Tools for valuing ecological, socio-cultural and monetary values of ecosystem services, spatial scales at which they can be applied, and
quantitative vs. qualitative nature
Spatial scale Methodological approach
Values Methods Local Regional Global Quantitative Qualitative
Ecological values Material flow analysis ●● ●● ●● X
Emergy analysis ● ●● ●● X
Embodied energy analysis ● ●● ●● X
Exergetic replacement cost ● ●● ●● X
MuSIASEM ●● ●● ● X
Land cover flows ●● ●● ●● X
Socio-cultural values Q-methodology ●● ● X X
Mental models ●● ● X
Social network analysis ●● ● X
Role playing ●● X X
Preference ranking/rating ●● ● X
Photo-elicitation surveys ●● ● X X
Photo-voice surveys ●● ● X
Monetary values Production function ● ●● ●● X
Avoided costs/restoration costs ●● ●● ●● X
Travel cost ●● ● X
Hedonic pricing ●● ● X
Contingent valuation ●● ● X
Choice experiment ●● ● X
Benefit transfer ● ●● X
Notes: MuSIASEM ¼ Multi-Scale Integrated Analysis of Societal and Ecosystem metabolism
● ● good fit with the spatial scale
● partial fit with the spatial scale
Reprinted by permission from Edward Elgar Publishing, from Gomez-Baggethun, E., Martin-Lopez, B., 2015. Ecological economics perspectives
on ecosystem services valuation, in: Martinez-Alier, J., Muradian, R. (Eds.), Handbook of Ecological Economics. Edward Elgar Publishing, pp.
260–282
traits and habitat types” (Gómez-Baggethun and Martín- analysis, deliberative valuation and scenario assessment
López 2015). These values have in turn been linked to (Walz et al. 2019). Such methods might ask a person to
concepts of ecosystem resilience and capacity to sustain rank their preferences towards specific ecosystem services
services amidst disturbance and change (i.e., an ‘insurance in non-monetary terms (e.g., Oteros-Rozas et al. 2014) or
value’) (Armsworth and Roughgarden 2003; Pascual et al. clarify how their perceptions differs between positive and
2010), and in the development of ‘critical natural capital,’ negative aspects associated to ecosystems (e.g., Ruiz-Frau
which when measured in and preserved based on ecosystem et al. 2018).
stocks and funds, keeps ecosystems away from ecological An integrated assessment of ecosystem services across
thresholds (Deutsch et al. 2003; Gómez-Baggethun et al. categories of value and related methods gives monetary val-
2013). Finally, ecosystem valuation is often linked to bio- uation a role but does not elevate it as an “all-encompassing”
physical measurements, quantifying flows of materials like solution for capturing the importance of ecosystem services.
calculating energetic requirements or entropic costs of human In 2018, the Intergovernmental Science-Policy Platform on
activity (Gómez-Baggethun and Martín-López 2015). Under Biodiversity and Ecosystem Services (IPBES) revised their
the Millennium Ecosystem Assessment’s definitions, ecolog- own framework on ecosystem services to emphasize the
ical values are often connected most frequently to regulating broader view of “nature’s contributions to people” (NCP),
and provisioning ecosystem services. and clarified the ways that ecosystem services impact overall
The sociocultural values are those which, when elicited, quality of life on a continuum from instrumental values (i.e.,
capture the “material, moral, spiritual, aesthetic” motivations food, energy and water security) to relational values (i.e.,
people have regarding the environment and its conservation environmental justice and equity) (Christie et al. 2019).
(Gómez-Baggethun and Martín-López 2015). As they often Although monetary valuation techniques predominantly con-
include emotional, affective and symbolic views of nature, ceive of values as instrumental, socio-cultural valuation
such cultural services are represented through a family of techniques may go beyond instrumental towards relational
methods and techniques that include psycho-cultural valua- indicators of wellbeing.
tion, social valuation, participatory mapping, content
11.5.3.2 Reconsidering Wealth as Pathway

to Protection
One of the most basic and generalizable expressions of the
relationships between the economy and the environment is
the so-called Ehrlich Identity, formalized by biologists Paul
and Anne Ehrlich (Ehrlich and Ehrlich 1990) as
I ¼ P A T;
where I is environmental impact, P is population, A is afflu-

ence (a measure of consumption), and T is technology
(an index of efficiency of resource use and pollution abate- Fig. 11.17 Generalized example of an Environmental Kuznet’s Curve
ment). The identity is of little value mathematically (for one (EKC), which proposes uncontrolled pollution during initial development
thing, it is very difficult to express I in meaningful units or stage of a nation and then a gradual shift to environmental protection and
find common units for all the variables), but it is useful remediation as income (GDP) improves. Each x represents a different
country. (Reprinted by permission from Springer Nature, from Gara, T.,
conceptually. 2019. Sustainable development or Environmental Kuznets Curve model:
The Ehrlich Identity asserts that environmental impact is Which route for Africa? Environment, Development and Sustainability 21,
not simply a function of human population density, but also 1341–1356. # 2018 Springer Science+Business Media B.V.)
of per capita consumption (A) and efficiency of resource use
(T ). Furthermore, the relationship between impact and other
variables is complex and often non-linear. For example, patterns, especially those in which affluent countries shift
extreme poverty (very low values of A) often results in their heaviest polluting industries to less developed countries,
great environmental damage because of the direct and a practice known among environmental economists as “envi-
destructive manner in which impoverished peoples obtain ronmental dumping.” Such examples of redirected rather than
resources, but environmental damage of some forms may reduced environmental harm, emphasize the façade that tra-
actually decrease as people become more affluent. This rela- ditional growth pathways present to developing economies.
tionship is displayed graphically in a family of expressions In a special issue of the journal Ecological Economics that
generally known as Environmental Kuznets Curves (EKC). explored the hypothesis of the EKC, many articles further
Originally developed as an expression of the economist insisted that more explanatory variables should be included
Simon Kuznets’ theory that economic inequality increases in the analysis, improving our understanding of both the full
over time, and then at a critical point begins to decrease array of causal mechanisms and the resulting policy
(Kuznets 1955), the basic inverted u-shaped Kuznets’ implications. Some additional variables might be “measures
Curve was increasingly adapted to environmental economics of power (in)equity and social factors (Torras and Boyce
and policy as a way of relating environmental quality to 1998), trade-related measures (Suri and Chapman 1998),
income levels, beginning in 1991 when economists G. M. and spatial intensity of economic activity (Kaufmann et al.
Grossman and A. B. Krueger noted the appearance of EKC in 1998). Further, Rothman (1998) argued for increased inves-
their analysis of the potential environmental impacts of the tigation of the relationship between ‘consumption based’
North American Free Trade Agreement (NAFTA). From this environmental indicators (e.g. ‘ecological footprints’) and
point on, EKC has been used to show how specific measures of economic growth” (Gawande et al. 2001). Solv-
measurements of environmental quality, such as sulfur diox- ing environmental problems associated with growth “must
ide and particulates in the air (an index of one kind of air mean more than ‘passing them off’ to people in other times
pollution) will at first increase with a population’s per capita and places” as wealth is accumulated (Rothman 1998).
income (an index of its affluence) but, at some critical point,
begins to decrease as per capita income rises further
(Fig. 11.17). 11.6 Broader Linkages Between Economics
Indirectly, EKC is a simple and general expression about and Development
the relationship between wealth and environmental quality.
But its happy result (the richer you are, the less environmen- 11.6.1 The Origins of Sustainable Development
tal damage you will do) holds only in specific settings and
populations, and only in those cases because it does not The concept of “sustainable development” has dominated
consider some of the additional costs of affluence to discourse around the environment-development problem
ecosystems (Arrow et al. 1995). Some have argued the because it “promises to defuse longstanding tensions between
many EKC for individual variables are a result of trade environmental protection and economic growth” (Carruthers
11.6 Broader Linkages Between Economics and Development 479
2001). Ever since the Brundtland Report (World Commission sustainability were replaced with a familiar call for techno-
on Environment and Development 1987) and the Rio 1992 logical fixes that erase the line between ‘sustainable develop-
Earth Summit, conservation, especially in the Majority ment’ and ‘sustained economic growth’ and promised a
World, has been increasingly concerned with people, eco- future where the fundamental principles of “business-as-
nomics, and poverty. In fact, Agenda 21, which arose from usual” could remain intact. Is there a way to reclaim a more
the Earth Summit (Chap. 12), emphasized that “if poverty is historically “authentic” version of sustainable development?
the environmental problem, then lifting people out of poverty
is the solution. The best means to that end is the promotion of
economic growth—the boundless expansion of the economic
Contemporary discourse around sustainable develop-
pie” (Carruthers 2001). This “triple-bottom-line” approach to
ment assumes that economic growth and equity are
sustainability has been widely championed as the solution to
simultaneously possible. If economic growth was no
simultaneously address human need, environmental protec-
longer the goal of development, how might that change
tion, and economic opportunity, all made possible by techno-
our view of the environment and its resources?
logical innovation and free trade. As political scientist, David
Carruthers, writes:
It is little wonder that sustainable development today holds such 11.6.2 Integrated Conservation
broad appeal. How could it be otherwise? It is universally and Development
applicable. It dissolves the old conflict between growth and
limits. It eliminates confrontation over who is entitled to the 11.6.2.1 Benefiting Local Communities Through
lion’s share of remaining growth. It averts the question of north-
ern overconsumption. It promises the compatibility of environ- ICDPs
mental preservation with the maximization of growth. It supports Efforts toward sustainable development in recent years have
technological development and scientific progress. It offers increasingly emphasized the creation of integrated conserva-
equity for both present and future generations. And it plays a tion and development projects (ICDPs) to ensure that con-
mutually supportive role with the other western universals—free
markets and democratic politics (Carruthers 2001:102). servation can proceed in an appropriate cultural context with
sensitivity to human need. Typically, ICDPs include means
Carruthers then goes on to ask, “what happens if the most through which local people share the benefits of plant or
basic premises of the new sustainable development are mis- animal resources in their environment at sustainable levels,
taken?” Carruthers describes this approach to sustainable take ownership of the conservation of such resources, and
development as “new” because the conceptual origins of have an active role in decisions affecting the use and man-
sustainable development in the 1970s were in fact much agement of these resources in ways that benefit them individ-
different, and paralleled larger conversations about the “the ually and culturally. Originally implemented within tropical
age of scarcity” and “limits to growth” (2001). As noted ecosystems, ICDPs focus on community-based natural
earlier, the development of ecological economics, with its resource management that elevate alternative livelihoods
concerns about earth’s carrying capacity, boundless growth and poverty reduction. For example, one strategy for doing
as a biophysical impossibility, and emphasis on qualitative so might be just compensation for restricted forest access in
rather than quantitative views on prosperity, was coupled protected areas (Abbot et al. 2001).
with a challenge to reconsider the business-as-usual approach As emphasized by Blom et al. (2010), common
to development. In the 1970s, the concept that all boats rise characteristics of the most successful ICDPs include:
with global economic growth could not satisfactorily explain
the economic inequalities that persisted. The emphasis on • measurable and clearly defined goals for both conserva-
economic growth “at any cost” was no longer palatable, and tion and development outcomes
new formulations such as grassroots development, • funding for a duration that reflects the time commitment
pro-peasant development, eco-development, bottom-up needed to achieve states goals
development, and people-centered development were • economic markets for participant products and services
conceived as an “alternative, ecologically sustainable, and resulting from alternative livelihoods
socially-just alternative for the [global] south” (Carruthers • appropriate methods for monitoring and evaluation of
2001). There was also an emphasis on “‘appropriate’ or progress and outcomes
‘intermediate’ technologies, designed with local inputs and • a focus on conservation threats that are largely local and
knowhow, much cheaper than the capital-, import-, and within the scope of community-based action
energy-intensive technologies of the modern sector, but still • collaborative decision-making between project planners
offering dramatic improvements upon indigenous tools and and communities
techniques” that resonated with global institutions like the • visible and sustainable benefits for communities at an
World Bank, USAID, and UN agencies (Carruthers 2001). early stage of project implementation
However, in the span of a decade, the more radical origins of
These principles and others stem from decades of research

on ICDP implementation and have actively informed the Information Box 11.2 (continued)
projects of newer international strategies such as global
framework for reduced emissions from deforestation and
forest degradation (REDD). Such efforts have tried to take a
top-down approach to community engagement on emissions
reduction projects, drawing from voluntary payments for
ecosystem services initiatives (PES) and lessons learned
from ICDPs (Blom et al. 2010). Although some have
criticized the ICDP approach for its underlying assumptions
about linkages between poverty and conservation (McShane
and Newby 2004), it remains an attractive tool for conserva-
tion practice because it recognizes needs and resource rights
of local communities (Blom et al. 2010). Information Box
11.2 highlights lessons learned from an ICDP in Serengeti
National Park in Tanzania. Information Box Fig. 11.2 A Blue Wildebeest (Connochaetes
taurinus) in the Ngorongoro Crater, Tanzania. (Photo by M.
Mahdi Karim, reprinted under a GNU Free Documentation
Information Box 11.2: Bushmeat Hunting License)
in Serengeti National Park
Bushmeat hunting is a threat to wildlife populations
Barrett and Arcese determined that wildebeest harvests
throughout Sub-Saharan Africa. For example, in the
could not be sustained at projected levels in the face of a
north-west Serengeti, bushmeat is consumed regularly
growing human population (Barrett and Arcese 1998).
by 45-60% of all households and has increasingly
In fact, the authors concluded that any ICDPs “that
become a commercial enterprise (Rentsch and Packer
reduce essentially to game cropping [taking of animals
2015). An ICDP developed for Tanzania’s Serengeti
from a wild herd in numbers that will not endanger the
National Park and surrounding environs has
viability of the wild population] are likely to collapse in
incorporated wildlife harvest for local economies, indi-
less than one generation in the absence of other
vidual subsistence to meet the needs of a growing
interventions to mitigate game meat demand and
population, and effective local enforcement procedures
poaching” (Barrett and Arcese 1998:456). The model
that discourage poaching and delay biodiversity loss.
conservatively assumed human population growth rates
Barrett and Arcese evaluated the plan with respect to a
of 3.4% or 3.9% and, not surprisingly, the wildebeest
population model of the Serengeti wildebeest
population collapsed faster when human population
(Connochaetes gnou and C. taurinus) (Fig. 11.2).
growth was more rapid. The plan’s fundamental flaw
They determined that the number of wildebeest was
was its failure to address endemic causes of rural pov-
erty. Barrett and Arcese recommended that sustainable
N tþ1 ¼ N t S8w S4dt þ N 1 Rt eut H t development and concurrent conservation would be
best served if local agriculture were developed to be
where Nt reflects the number of wildebeest at time t, Sw more profitable and more sustainable, reducing the need
and Sdt are survivorship in the eight-month wet season for wildlife harvests and incentives for poaching. They
and four-month dry season, respectively, and Rt is the suggest that “successful ICDPs will combat poverty,
current recruitment rate. Losses are represented by risk, and food insecurity by changing the capabilities
harvest rate (Ht) of wildebeest by humans. Survival and incentives facing human populations on parks’
and recruitment rates are assumed to be functions of peripheries. . .” (Barrett and Arcese 1998:462).
food availability, determined from separate functions In general, the ICDPs involving wildlife harvests
(Barrett and Arcese 1998). Food availability is a func- share three problems. First, while human populations
tion of area, grass production, and rainfall. An average were typically growing, the harvested wildlife
household of seven consumes one-third of a wildebeest populations often were not. Second, no wildlife harvest
per year, leading to a regional take of 60,000 wilde- may be available during years of “environmental
beest annually. shock,” such as drought, that reduce recruitment in
Under their assumptions, the model predicted col- wildlife populations to near zero, yet low recruitment
lapse of the wildebeest population in 9–14 years, and in the wildlife population may coincide with periods of
reducing rural poverty, and achieving both objectives on a

Information Box 11.2 (continued) sustainable (self-financing) basis (Kiss 2004).
greatest human need, creating additional pressure on Effective ecotourism can lead to the protection of biodi-
wildlife and initiating a spiral of population decline versity in one of two ways. In the best scenario, earnings from
caused by having the heaviest harvests coincide with ecotourism are so high that local people give up all forms of
the lowest levels of recruitment. Finally, increased per destructive environmental labors or occupations and deliber-
capita income associated with development may lead to ately protect their environment to protect their new source of
increased demand for wildlife products in very poor income. Alternatively, ecotourism can protect biodiversity if
populations. Although such problems suggest that an outside group (business, NGO, or government agency)
induced demand for natural resources can impact provides initial funds and capital to a community to develop
resource stock levels and household welfare in com- ecotourism in exchange for the community’s pledge to pro-
plex ways, it also reinforces the importance of includ- tect local biodiversity. In this scenario, money does not, at
ing economic expertise to ensure household income least initially, come from ecotourism revenues but from
gains do not result in unintended consequences for investment capital, given to community members on the
conservation (Gilliland et al. 2019). expectation of future earnings. This kind of linkage is more
risky, since, if revenues fail to materialize, local residents
may nevertheless consider the initial revenues an entitlement
11.6.2.2 Ecotourism as a Payment for Protection
for protecting biodiversity that should continue whether suc-
Ecotourism has been defined by its own practitioners, such as
cessful ecotourism develops or not. In this case, if payments
the International Ecotourism Society, as “travel to natural
do not continue, local protection of biodiversity may be
areas that conserves the environment and sustains the well-
stopped. However, this second approach can be very effec-
being of local people” (IES 2019) (Fig. 11.18). In particular,
tive if properly managed. For example, the Amboseli Com-
such activities that sustain “the well-being of local people”
munity Wildlife Tourism Project in Kenya pays a “land
are usually identified with a specific form of ecotourism
holding rental” as soon as a village agrees to dedicate an
known as community-based ecotourism, or CBET. As
area of land for wildlife tourism. This rental is expected to
Agnes Kiss of The World Bank noted “. . .by the
stop once the tourism begins to generate revenues, but few
mid-1990s, USAID had 105 projects, totaling >US$2 billion,
communities have been willing to set aside land without this
with ecotourism components. . ., and 32 of 55 World Bank-
initial direct payment. However, once engaged, most
financed projects that supported Protected Areas (PAs) in
communities have been successful in protecting biodiversity
Africa between 1988 and 2003 included a CBET
and contributing to other conservation objectives and have
component. . .” (Kiss 2004:232). CBET is attractive because
captured and benefited from increased revenues from eco-
it offers the prospect of linking conservation and local
tourism (Kiss 2004).
livelihoods, preserving biodiversity while simultaneously
Fig. 11.18 A hanging bridge in

an ecotourism area of Thenmala,
Kerala in India. With
governmental support, Thenmala
is considered to be the first
planned ecotourism destination in
the country. (Photo by Augustus
Binu/www.dreamsparrow.net,
reprinted under CC BY-SA 3.0)
Ecotourism also can achieve biodiversity conservation on (3) achieve conservation objectives at the scale of
larger scales because, if it becomes embedded in the fabric of ecosystems; (4) provide clear, direct incentives for residents
a national economy as a major source of revenue, it begins to to actively protect habitat; (5) deter immigration; and
influence national policy. One tourism lobby persuaded the (6) reduce the social and political conflicts over resource
government of Ecuador to resist efforts to open a allocation that often endanger ecosystem survival (Ferraro
biodiversity-rich site in that country to oil exploration 2001). Can traditional ICDPs do all these things? Conserva-
(Wunder 2007). The government of Mozambique has tion policy expert Paul Ferraro summarizes the problem with
established large conservation areas as a key element of its ICDPs in a few words. “Experience with development
tourism development strategy (Kiss 2004). interventions over the last four decades indicates that simply
Although ecotourism has its share of success stories, the raising standards of living and encouraging economic growth
possible outcomes and linkages between biodiversity and is a major undertaking in many countries. . .Advocates of
ecotourism are only one manifestation of an intensifying development-based conservation interventions propose a
debate in the conservation community about the best ways much more difficult task. They propose, in effect, to guide
to use economic incentives to achieve conservation or control the development process so that specific behavioral
objectives. Ecotourism might be considered a form of pay- changes will occur and precise conservation objectives will
ment for ecosystem services. In this case the service is recre- be achieved. They are attempting not only to affect change,
ation or aesthetic experience, and the amount and kind of but to control the precise evolution of the change” (Ferraro
payment is determined by the private preferences of affluent 2001:992).
ecotourists. But preferences also can be expressed at national A frequent problem associated with the ICDP approach is
levels, such as through laws that protect endangered species, the high level of uncertainty regarding outcomes. Even if the
or even through international conventions, such as treaties development effort succeeds, its conservation benefits may
that aim to restrict global carbon emissions. The question is: be unexpectedly elusive and its unintended consequences
what form should such payments for these services take? unfavorable for conservation. Even when successful, ICDPs
ICDPs take the approach that, to succeed, conservation are problematic to conservation because of disjuncture at
must become an intrinsic value of local human communities temporal and spatial scales. Conservation needs are often
and must be fully integrated into community economic struc- urgent. Integrated economic-conservation development may
ture. The community that succeeds in such integration will take years to bear fruit. Conservation usually demands
receive the direct benefit of the ecosystem services they have ecosystem-wide strategies to preserve habitat and
conserved, which will come to be seen as costly goods vital to populations. ICDPs are community-specific, and their sphere
their own welfare. But an alternative approach takes a more of influence and effect no greater than that of the community
direct path to the solution. Why not make direct payments to in which they reside. Some ICDPs do attempt to operate on a
individuals or local communities for protecting or restoring larger scale, and some try to move development forward at
ecosystem services which benefit others, whether the local rapid rates, but as Ferraro points out, “When practitioners
community has fully integrated the benefits and values or quickly introduce new technologies, markets, and attitudes at
not? That question has sparked a vigorous debate that broadly large scales, they spread their resources thinly over a large
revolves around these two kinds of approaches: “integrated territory, thereby diluting or misdirecting their impact”
conservation development projects” (ICDPs), of which (Ferraro 2001:992).
CBET is one form, and performance payments, in which An alternative to the ICDP approach is the direct payment
money is given directly to individuals or communities for (DP) strategy, or payments for ecosystem services (PES).
meeting specified conservation objectives. Which of these is PES is a very specific and direct form of subsidy, a market-
the most effective way to achieve conservation goals? based approach to conservation we have covered previously
(Sect. 11.4.2.2). We give PES particular attention here
11.6.2.3 The Broader Debate: Integrated because it is becoming increasingly popular and provides
Development or Direct Payments stark contrasts to an ICDP approach in both its methods and
for Ecosystem Services? its assumptions.
Almost all conservation efforts today could be described as PES approaches are voluntary, conditional agreements
“interventions” that attempt to save species or habitats by between at least one “seller” and one “buyer” with regard to
trying to change human behavior at some level. An ideal a well-defined environmental service or, less directly, to a
“conservation intervention” that leads to local communities land use presumed to produce that service (Wunder 2007). In
achieving measurable conservation objectives should possess contrast to integrated development approaches, PES
the following characteristics: (1) be relatively simple in the strategies aim at short-term, sometimes immediate results,
senses that they allow practitioners to focus their energy on a and can often achieve them because they combine clear
few activities with high probabilities of success; (2) achieve performance indicators with explicit assumptions and
conservation objectives in both the short and the long term; informed consent of all parties. One category of PES schemes
Table 11.5 Profitability of Costa Rican forest protection against the profitability of logging under two PES scenarios. Values in US$
NPV, 5 years NVP, 15 years
Forest protection, Forest protection,
ha Logging PES ¼ $130 Logging PES ¼ $75
10 2108 2107 1741 2517
50 20,126 18,007 22,087 21,564
100 42,649 37,882 47,644 45,373
150 65,172 57,756 71,810 69,182
200 87,695 77,631 98,387 92,990
250 110,218 97,506 123,820 116,799
300 132,741 117,380 149,254 140,608
Reprinted by permission from Elsevier, from Ibarra Gené, E., 2007. The profitability of forest protection versus logging and the role of payments for
environmental services (PES) in the Reserva Forestal Golfo Dulce, Costa Rica. Forest Policy and Economics 10, 7–13. # 2007 Elsevier B.V.
For example, if forest stands (50–300 ha) were enrolled in a PES program for 5 years at the current PES rate ($130/ha), forest protection would be
less profitable than logging (assuming 2005 prices). If profit is a landowner’s main concern, it is unlikely that the current PES will motivate a
conscious choice of forest protection over logging, especially when the payments are issued for 5 years and income from logging is more certain in
the short run. Forest protection becomes an attractive alternative against logging when the annual award of PES matches a period of 15 years. In the
long run, a PES of $75/ha/year results in forest protection that is only marginally less profitable than logging. NVP = Net Present Value
for conservation are various forms of international habitat institutions and payment schemes. For example, Costa
reserve programs (IHRPs). An IHRP is “a system of institu- Ricans have created institutional mechanisms through
tional arrangements that facilitates conservation contracting which local, national, and international beneficiaries of eco-
through multiple actors and individuals or groups that supply system services compensate those who protect ecosystems.
ecosystem services. The contracts specify that the outside Costa Rica’s Forestry Law (no. 7575) recognizes four eco-
agents will make periodic performance payments to local system services: carbon fixation and sequestration, hydrolog-
actors if a targeted ecosystem remains intact or if target levels ical services, biodiversity protection, and scenic beauty. The
of wildlife are found in the ecosystem” (Ferraro 2001:994). law gives landowners the opportunity to be compensated for
For example, PES schemes might take the form of paying provision of these services. Funds for the program come from
landowners for conserving existing forests for their value in the National Forestry Financial Fund, sources for which
carbon sequestration. As Swen Wunder of Brazil’s Center for include fuel taxes and direct payments from other countries
International Forestry Research puts it, “The core idea of PES (e.g., Norway). The Fund establishes contracts for three
is that external beneficiaries of environmental services make management categories: reforestation, sustainable forest
direct contractual quid pro quo payments to local landowners management, and forest preservation, with each receiving a
and land users in return for adopting land and resource uses fixed annual payment per hectare. (Ferraro 2001).
that secure ecosystem conservation and restoration. . . this Direct payments have some advantages over ICDPs.
contingent conservation approach explicitly recognizes hard These are well-summarized by Ferraro. “Because perfor-
trade-offs and seeks to bridge conflicting interests by means mance payments can be targeted more precisely than devel-
of compensation” (Wunder 2007:49). opment interventions, practitioners can be more confident
Countries with well-developed government conservation that their interventions will have an effect on the areas
structures have some of the most effective payment targeted for conservation” (Ferraro 2001:995).
programs. In Canada, the United States, and much of western PES programs are far from perfect. The PES must provide
Europe, federal governments provide financial incentives to a profit incentive. If it does not, it will fail to protect land
farmers to keep land out of agricultural production or shift it areas where conservation is most urgent due to the projected
to alternative uses, thereby augmenting the supply of envi- revenue of competing land-use practices (Ibarra Gené 2007,
ronmental services. In Europe, 14 nations spent an estimated Table 11.5). In some cases, those enrolled take advantage of
US$11 billion from 1993–1997 to divert over 20 million ha agency or NGO incentives, particularly in PES schemes
of land into long-term set-aside and forestry contracts. In the centered on land use, by taking money for not doing some-
US, the Conservation Reserve Program spends about thing to the land that they never intended to do in the first
US$1.5 billion annually on contracts for 12–15 million ha, place. For example, in the Costa Rican program previously
an area twice the size of all national and state wildlife refuges discussed, Newburn et al. (2005:1417) note that, “In many
in the lower 48 US states. cases, the costs of forest conversion exceeds the expected
A similar approach can work in developing countries. returns from alternative uses (pasture, agriculture), meaning
Using direct payments, practitioners can focus their scarce these landowners have no intention of forest clearing during
resources on two key tasks: the design of appropriate the contract period and the opportunity costs are effectively
zero. The result may be that the Costa Rican government was
allocating funds largely to protect forestlands that are not at Specifically, how can we better choose what we shall
immediate risk of deforestation.” value, instead of treating our appetites, wants, and
Some believe that the problems with PES run deeper. By desires as givens that must be satisfied regardless of
delinking conservation from development, PES schemes environmental cost? And how can we restructure the
could deprive local communities in developing countries human economic enterprise so that it not only ceases to
from their own legitimate aspirations for development of degrade the world, but makes the human presence an
their own land and associated economies. Affluent conserva- agent of biodiversity conservation?
tion consortia could “buy off” local residents for relatively We can see, in individual communities and isolated
little money, preserving habitat and biodiversity but keeping efforts, that it is possible to make economic activity the
local people at low economic levels. A second concern, reflection of value rather than the determinant of it, and
essentially the opposite of the first, is that paying people to it is possible to make human activity a restorative
be good conservationists and land stewards will erode tradi- ecological force rather than an agent of ecological
tional, culturally-rooted, not-for-profit conservation values destruction. Today private economic incentives can
historically held by the community (Wunder 2007). PES is aid conservation because social values have been
not always the best strategy, but it may be “best suited to changed, and that change has itself been shaped by
scenarios of moderate conservation opportunity costs on laws and policies of the government that set certain
marginal lands and in settings with emerging, not-yet realized environmental and conservation values, such as
threats” (Wunder 2007). In fact, one recent review suggests endangered species, wetlands, clean air and clean
that while cash transfer programs can assist hundreds of water, above and beyond market forces. Markets and
millions of poor individuals, they must be implemented in property rights can be harnessed to achieve conserva-
tandem with environmental policies to “achieve welfare gains tion goals when they are made to serve socially norma-
and mitigate environmental damages that in turn economi- tive conservation values enforced by law and policy.
cally harm communities” (Gilliland et al. 2019:6737). But markets and property rights cannot intrinsically
generate conservation value, and their historic failure
to do so is an inarguable witness of the human experi-
Synthesis
ence. Conservation biologists, working with
Some conservationists, eager to make endangered spe-
economists, must offer a careful and well-designed
cies, critical habitats, and rare ecosystems able to stand
integration of conservation as an expression of human
toe-to-toe with industrial output, residential real estate
economic behavior that is guided toward conservation
development, and intensive agriculture, have devel-
goals established outside of the economic process
oped or employed a variety of creative measures to
itself. And in doing so, conservation biologists must
document the dollar values of their concerns, while
work to make conservation itself a normal pattern of
others, equally creative and passionate, have laid elab-
economic behavior, not simply a series of heroic but
orate plans through which humanity can continue to
ultimately futile efforts to save things that no one ever
take more but, through its increased ingenuity, degrade
really valued. This persistence towards a holistic and
the environment less via technological innovation.
ecologically-sensitive economic system will also help
Both these approaches, although well intentioned
us in our efforts towards “sustainability,” which will
and passionately advocated, have got the question
ultimately depend on how much “business-as-usual”
backwards. The first because it fails to ask whether
we believe our planet can tolerate.
current systems of individual-preference market-driven
valuations can ever rightly determine what is good for
many, or how people will ever become better than their
own self-centered appetites if those appetites are all
that determine their economic behavior. The second
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The Legal Foundations of Conservation Biology
12
To make the changes we need to make and to reach a safer future, we will need the resources of everybody here
— the scientists, the policy makers, and the industrialists — all working together towards a common goal. And
that goal is a planet that can continue to support life.
Dr. Piers Sellers, American astronaut
Keywords 12.1 Law and Policy as a Framework

Endangered Species Act · National Environmental Policy for Conservation
Act · Habitat conservation planning · Stockholm Confer-
ence · Conservation policy · CITES · Convention on 12.1.1 Nexus Between Conservation Science
Biological Diversity · UNFCCC · UNEP · International and Policy
conservation law
Conservation biology is a legally empowered discipline; that
is, it represents a scientific community that has received legal,
political, and cultural incentives and reinforcements towards
Overview the accomplishment of its goals (as outlined in Chap. 1).
In this chapter, you will learn about: Indeed, some have gone so far as to call conservation biology
a “regulatory science” that “seeks to develop scientific
1. The relationship between conservation law and pol- standards that can be applied to regulatory criteria and then
icy and the science of conservation biology. to develop management strategies to meet those standards”
2. Examples of national environmental laws in the (Tarlock 1994:1130). Throughout the world, the goals of
United States that have provided models for conser- conservation biology, including the preservation of biodiver-
vation in other countries. sity, protection of endangered species, and conservation and
3. Several key international conservation laws and management of ecosystems, are increasingly established in
how they define and empower conservation. and enabled by laws. This phenomenon speaks not only to
4. Specific case histories in which national and inter- the strong societal values these laws are meant to espouse and
national conservation laws have together influenced protect (as outlined extensively in Chap. 10), but also to the
the goals and practices of conservation biology. close connections between conservation science and public
policy.
In fact, conservation biology owes much of its early suc-
cess and continuing vitality to legal empowerment and sup-
port. Major environmental laws, such as the US Endangered
Species Act (ESA) (1973) and the United Nations Conven-
tion on the International Trade in Endangered Species
(CITES) (1973), among others, preceded the earliest

490 12 The Legal Foundations of Conservation Biology
organizational efforts to define the discipline. Although con- Oelschlaeger 1996). While traditionally, many scientists
servation biology might still have developed without national have avoided involvement in law- and policy-formulation
and international environmental legislation, it would have because they believed that such involvement would under-
been substantially less influential. Modern national and mine their professional objectivity and public credibility
global environmental legislation has affected and continues (much like the “pure scientist” Pielke describes, Information
to affect conservation biology in at least three ways. First, it Box 12.1), many conservation biologists disagree. Reed
has given legal incentives and approval for biodiversity pres- Noss, a former editor of the discipline’s most well-known
ervation. Second, it has affirmed the goals of conservation journal Conservation Biology said, “I believe that conserva-
biology and influenced the public to value conservation. tion biologists have a responsibility to enter the policy arena
Third, it has provided an environment that requires and and advocate both general principles and specific actions
sustains scientific research, management and monitoring as needed to conserve biodiversity” (Noss 1993:216). Putting
a function of good public policy. the matter more forcefully, conservationists Dwight Barry
Good science and its attendant empirical data are neces- and Max Oelschlaeger stated, “advocacy for the preservation
sary, but insufficient, for achieving conservation biology’s of biodiversity is part of the scientific practice of conservation
goals of stemming species extinction and ecosystem degra- biology” (Barry and Oelschlaeger 1996:905). Such
dation (Meffe and Viederman 1995). Conservation biology, statements might go too far if understood to mean that all
as a discipline, asserts that scientists can and should influence conservation biologists ought to be “issue advocates,”
environmental policy. Some even suggest that identifying as but they do clearly suggest that conservation biologists are
a conservation biologist “implies that we have an ethical uniquely motivated to clearly communicate outcomes for
obligation to provide decision makers with explanatory conservation along a range of policy choices (similar to
knowledge and prescriptive recommendations” (Barry and Pielke’s “honest broker”, Information Box 12.1).
Information Box 12.1: A Scientist’s Engagement

with Law and Policy politics: pure scientist, science arbiter, honest broker,
Roger A Pielke, Jr., a political scientist known for his and issue advocate (2007; Information Box Fig. 12.1).
work on environmental science-policy interactions, Although these roles are not always quite so distinct in
offers four idealized roles for scientists in policy and practice, they offer each scientist a degree of choice for
Criteria for determining the roles

of science in policy and politics
Is the decision context characterized

by both values consensus
and low uncertainty?
yes no
Connected to Reduce scope

policy? of choice?
yes no yes no
Science Arbiter Pure Scientist Issue Advocate Honest Broker
Information Box Fig. 12.1 Flow chart illustrating the logic of four idealized type, organizations and individuals working from the per-
roles for scientists in policy and politics. Often the political context spective of the honest broker, would provide a broad range of
shapes the role of science in policy and politics, including the degree options in support of non-politicized science. (Reprinted by permis-
to which society has a sense of shared values about desirable sion from Cambridge University Press, from Pielke, R.A., Jr. 2007.
outcomes and the means to achieve those outcomes. Furthermore, The honest broker: making sense of science in policy and politics.
science is always uncertain, and even in cases where a threat is Cambridge University Press, Cambridge)
clearly identified, science by itself cannot tell us what to do. As an
12.1 Law and Policy as a Framework for Conservation 491
generations, codifying aspirations or preferences into some-

Information Box 12.1 (continued) thing more lasting and transcendent. Laws empower action,
engagement. A pure scientist is one who may have no providing political resources and social force to achieve spe-
interest in the decision-making process, and simply cific goals, but laws also limit action by setting arbitrary and
wants to share fundamental information about a scien- fixed boundaries that may not correspond to the needs of
tific phenomenon with no consideration for its use or dynamic social or ecological systems. Because laws can be
utility. The pure scientist’s research may provide infor- difficult to repeal, they provide a sense of permanence to the
mation of value to decision-makers but the responsibil- values they establish. But laws also can become rigid, unre-
ity for information use lies solely with the decision- sponsive to changing conditions, and ultimately ineffective in
maker. A science arbitrator is one who serves as a solving the problems they were enacted to address. For these
resource, “standing ready to answer factual questions reasons and others, it is important for conservation biologists
that the decision-maker thinks are relevant” (Pielke to appreciate and accept the socio-political dimensions of this
2007). Such a person may clarify the meaning of tech- work. However, more than being overcome with fear of chal-
nical words and explain technical concepts, but like a lenge and compromise that participation in advocacy may
pure scientist, does not take an interest in any specific appear to bring, Harvard Professor of the History of Science,
policy outcome. Naomi Oreskes, suggests, “that society needs scientists to
In contrast, an honest broker is a scientist who speak up to alert society to challenges that, without science,
explains all policy options to decision-makers, we would not understand and might not even recognize”
clarifying the semantics, science, history, and potential (2017). In 1863, The National Academy of Sciences was
ecological, social, and economic consequences. In this chartered by the US Congress as a private, non-governmental
role, scientists work to provide comprehensive infor- organization to act as an advising body to the government and
mation and clarify outcomes, allowing decision-makers nation on scientific and technological matters (National
to sort through and reduce the scope of viable Academies 2019). This organization, and other national
alternatives according to their own preferences and equivalents like the Royal Societies in the Commonwealth of
values. Finally, an issue advocate is a scientist who Nations, provides one formal venue for scientists to “speak up”
promotes a preferred policy or policy alternative, effec- as they review research on a particular topic and draft white
tively reducing the scope of choice available to deci- papers on policy issues of special interest. However, there are
sion makers from the beginning of engagement. There many other informal avenues for science communication as
are multiple categories of issues advocacy ranging well, including public comment periods for new proposed
from “first-degree” examples, where scientists actively polices and civic engagement more broadly.
attempt to convince politicians or policy makers to As evidenced throughout this chapter, scientific informa-
adopt their preferred policies, to “second-degree” tion is essential to the development and implementation of
examples, where a scientist promotes a preferred effective law and policy. Closing the knowledge gaps
policy within a governmental or non-governmental between scientists and policy-makers, and increased commu-
organization, with the knowledge that the organiza- nication throughout the policy-making process will together
tion will actively attempt to convince politicians work to “support decision making through scientific inquiry
(Wilhere 2012). An example of “third-degree” advo- that is aligned with the medium- and long-term policy
cacy is when a scientist might transmit their policy priorities” (Fleishman et al. 2011:290). New generations of
preference through an article in a peer-reviewed journal conservation biologists who are both policy-savvy and scien-
or via a government technical report, but does not tifically grounded, will be on the front lines of solving the
attempt to convince politicians or policy-makers them- world’s most challenging conservation problems.
selves (Wilhere 2012). An individual scientist may
inhabit each of these four roles over time, depending
on their own interests and the perceived sense of 12.1.2 Defining Terms: Legal Frameworks
scientific uncertainty and value consensus around the and Linkages to Policy
issue.
Globally, there are thousands of environmental laws, and
corresponding policies, that regulate human interactions
with the environment. Generally speaking, environmental
To be most effective, one must not only comprehend both
law is rooted in three conceptual frameworks: ethical rights,
science and policy, but understand the inherent challenges
utilitarian interests, and equitable distribution of risks
that advocacy brings. On the one hand, laws represent current
(Salzman and Thompson 2013). As noted in Chap. 1, the
social values. But these laws also shape values for future
earliest laws addressing the use or treatment of plants and
animals were rooted in concepts of ethical rights. In ancient goal. More specific to our context, environmental legal
Roman, Chinese and Jewish legal traditions, animals and, in scholars James Salzman and Barton Thompson define envi-
some cases, even the land itself, were protected from certain ronmental law and policy as “the use of government author-
forms of abuse and mistreatment. Although the intention of ity to protect the natural environment and human health from
such laws, particularly toward animals, was not directed the impacts of pollution and development” (Salzman and
toward “conservation” as we understand it today, but rather Thompson 2013).
towards dispensing justice, these traditions did establish a At any level of government, the policymaking process is
basis for treating non-human creatures and ecosystems as complex, and includes a wide variety of values, stakeholders,
moral subjects. That is, non-human entities in the natural and sources of information as decision-makers try to select a
world were perceived as “morally considerable,” they could definite course of action among a range of alternatives. In
be treated in a morally right or wrong manner (Chap. 10). democracies, laws originate with issues that gain the attention
A second category of laws were prohibitions against the of politicians and government bureaucrats. But issues do not
use of plants or animals found on private property, especially become law and laws are not translated into policies without
if the property belonged to nobility. In some ways, these laws lengthy examination and development by all concerned
also were concerned with rights, in particular property rights, parties because one of the main functions of law is to provide
and what was protected was the right of the landowner to a framework for legitimating social norms. Laws are specifi-
enjoy a healthy, productive, or aesthetically beautiful envi- cally aimed to influence behavior and reinforce approved
ronment. Although such laws achieved a measure of protec- values by establishing normative rules that everyone must
tion for non-human species, the rights they protected were follow or else face punishment. Environmental and natural
expressions of privilege, not expressions of conservation. resource law often drive ongoing conservation efforts and
Laws of this type were rooted in utilitarian interests of the environmental protection, but to be effective, law must even-
landowner. Generally, conservation laws arising from tually be supplemented by attendant policies that support and
concepts of rights, grounded in moral values, tend to advo- clarify its intentions.
cate complete protection for the entity to be conserved, On a global stage, international cooperation through inter-
regardless of costs. In contrast, laws rooted in utilitarian national conservation law is critical to the world conservation
interests often use cost-benefit analyses (as discussed in enterprise. Much of international conservation law has been
Chap. 11) as the primary guide to making the correct or crafted from laws that were first developed in individual
“right” decision. In such a view, costs are not irrelevant, but nations. Even today, with a strong and growing body of
rather the most relevant and decisive decision-making factor. international conservation law designed to empower the
The third category of environmental laws focus on equita- world conservation effort, international laws and treaties
ble distribution of risks. Here, environmental justice, and its invariably suffer constraints. By the very nature of the diver-
intersection with civil rights law, provides a frame for sity of nation states, international conservation agreements
preventing the unequal distribution of environmental risk often descend to a ‘least common denominator’ approach in
and harm, focusing on the role of socioeconomic status species and habitat protection, usually united around trade or
along with race (American Bar Association 2017). While other forms of economic interests.
many of these laws traditionally focus on environmental The actions needed to preserve endangered species and
pollution and associated health risks, intergenerational equity their habitats must almost always be resolved at national and
and the rights of future generations has been a focus of recent local levels, not only because that is where local breeding
discussion with respect to the ongoing impacts of climate populations are resident, but because national and local
change and the sustainability of both our current economic communities are more likely to achieve a consensus of shared
and ecological systems (Mintzer and Michel 2001). values that support more aggressive and effective actions
In democratic governments such as the United States, laws needed to achieve real conservation goals. Further, even
are grounded in a constitution, a central document outlining international conservation laws become meaningless without
basic rights and foundational law, and carried out through national and local enforcement. The engagement of conser-
specific policies. One way to think of policy is as “a definite vation biologists is required at all of these levels, as such
course or method of action selected from among alternatives participation is essential for evaluating the effectiveness of
and in light of given conditions to guide and determine monitoring and enforcement. Here we review several
present and future decisions” (Merriam-Webster 2003). examples of foundational conservation law in the United
While many types of organizations can have policy, the States, before discussing several international agreements
specific actions of the government are generally considered developed to address conservation issues at global scales.
to be examples of public policy, or actions taken by the
government on behalf of the public in service of a specific
12.2 Foundational Conservation Law in the United States 493
12.2 Foundational Conservation Law non-economic values for resources and non-human creatures
in the United States (NEPA, ESA); (3) emphasized the status of individual species
and affirmed that extinction is undesirable (ESA); (4) stated
12.2.1 Common Characteristics of Effective that renewable resources were to be managed sustainably, and
Conservation Law that managers of non-renewable resources must take into
account the permanent consequences of present management
Although laws addressing conservation issues are diverse, actions (NEPA); (5) made federal funding available for
the most powerful and effective among them share important research and habitat acquisition (ESA); (6) provided citizens
characteristics that are now common in the environmental and NGOs with avenues for participation in decision-making
laws of individual nations throughout the world. Their shared and litigation against federal agencies (ESA, NEPA); and
traits include an inspirational and forward-thinking message, (7) given additional power to agencies to protect resources
the potential for growth in influence, an ability to attract and (ESA, NEPA). More than any other legislation, the radical
hold the interest of scientists because they raised questions transformation of conservation law achieved by these two acts
that must be answered by research, and a requirement for created the legal environment and social values in which
monitoring (Rodgers 1994). conservation biology operates today.
The inspirational and radical message of the strongest
modern environmental and conservation laws built a firm
foundation of moral and social support. Although court inter- 12.2.2 The US National Environmental Policy
pretation often has been necessary for the message to be Act (NEPA)
clarified and implemented, such a message has been latent
within all truly effective conservation legislation. According 12.2.2.1 NEPA’s History and Content
to legal scholar and law professor William H. Rogers Jr., In 1966, Indiana University (USA) professor of public
environmental laws “lack the compromised and ambiguous administration, Lynton K. Caldwell, published a paper enti-
form normally associated with an act of Congress” (Rodgers tled “Administrative Possibilities for Environmental Control”
1994:1014). Indeed, the most effective statutes in (Caldwell 1966) (Fig. 12.1). In his paper, Caldwell suggested
US environmental law were almost brazen in their language that qualitative environmental standards could provide the
and inspired popular support. The potential for growth in administrative coherence historically lacking in natural
influence allowed such laws to alter social values, and they resource policy (Caldwell 1966; Tarlock 1994). Caldwell’s
gained and held scientific support because they defined tasks paper, published in the book Future Environments of North
for scientists to perform and questions for them to answer. America (Darling and Milton 1966), would become one of
Many laws apply to topics of conservation concern, such the most influential publications on environmental policy of
as to forestry, soils, water, fish and wildlife, agriculture and the late 1960s.
parks. Sometimes these statutes are known collectively as The US Congress employed Caldwell as the principal
natural resource law, relating specifically to land manage- drafter of a law that was designed to be the centerpiece of a
ment and wildlife protection. Although some federal natural new era of environmental and conservation legislation, the
resource laws exist (Table 12.1), there are many more state National Environmental Policy Act of 1969 (Tarlock 1994).
laws that represent a range of resource use and protection In writing NEPA, Caldwell mandated that a “detailed state-
preferences based on state interests. The two notable ment” must accompany “proposals for legislation and other
exceptions to this patchwork of statues are the US National major federal actions significantly affecting the quality of the
Environmental Policy Act of 1969 (NEPA) and the human environment” (emphasis added; 42 U.S.C. §§4321-
US Endangered Species Act of 1973 (ESA), which are 4370h (1970)). This requirement led to the development of
often considered hybrids of natural resource law and regu- the now-familiar environmental impact statement (EIS) that
latory environmental law (which seeks to curb or “regulate” describes the possible environmental effects of actions pro-
negative environmental impacts like pollution). posed by federal agencies. Ultimately, policies and
NEPA and the ESA, passed and enforced separately but procedures associated with preparation of an EIS led to
often interacting legally, have radically altered the practice pervasive and well-defined procedures for public involve-
and enforcement of conservation values in the United States ment, as well as for challenging an EIS in court.
and, by imitation, throughout the world. This is why in this NEPA was signed into law by President Richard Nixon on
chapter we refer to these two statues as foundational conser- January 1, 1970, a fitting beginning to what would be called
vation law. These laws have (1) required that pollution or “the decade of the environment.” NEPA stated a national
environmental degradation be evaluated in the context of policy for the environment and formally established environ-
ecosystem function (NEPA); (2) endorsed intrinsic and mental quality as a leading national priority. NEPA expressed
its “inspirational and radical message” (Rodgers 1994) in
Table 12.1 Major US environmental and natural resource laws affecting biological conservation. There is no single “conservation law” within the
US and many other countries have parallel versions of these signature laws
Legislation Yeara Implementing agencyb Primary objective
National Environmental Policy 1969 (1975, 1982) Any applicable federal Promotes environmental protection by requiring federal
Act agency via the White agencies to evaluate the consequent environmental effects of
House CEQ proposed actions. These evaluations are conducted as
environmental assessments (EAs) and environmental impact
statements (EISs).
Endangered Species Act 1973 (1978, 1982, USFWS, NMFS To protect and recover imperiled species and the ecosystems
1988) upon which they depend. Among the benefits authorized for
listed species are: protection from being jeopardized by
federal activities, protection of critical habitat from
destruction or adverse modification, and required species
recovery plans.
Marine Mammal Protection 1972 (1994) USFWS, NOAA, Prohibits taking of marine mammals, and enacts a
Act MMC moratorium on the import, export, and sale of any marine
mammal, parts, or products.
Wild Bird Conservation Act 1992 USFWS Prohibits exotic bird species from being imported into the
United States unless there is an approved plan for sustainable
use of the species.
Lacey Act 1900 (2008) USFWS Prohibits trade of fish, wildlife or plants that have been
illegally taken, possessed, transported, or sold.
Pelly Amendment, Section 8 of 1978 USFWS Authorizes the President to limit the importation of any fish
the Fishermen’s Protective Act and wildlife products from a county of which nationals are
engaged in trade or taking that diminishes the effectiveness
of international conservation of threatened and endangered
species.
Migratory Bird Treaty Act 1918 (1936, 1960, USFWS Treaty between the United States and four countries
1974, 1978, 2004) (Canada, Mexico, Japan and Russia) prohibiting take,
possession, importation, exportation, transportation, and
purchase of any migratory bird and its parts expect under
federal permit; originally implemented the 1916 Convention
between the US and Great Britain (for Canada).
Antiquities Act 1906 NPS Authorizes the President to create national monuments from
federal lands that protect environmental, cultural, or
scientific values.
Magnuson-Stevens Fishery 1976 (1996, 2006) NMFS Establishes regional fishery management councils and
Conservation and Management includes national standards for management of fishery stocks
Act (MSFCMA) such as catch limits.
Multinational Species 1988–2004 USFWS A set of acts that provides grants to international projects that
Conservation Acts benefit elephants, rhinos, tigers, great apes, and marine
turtles and their habitats.
a
Year enacted; major amendments follow in parentheses
b
CEQ Council on Environmental Quality, USFWS US Fish and Wildlife Service, NMFS National Marine Fisheries Service, NOAA National Oceanic
and Atmospheric Administration, MMC Marine Mammal Commission, NPS National Park Service
these words: “It is the continuing responsibility of the federal without degradation,. . .(d) preserve important historic, cul-
government to use all practicable means, consistent with other tural, and natural aspects of our natural heritage,. . .
essential considerations of national policy, to improve and (e) achieve a balance between population and resource use
coordinate federal plans, functions, programs, and resources which will permit high standards of living and a wide sharing
to the end that the nation may: (a) fulfill the responsibilities of of the amenities of life, and (f) enhance the quality of renew-
each generation as trustee of the environment for future able resources and approach the maximum attainable recycling
generations, (b) assure for all Americans safe, healthful, proof depletable resources” (42 U.S.C. §§4321-4370h (1970)).
ductive, and esthetically and culturally pleasing surroundings, Robed in such positive platitudes, NEPA passed both houses
(c) attain the widest range of beneficial uses of the environment of Congress with relatively little opposition.
consider the value of non-economic resources, ensuring that

conservation would be considered in evaluating the proposed
action. Finally, NEPA introduced environmental assessment
as a means to guide administrative decision-making
(Caldwell 1966; Tarlock 1994). Thus, NEPA not only
established a mechanism for environmental review, but also
stimulated an increased level of citizen involvement in envi-
ronmental decision-making. Policy analyst Richard A. Liroff
summarized the true significance of the act when he noted,
“Implicit in NEPA was the notion that the public was to be
informed of the rationale underlying environmentally
impacting administrative actions. NEPA’s architects also
sought public involvement in decision making, but they did
not indicate when it should occur or what form it should take“
(Liroff 1976:88). It is also noteworthy that NEPA was
strongly linked to the kind of “ideal” role of government in
conservation that had first been developed by Theodore
Roosevelt (Chap. 1), embodying his vision of environmental
protection resting on a foundation of scientifically-informed
government decisions modified by citizen input.
These implicit notions of public participation ultimately
became explicit directives for public involvement, first
addressed by the courts in the case of Calvert Cliffs v. the
Atomic Energy Commission of 1971. In this case, the
Fig. 12.1 Lynton Keith Caldwell, Professor of political science and US District of Columbia Court of Appeals ruled that federal
public and environmental affairs at Indiana University from 1965–1984. agencies must comply with the procedural requirements of
One of Caldwell’s enduring legacies is his contribution to the formation
NEPA, including compliance with the preparation of a
of the National Environmental Policy Act of 1970 (Caldwell at home,
1972). (Photo courtesy of B. Cook, Indiana University Archives) detailed statement describing the environmental impact of a
proposed action, and that this requirement was in force even
It was not NEPA’s lofty opening rhetoric, however, that for an action by a private company or private individuals on
would have significant impact on US environmental policy. private land if the action required a permit from a federal
Hidden in the more mundane language of the law were words agency. Calvert Cliffs added legal precedent and enforcement
that would profoundly affect the practices and decisions of toward motivating US federal agencies to take seriously the
every US federal agency. The requirement that all federal requirement for an EIS for all “major federal actions.” To
agencies develop information, in the form of a “detailed better understand the scope of NEPA’s effect, one must
statement,” on the ecological consequences of their actions understand what constitutes a “major federal action.”
and weigh these impacts in their decision- and policy-making
would become the heart of NEPA’s power. Each such 12.2.2.2 NEPA’s Environmental Impact Assessment
“detailed statement” must describe (1) the environmental Process
impact of the proposed action, (2) any adverse environmental Under NEPA, a “federal action” takes place on federal lands,
effects that cannot be avoided should the proposed action be or is envoked if a proposed activity requires a federal agency
implemented, (3) alternatives to the proposed action, (4) the permit, is funded in significant part by federal money, or if
relationship between local, short-term uses of the environ- federal agencies have sufficient control or responsibility over
ment and the maintenance and enhancement of long-term a project to influence the project outcome. Any of these
productivity, and (5) irreversible or irretrievable situations constitute the “federal hook” that activates the
commitments of resources involved in the proposed action NEPA process. All federal agencies must comply with
should it be implemented. NEPA, but it has had a particularly significant impact on
NEPA was unique among environmental and conserva- the management of federal lands. The US government is
tion legislation in several ways. First, it was proactive rather the nation’s largest landowner, with responsibility for more
than reactive, forcing government agencies to consider the than 640 million acres (260 million ha), or 28% of total
environmental effects of proposed actions in advance. Sec- US land area (Vincent et al. 2017) (Fig. 12.2). Many of the
ond, NEPA forced government agencies to explicitly country’s western states are largely public domain, and more
than half of the land in Alaska, Nevada, Idaho, Oregon, Utah,
Fig. 12.2 The US government is the nation’s largest landowner, with of Land Management (BLM) and Forest Service (USFS) managing the
responsibility for more than 640 million acres (260 million ha). There is highest number of acres. (Image from the National Atlas of the United
a clear difference in the sum and distribution of federally owned land States, public domain)
between the western and eastern coasts of the country, with the Bureau
and Wyoming is federally owned. Four major federal land within a National Park. If a CE does not apply, the agency
management agencies collectively manage 95% of this area, must prepare an EA to determine whether the proposed action
including the Bureau of Land Management (BLM), Fish and or its alternatives have potentially significant environmental
Wildlife Service (USFWS), the National Park Service (NPS) in effects. The key and highly subjective word here is “signifi-
the Department of the Interior (DOI) and the Forest Service cant.” In determining significance, NEPA requires agencies
(FS) in the Department of Agriculture. This land is managed for to consider the geographic, biophysical, and social context of
many purposes, including preservation and recreation. Under- the action as well as the intensity or severity of the projected
neath this land area also lies a wealth of natural resources, impacts. Beneficial or adverse impacts may include those
including oil and natural gas, which is often subject to requests related to public health and safety, unique risks, effects on
for development (Vincent et al. 2017). endangered or threatened species, proximity to cultural or
The agency involved in a “federal action” initially historic resources, and cumulative impacts (Lamb 2018).
assesses the project proposal to determine the appropriate An EA must contain (1) a clear and concise description of
level of analysis under NEPA: Categorical Exclusion (CE), the proposed action; (2) a detailed description of the environ-
Environmental Assessment (EA) or Environmental Impact ment affected by the proposed action; (3) an assessment of
Statement (EIS). A CE applies to any routine action which the probable effects of the proposed action; (4) an evaluation
is already determined by the agency to cause no individual or of the probable cumulative and long-term environmental
cumulative effect on the quality of the human environment. effects, both positive and negative; (5) an assessment of the
An example of a CE may be the rehabilitation of hiking trails risk of credible potential accidents; (6) a description of the
Fig. 12.3 The process by which an Environmental Impact Statement clear if the action will have a potential significant impact on the envi-
(EIS) is determined to be necessary and then conducted under the ronment, an Environmental Assessment (EA) will be conducted.
direction of a supervising agency with public input. If the proposed (Graphic created by K. DeVoss, modified in style from USACE 2020)
action does not need environmental review, it might receive a categori-
cal exclusion (CE). If an environmental review is necessary but it is not
relationship of the proposed action to any applicable federal, responsible agency (if an action involves two or more
state, regional, or local land use plans and policies; and (7) a agencies, one is designated the “lead agency” and assumes
brief description of reasonable alternatives and their probable responsibility for the EIS) and describes the proposed action.
environmental effects, one of which is required to be that of Invitations, procedures, dates, times, and locations of public
not implementing the proposed action, the so-called “no meetings, with availability of related documents, also are
action” alternative. An EA differs from an EIS in scope, listed. Minimally, the NOI will be published in the Federal
length, and detail; however, an EA also requires substantial Register (online and in print) and sent to individuals who
agency investments of time, effort, and money. Many request it, individuals known to be interested in the proposed
agencies lack proper scoping in their analyses and often action, and national organizations expected to be interested in
create a mini-EIS in lieu of an EA, costing more time and the action. The NOI may also be in local newspapers,
money than originally intended (Lamb 2018). publicized through local media, and posted on the site to be
Each year, thousands of EAs are prepared, and many are affected.
presented to citizens and communities to invite public feed- As a first step in preparing the EIS, the lead agency will
back. However, unlike an EIS, a formal review and comment assemble an interdisciplinary team of professionals capable
process is not mandated. The outcome of an EA is either a of assessing the scientific, social, and economic issues likely
Finding of No Significant Impact (FONSI) or a determination to be addressed in the EIS. A team leader coordinates the
that an EIS is needed. If an agency submits a FONSI, the group’s activities to produce the EIS within specified
White House Council on Environmental Quality (CEQ) guidelines and deadlines, and assembles comments from
requires that the analysis is posted within the Federal Regis- other team members, other agencies, experts, and the public.
ter for a 30-day review period (CEQ 2007). The FONSI must EIS preparation requires regular contact among the lead
describe the action, the alternatives considered, and the envi- agency, and other relevant stakeholders. Public-issue identi-
ronmental effects and the reasons why they are not signifi- fication or “scoping” meetings involve the public early in the
cant. Individuals or groups unsatisfied with the FONSI or process. As part of the scoping process, the agency will
with the EA in general, can take the agency to court for not identify any existing or required studies, define the role of
preparing a full-scale EIS. all agencies involved, determine the relevant environmental
If the EA results suggest that an EIS is required, the most issues and describe the project’s rationale (Lamb 2018). After
rigorous review of NEPA compliance, the agency publishes a it is completed, the lead agency prepares an EIS implementa-
notice of intent (NOI) (Fig. 12.3). The NOI identifies the tion plan and uses it to produce a draft EIS (DEIS).
The lead agency conducts an internal review of its DEIS public in 2016, but they do not imply that these EISs were
and then publishes a “notice of availability” (NOA) in the prepared within a year. In fact, the average EIS preparation
Federal Register. Public comment on the DEIS, including time of those published in 2016 was 1378 days (~3.7 years);
comments received at public meetings where the DEIS is this is 698 days longer than the annual average recorded in
presented and explained, is then received, considered, and, 2000 (Nicholson 2017).
if appropriate, incorporated into a revision of the EIS. From In this same 2016 report, the NAEP found that the federal
this effort, a review draft of a final EIS is prepared, reviewed government prevailed in 83% of all NEPA cases brought
within the agency, and made available to the public for before the US Courts of Appeal (27 cases in total). Since
comment. Considering information presented in the final 2012, federal agencies have increasingly prevailed in the
EIS, the responsible official of the lead agency decides challenges to NEPA brought against them. Although 16 of
whether to implement the proposed action or one of the these controversies involved EISs, there were also 10 cases
alternatives (including the possibility of the “no action” alter- involving EAs and one involving a CE (Nicholson 2017). In
native) and publishes the decision in the Federal Register as many of these disputes, federal agencies were being
their “Record of Decision (ROD).” Usually, the ROD challenged over the adequacy of the NEPA documents they
initiates the ability for parties to sue in court to challenge produced, including assessment of impacts and inadequate
the EIS. However, the judicial review process in these cases mitigation of harm. In at least five cases, the federal govern-
is based strictly on whether the agency followed the proce- ment was being sued due to insufficient consideration of
dural rules for preparing the EA/EIS. There is no substantive alternatives (Nicholson 2017).
review of a final EIS.
Although the environmental impact assessment process is 12.2.2.3 NEPA’s Strengths and Weaknesses
intended to aid the agency in making a well-informed deci- It has been almost five decades since NEPA became law in
sion, the agency is not legally bound by the assessment the United States. Policy analyst Richard A. Liroff provided a
findings. This means that an agency can continue with a key to understanding NEPA’s profound effect on national
project that is found to have significant impact on the quality environmental policy by noting that “. . . NEPA laid the
of the human environment if plans for mitigating related groundwork for a series of procedures whereby environmen-
impacts are discussed. Two of the most important and often tal considerations could be fed into agency decision-making
controversial considerations of the NEPA process, therefore, routines” (Liroff 1976:210). NEPA procedures for environ-
are impacts and alternative analysis. Those developing a final mental assessment have radically changed the pattern and
EIS need to consider whether all impacts are truly being process of agency decision making. Before NEPA, major
considered and evaluate whether all reasonable alternatives agency initiatives, such as building a dam, executing a timber
have been carefully reviewed in light of agency goals and sale, or building a road in an environmentally sensitive area
concerns raised during public comment. Alternative analysis could be implemented with little or no consideration of their
is considered to be “the heart” of the EIS because to “rigor- environmental impact. That is no longer the case. And since
ously explore and objectively evaluate” the options means to NEPA, agency analyses of impacts and alternatives, coupled
be certain that the ROD is based on clear and convincing with public comment, have often led agencies to redesign
evidence and support (40 CFR § 1502.14). their projects in ways that are less environmentally damaging
There has been significant and consistent engagement by (Adler 2006; Pepper 2015). Such analysis has profoundly
the public and environmental and conservation affected the development of conservation biology because it
organizations in the process of EIS creation and review, and makes conservation issues relevant and legally mandated
plenty of opportunities for it. In a 2016 report, the National considerations in many proposed government actions.
Association of Environmental Professionals found a Notice And many individual US states have developed their own
of Availability (NOAs) for 312 environmental impact version of NEPA, extending environmental assessment to
statements published in the Federal Register. Of these state agencies and their funded projects.
312 published notices, 144 were draft EISs and 168 were NEPA has also transformed US environmental and con-
final EISs. Although 45 agencies were responsible for pub- servation policies into arenas for public participation rather
lishing at least one EIS, the majority of them came from eight than simply expressions of elected representatives.
agencies, with the US Forest Service responsible for Procedures for public input established by agencies and by
67 (Nicholson 2017). Considering the geographic breakdown US courts in response to NEPA set precedents regarding
of these draft and final EISs, seven addressed nationwide requirements for public input in most subsequent environ-
actions, and 89 addressed actions in multiple states. Regard- mental and conservation legislation. There are many ways to
ing their geographic distribution, 42 of the 50 states had at get involved in the NEPA process: (1) when the agency
least one EIS, the highest number (42) found in California. prepares its NEPA procedures, (2) prior to and during prepa-
These numbers focus on the EISs made available to the ration of a NEPA analysis, (3) when a NEPA document is
published for public review and comment, and (4) when Most recently, NEPA has been subject to critique from
monitoring the implementation of the proposed action and lawmakers who want to reform the statue to eliminate project
the effectiveness of any associated mitigation (CEQ 2007; delays that cost companies time and revenue (Hanson 2018).
McCuin et al. 2009). During each of these steps, non-agency In the words of the US House Committee on Natural
scientists can play a role in sharing appropriate scientific Resources, “. . .Germany, Canada, and Australia are all able
findings and argue for specific alternatives based on scientific to approve most major infrastructure projects within two
evidence. years. By contrast, a major infrastructure or energy project
No law is without challenges in interpretation and imple- in the United States can undergo a decade of environmental
mentation. One of the most foundational tensions in NEPA is review with no guarantee that the project will ever be
that it assumed an ecosystem management approach before approved” (as quoted in Hanson 2018). However well-
there were well-developed concepts and procedures of eco- intentioned, such changes, if motivated primarily by eco-
system management (Chap. 9). Specifically, NEPA’s intent nomic considerations, could reduce opportunities for public
was to provide for functioning, sustainable ecosystems and participation and erode a comprehensive review process
long-term environmental quality. However, its highest-level which relies on access to scientific assessment and full exam-
mechanism, the EIS, is usually prepared by one administra- ination of alternatives to reduce environmental impacts.
tive unit of a single federal agency, such as the staff of a Streamlining a 50-year old statute might be warranted;
national forest within the US Forest Service, operating doing so while retaining its original intent and navigating
within fixed spatial boundaries, limited jurisdiction, and the environmental stressors of the twenty-first century (such
strong vested interests in particular commodities. Some as land-use change and climate change) will be difficult.
experts now argue that NEPA will become more effective
as US resource management agencies mature in their under- 12.2.2.4 Adoption of NEPA in Other Countries
standing of and commitment to ecosystem management On NEPA’s 25th Anniversary, the White House CEQ
approaches, particularly those which acknowledge and incor- conducted a review on the effectiveness of Act, focusing on
porate climate change impacts. There is some evidence in how its performance aligned with the intent of NEPA’s
individual agencies that, in fact, this is the case, with more framers, and its impact on the global stage (CEQ 1997). At
recent EISs and decisions, particularly in the Forest Service, this point in time, NEPA had been emulated by over
reflecting more fully the true intent of NEPA in ecosystem 80 countries around the world and served as a model for
protection and less of simply following the rules of an admin- environmental impact assessments (EIAs) for global
istrative procedure (Goetz 1997). institutions like the World Bank. Some of the first countries
Additionally, NEPA mandates that the lead agency to adopt similar environmental impact policy in the 1970s
identify and inform stakeholders, but its procedures do not were Australia (Environmental Impact Assessment Policy/
always truly involve stakeholders as full partners in the Environment Protection Act), New Zealand (Mark I Environ-
decision-making process. Consequently, there can be dissat- mental Protection and Enhancement Procedures) and Canada
isfaction with the outcome of the final EIS and Record of (Federal Environmental Assessment and Review Process/
Decision that accompanies it. Judicial review does give the Canadian Environmental Assessment Act) (Clark and Canter
public some recourse to the perception of bad agency 1997). The environmental impact assessment process also
decision-making, especially if the agency did not address continued to develop across Europe, with efforts to integrate
substantive comments or include key stakeholders in the EIA procedures into member state decision-making pro-
decision-making process as mandated. However, some pol- cesses without the disruption or litigation that
icy analysts have argued for new, more creative approaches has characterized the American experience with NEPA
to the NEPA process. These have included such novel (Clark and Canter 1997). In contrast, the first efforts
propositions as the “citizen jury,” in which members of the in establishing EIA policies in the Majority World were
public evaluate the EIS and determine the decision by con- categorically different.
sensus, rather than the agency (Brown and Peterson 1993), or Most countries in the Majority World were asked to
the use of informal advisory groups that would have develop or follow EIA processes as a condition of receiving
continuing input to the agency’s interdisciplinary team (Sam- aid from development assistance agencies. Given that EIA
ple 1993). In 2015, the White House Council on Environ- procedures were perceived as a top-down requirement
mental Quality used a series of pilot projects to identify imposed by external organizations, there was often resistance
specific recommendations for improved stakeholder engage- and an accompanying lack of intrinsic political will from
ment, including refining and developing IT tools such as the leaders within these countries, especially given the percep-
EPA’s NEPAssist geospatial IT tool which allows tion that adoption of EIA procedures was a pre-requisite for
stakeholders to interact with assessment-relevant geospatial economic assistance (Clark and Canter 1997). Furthermore,
data (CEQ 2016). the legal foundation for elements of EIA processes at the
national level remained weak, particularly the inclusion of might be considered one of the strongest pieces of conserva-
public participation and the comprehensiveness of the tion legislation, it is also one of the most controversial,
impacts assessed. Today the EIA process remains unclear in enduring hostility and withering criticism from a range of
many countries, but the desire for environmental protection stakeholders.
within global sustainable development initiatives has First passed in 1966 as the Endangered Species Preserva-
given renewed focus to improving the transparency and tion Act, the original law was adopted with little controversy
effectiveness of EIAs throughout the world. At the 2017 or fanfare, and relatively little power. This version of the Act
Meeting of the Parties to the UN Economic Commission for limited protection to vertebrates native to the US, provided
Europe, Convention on Environmental Impact Assessments authority for only modest land acquisition for habitat,
in a Transboundary Context (i.e., Espoo Convention), focused on populations in existing wildlife refuges, created
participants stressed the importance of environmental no new programs or legal power, and was vague in its
assessments towards achieving the UN Sustainable Develop- language and intent. Its immediate successor, the Endangered
ment Goals, in particular Goal 13 which focuses on climate Species Conservation Act of 1969, was not much better,
action (Mead 2017). although it broadened the definition of “fish and wildlife” to
include invertebrates and prohibited the importation of
endangered foreign species except for scientific purposes
12.2.3 The US Endangered Species Act (ESA) (Nash 1989; Smith 1992). These legally-toothless statues
were rewritten in 1972 by E. U. Curtis Bohlen, then Under-
12.2.3.1 Historical Origins and Content secretary of the US Department of Interior, in ways that
The Endangered Species Act of 1973 (ESA) has been called profoundly changed the legal landscape of conservation in
the “strongest and most comprehensive species conservation the United States. In 1973, the Endangered Species Act
strategy” in the world (Rohlf 1991: 273). The ESA affirms (ESA) passed both houses of Congress with near-unanimous
the value of biodiversity, and actions authorized under the support.
ESA have contributed to the persistence of many endangered The 1973 statute expanded the jurisdiction of the ESA
species, and even the complete recovery of a few, such as the from vertebrates to most plant and animal species, legally
bald eagle (Haliaeetus leucocephalus) (Fig. 12.4). As of defining a “species” as “any subspecies of fish or wildlife or
April 2019, 1663 native species have been formally listed plants, and any distinct population segment of any species or
under the Act as either threatened (388) or endangered (1275) vertebrate fish or wildlife which interbreeds when mature”
in the United States (USFWS 2019a). Although the ESA (Sect. 3, 16 U.S.C. § 1531 et seq). Although this definition
was not scientifically satisfying (it assumes an understanding
of the very concept it is attempting to define), it was compre-
hensive in specifying an enormous array of organisms eligi-
ble for protection. The ESA also created a new category for
legal protection called “threatened species,” and even
allowed the listing of species that were threatened only in a
portion of their range.
The ESA gives primary authority for enforcement to the
Department of Interior’s Fish and Wildlife Service (USFWS)
for cases involving terrestrial and freshwater species and to
the National Marine Fisheries Service (NMFS) (sometimes
referred to as NOAA Fisheries) of the Department of Com-
merce for marine species. Introducing the concept of
“designated critical habitat” into environmental law, the Act
also requires the federal agencies to not only protect the
species themselves, but also the land or water in which it
lives. USFWS is given authority to identify, and in some cases
Fig. 12.4 The bald eagle (Haliaeetus leucocephalus), one of the first
purchase, such critical habitat, and to stop activities on such
species to be protected under the Endangered Species Act in 1967, was habitat that harm the species, even if the habitat is privately
officially delisted in 2007. While listed under the ESA, bald eagles owned. In Section 7, federal agencies are prohibited from
exhibited a dramatic recovery, from a low of barely 400 nesting pairs jeopardizing the continued existence of any threatened or
in the lower 48 states in 1963, to nearly 10,000 nesting pairs today. Bald
eagles continue to receive federal protection under the Bald Eagle
endangered species or destroying or adversely modifying its
Protection Act and the Migratory Bird Treaty Act. (Photo courtesy of critical habitat. And Section 9 makes it unlawful for a person
A. Morffew, reprinted under CC BY-SA 2.0) to “take” any federally listed fish or wildlife species.
Fig. 12.5 (A) The snail darter (Percina tanasi), a fish that delayed the an exemption from the ESA and construction was finally completed.
construction of the multi-million dollar Tellico Dam (B) on the upper (Photo of the snail darter courtesy of USFWS, public domain; photo of
Tennessee River, USA, by virtue of its protection under the Tellico dam courtesy of Tennessee Valley Authority, public domain)
US Endangered Species Act (ESA). Later, Congress granted the dam
Although perhaps an inconspicuous word, “taking” here is agreements “with any State for the administration and man-
legally defined as “to harass, harm, pursue, hunt, shoot, agement of any area established for the conservation of
wound, kill, trap, capture, or collect or attempt to engage in endangered species or threatened species,” and is authorized
any such conduct” (16 U.S.C. § 1531 et seq). to “enter into a cooperative agreement . . .with any State
The most famous challenge to the ESA occurred in 1978, which establishes and maintains an adequate and active pro-
only several years after being signed into law. In Tennessee gram for the conservation of endangered species and
Valley Authority v. Hill, the Supreme Court ruled that the threatened species” (16 U.S.C. § 1535). In fact, the ESA
Tellico Dam on the Little Tennessee River could not be actually helped stimulate the kind of federal-state cooperation
completed because the dam would destroy the habitat of a it envisioned by its very existence because, after its passage,
listed endangered fish, the snail darter (Percina tanasi) many states passed state endangered species laws modeled on
(Fig. 12.5) and cause its extinction. The Supreme Court’s the ESA.
decision substantially strengthened the ESA, particularly the
prohibition in Section 7 – at least on paper. Although 12.2.3.2 Species Listing and Delisting Process
environmentalists won the battle in court, their victory cost Under the ESA
them the war in congressional backlash to what many The ESA gives the USFWS and NMFS responsibility for
representatives now perceived as an act that was too restric- identifying endangered species and proposing these species
tive and insensitive to human need. Within a year, Congress for protection through the “listing” process (Fig. 12.6a).
had amended the ESA to create a committee that could waive However, actual listing is normally accomplished through
the law’s regulations under special economic conditions. interagency consultation, as specified in Section 7 of the
Although officially called the Endangered Species Commit- Act, because the ESA authorizes all federal agencies to
tee, this group soon became known as the “God Squad” “utilize their authorities in furtherance of the purposes of
because of its power to revoke the ESA’s protection for this Act by carrying out programs for the conservation of
selected species. The Committee ruled in favor of the fish, endangered species and threatened species. . .” The ESA
but Congress responded by excluding the snail darter from defines an “endangered” species as one that is “in danger of
protection under the ESA through a rider on an appropriations extinction throughout all or a significant portion of its range.”
bill. As for the obscure species that caused all the trouble, A “threatened” species “is likely to become an endangered
snail darter populations were transplanted and established in species within the foreseeable future throughout all or a
other streams, and the Tellico Dam was completed. significant portion of its range.” The term “foreseeable
The ESA also offers incentive for the federal government future” has been subject to much debate given the potential
to initiate cooperation with state programs as well as to impacts of climate change on species habitat (Wentz 2018,
cooperate fully with existing state programs to protect species more in Chap. 4). Proponents view climate change
(Section 6). For example, the Act states explicitly that the projections as providing sufficient evidence to consider a
Secretary of the Interior shall “cooperate to the maximum species for listing given the law’s definition, and such sci-
extent practical with the States,” may enter into management ence has contributed to the proposed listing of several species
Fig. 12.6 The process through which a species is “listed” (a) as Threatened or Endangered and subsequently “delisted” (b) under the provisions of
the US Endangered Species Act. (Diagram courtesy of US Fish and Wildlife Service, public domain)
such as the wolverine (Gulo gulo) and Arctic bearded seal as a potential “candidate” for listing as either “threatened” or
(Erignathus barbatus). Opponents claim that these projections “endangered.” Through the NOR, the agency makes an open
are not sufficiently reliable for ESA determinations. In 2018, call for all biological or scientific information on the species
USFWS and NMFS proposed changes to the regulatory frame- that could aid in their review. When considering whether
work they use to determine the “foreseeable future” that would or not to list a species, evidence is reviewed according to a
consider impacts of environmental change on a case-by-case five factor analysis: (1) present or threatened destruction,
basis (USFWS and NOAA 2018). modification, or curtailment of its habitat or range, (2) over-
As the first step in the listing process, the USFWS or utilization for commercial, recreational, scientific, or educa-
NMFS publishes a Notice of Review (NOR) via the Federal tional purposes, (3) disease or predation, (4) inadequacy
Register, specifying a species that the agency itself identifies of existing regulatory mechanisms, and (5) other natural or
manmade factors affecting its survival (USFWS 2016). These requires that listing determinations be based solely on the
agencies also re-assess species previously identified as best scientific and commercial information available; eco-
candidates but not yet listed, to update their status and deter- nomic impacts are not considered in making species listing
mine if they can either be removed from the candidate list due determinations and are prohibited under the Act. The agency
to ongoing conservation efforts or if their listing priority must publish their final rule in the Federal Register or
should change. As of April 2019, there were 22 species announce their decision not to list within one-year of the
across both the USFWS and NMFS that were actively date of the proposed rule. However, this time frame may be
being considered for listing as “candidates” (USFWS extended if there is substantial scientific disagreement. The
2019b). It is important to emphasize that species are not number of species listed each year has varied considerably
protected just because science says they are endangered or since 1967, with a high of 129 species in 1994 and a low of
threatened. They are protected only if they are officially one in 2007 (USFWS 2019b). Once a species is listed, it
listed. benefits from all legal protections afforded under the ESA.
The public can also suggest a species for listing via formal The species remains listed until it is considered “recov-
petitions, which is how most candidate species are now ered” and the protection provided under the ESA is no longer
identified. Once received, the USFWS or NMFS must review viewed as necessary. There is consequently a tight relation-
the supporting data and make a finding within 90 days of ship between the delisting process and species recovery.
receiving a petition (to the extent practicable) as to whether or Under Section 4 of the ESA, the USFWS and NMFS are
not there is “substantial information” indicating that the peti- responsible for the conservation and survival of each feder-
tioned listing may be warranted (USFWS 2018a). If this ally listed species by developing and adminstering species-
preliminary review finds that the data supports the need, a specific recovery plans (16 U.S.C. § 1533(f)). These plans are
more comprehensive status review is conducted using intended to outline objective and measurable criteria which,
biological and other scientific information to make a determi- when met, will result in the species’ removal from the list of
nation on the five listing criteria. Within one year of receiving endangered and threatened species (Himes Boor 2014). Full
the petition, the agency must decide whether a species’ listing recovery for most protected species is projected to take up to
is “warranted,” “warranted but precluded” or “not 50 years (Gies 2012; Nazzaro 2006). Consequently, recovery
warranted.” As of April 2019, there were 533 petition plans are required to undergo revision and review every five
requests under review by either agency (USFWS 2019b). years to more accurately assess and track a species’ progress
Due to the volume of species under consideration, many towards long-term survival.
species receive a “warranted but precluded” determination To delist a species, the USFWS or NMFS follows a process
and require re-review for each successive year until either a similar to when a species is initially listed (Fig. 12.6b). The
proposed listing rule is published or a “not warranted” goals of recovery are to reduce or eliminate threats, as well as
finding is made. With this designation, many species are to see populations achieve stable and healthy levels, so each
essentially placed on the candidate list indefinitely without of the five criteria used in the listing process is evaluated
the formal protections provided by the ESA. Given limited continuously during recovery and again in the process of
resources, USFWS works to evaluate all candidate species delisting (USFWS 2016). If an agency feels a species has
and prioritize them according to the degree and magnitude of sufficiently achieved these goals, it publishes a proposed rule
threat, followed by the immediacy of the threat, and, finally, in the Federal Register, which is open for public comment
the taxonomic distinctiveness of the species. In principle, no and includes a call for further scientific information. The final
preference is given to popular species or so-called “higher life rule (decision to delist) is posted again in the Federal Regis-
forms” (USFWS 2016). Often due to “higher-priority actions ter. If delisted, the agency must monitor the species
or listings,” some species linger on the candidate list for according to a post-delisting monitoring plan that evidences
decades, potentially declining as they wait. Several cooperation with States, Tribes and any other landowner or
US environmental organizations, such as Defenders of Wild- entity that has made commitments to long-term protection
life, Center for Biological Diversity, and WildEarth and management of the species or its habitats. However, the
Guardians not only submit petitions but have also sued the legal protections afforded to this species under the ESA are
agencies for delays in reviewing candidate species for listing. otherwise removed.
If the USFWS or NMFS find that a listing is “warranted”
the agency publishes a proposed listing (or proposed rule) via 12.2.3.3 Controversial Delistings: The Case
the Federal Register. This listing is then made available for of the Gray Wolf
public comment for a period of 60 days. In cases of high Lawsuits can be used to challenge both listing and delisting
public interest, the agency may also hold public hearings. All decisions under the ESA. One well established controversy is
substantial public comments and any new data received dur- over the proposed delisting of the gray wolf (Canis lupus). As
ing this public comment period is evaluated. The ESA a member of the “Class of 1967,” the first group of species
Fig. 12.7 Historical range of the

gray wolf (Canis lupus) across the
continental United States. Distinct
population segments (DPS) in the
Northern Rocky Mountains and
Western Great Lakes are all that
remain today. The Mexican gray
wolf population in the southern
US is considered a subspecies of
gray wolf and is not being
considered for delisting.
(Distribution map adapted from
CBD and THS 2018; photo of
gray wolf courtesy of Gary
Kramer/USFWS, public domain)
given federal protections, the gray wolf was suffering from in the 2011 budget that delisted the Northern Rockies popu-
declining populations across its entire historical range (much lation in Idaho and Montana, making further judicial review
of the contiguous United States) due to declines in prey impossible. This is the only species to ever lose its legal
populations and frequent conflicts with farmers and ranchers protections in this manner. Although unprecedented as a
(Jackson 2019). In fact, a population originally totaling in the method of delisting, this move reflects a deference towards
hundreds of thousands had been reduced to a few hundred states who want to manage the wolves themselves without
individuals located in northern Minnesota and Isle Royale federal interference and to protect the interests of their
National Park in Michigan (USA) (Collier 2018). Thanks to farmers (Furman 2011).
protections afforded as a listed species, including In March 2019, USFWS proposed to delist gray wolves
reintroductions of Canadian populations, the gray wolf pop- throughout the contiguous US, marking the ninth time the
ulation has rebounded. The USFWS notes that there are now agency has attempted to delist gray wolves since 2003 (Bies
more than 6000 individuals across the conterminous United 2018). This effort followed a successful initiative by the
States, exceeding the USFWS’s combined recovery goals for agency in 2017 to remove protections for the last remaining
the US Northern Rocky Mountains and Western Great Lakes listed population of the Northern Rocky Mountain DPS in
populations (USFWS 2019d, e; Fig. 12.7). Wyoming (USFWS 2017a). The proposed delisting has gar-
Originally listed as regional populations of subspecies, nered significant attention from both proponents of delisting
USFWS reclassified the gray wolf as a single endangered as well as opponents who had serious concerns about the
population at the species level in 1978 (except for the agency’s definition of species recovery. In May 2019, over
Minnesota gray wolf population, which was classified as 100 scientists and scholars issued a statement urging USFWS
threatened; USFWS 2019d). Although protected species to rescind its proposal citing concerns about (1) the definition
may be divided into “distinct population segments” (DPS) of grey wolf recovery, as current populations only inhabit
for purposes of listing and delisting, the agency has consis- 15% of their historic range; (2) inadequate consideration of
tently faced challenges in the courts for their efforts to delist the impact that genetic health has on the recovery of wolves,
“recovered” segments. For example, the Western Great Lakes as the current population is at only 5% of historic levels;
DPS and the Northern Rocky Mountain DPS were delisted (3) lack of inclusion of best-available science, leading to an
from the ESA in 2007 and 2008, respectively, but were put inappropriate shortcut of scientific review; and
back on the list by the courts in 2011 due to USFWS’s lack of (4) motivations for delisting based on local and special
consideration for how delisting a given regional population interests rather than the national interest (CBD 2019).
might affect other wolf populations remaining under federal In defense of the delisting, David Bernhardt, acting secre-
protection (Miller 2017). In that same year, Congress tary of the US Department of Interior, argued, “The facts are
responded to the court’s decision by including a provision clear and indisputable – the gray wolf no longer meets the
definition of a threatened or endangered species. Today the weakened and betrayed the Act’s original intent to preserve
wolf is thriving on its vast range and it is reasonable to endangered species regardless of economic cost (Nash 1989).
conclude it will continue to do so in the future. . .Today’s In 2018, the USFWS and NMFS proposed changes to the
action puts us one step closer to transitioning the extraordi- implementation of critical habitat designations in order to
nary effort that we have invested in gray wolf recovery to initially limit the amount of land being considered. For
other species who actually need the protections of the example, the agencies had previously considered critical
Endangered Species Act, leaving the states to carry on the habitat in areas both currently occupied and unoccupied by
legacy of wolf conservation” (USFWS 2019e). Here, the listed species simultaneously, but the proposed change
Bernhardt is claiming that the USFWS has fulfilled its obli- would require agencies to evaluate occupied habitat first. If
gation under the ESA and is arguing that a transfer of protec- such changes were adopted, the agency would be required to
tion is appropriate in order to allow more management first determine that occupied areas were inadequate to ensure
options for landowners and state agencies, including hunting. the conservation of the species before classifying unoccupied
Based on these public comments and others, the legal battle areas as critical habitat (Jarman 2018). The proposed rule also
over grey wolf delisting is likely to persist for years. With lists several circumstances under which no critical habitat
implications for other candidate and listed species, the fate of may be designated by the agency, including if the designation
the gray wolf will be of particular interest to conservation would increase the likelihood that species would be harmed
biologists, who have long advocated for its recovery. by human activity via poaching or vandals, or if there are no
habitat-based threats to the species (CRS 2018). Previous
12.2.3.4 Critical Habitat Designation for Listed litigation provides further context for this proposed change.
Species In 2010, USFWS designated areas of Louisiana (USA),
In managing an endangered or threatened species, the where the dusky gopher frog (Lithobates sevousus) had not
USFWS or NMFS must identify and protect a species “criti- been found in decades, as critical for the species’ long-term
cal habitat.” The ESA defines critical habitat as “specific conservation. Such designated areas included a privately-
geographic areas that contain features essential to the conser- owned parcel of land, and the USFWS was sued by the
vation of an endangered or threatened species and that may landowners as well as by a private company who had intended
require special management and protection” (16 U.S.C. § to use the land for residential and commercial development
1532, USFWS 2017b). In determining which features are (Weyerhaeuser Company v. United States Fish and Wildlife
essential, USFWS recovery biologists consider (1) space for Service). The argument was that (1) the closed-canopy timber
individual and population growth and for normal behavior; plantation on their land could not be critical habitat for the
(2) cover or shelter; (3) food, water, air, light, minerals, or dusky gopher frog, which lives in open-canopy forests and
other nutritional or physiological requirements; (4) sites for (2) USFWS had failed to adequately weigh the benefits of
breeding and rearing offspring; and (5) habitats that are designating the property against the economic impact. Lower
protected from disturbances or are representative of the his- courts upheld the USFWS designation as critical habitat,
torical geographical and ecological distributions of a species rejecting the landowners’ argument that the USFWS had
(USFWS 2018b). As of January 2015, critical habitat has acted arbitrarily and capriciously (Cornell Law School 2018).
been designated for 704 of the more than 1500 US species In November 2018, however, the Supreme Court unanimously
listed as endangered or threatened (USFWS 2018c). ruled that the appeals court must consider whether the USFWS
Agencies generally intend to finalize critical habitat had been excessive in its designation of “critical habitat.” Chief
designations concurrent with final listing rules, “to the extent Justice Roberts wrote, “according to the ordinary understand-
prudent and determinable” (USFWS 2016). Although the ing of how adjectives work, ‘critical habitat’ must also be
agency is prohibited from considering economic effects in ‘habitat’. . .Adjectives modify nouns – they pick out a subset
decisions regarding the listing of a species, amendments to of a category that possesses a certain quality. It follows that
the ESA in the 1980s added the requirement that the USFWS ‘critical habitat’ is the subset of ‘habitat’ that is ‘critical’ to the
conduct an economic analysis of the effects of designating conservation of an endangered species” (Supreme Court
critical habitat. Because such designation usually, but not 2018). Roberts argued that lower courts had concluded that
always, involves suspension of other activities in the area, “critical habitat” was “not limited to areas that qualified as
including economically profitable ones, the amended ESA habitat” and instructed them to decide whether the land in
includes an “exclusion process” through which all or part of question indeed qualified as the frogs’ “habitat” (Liptak
the critical habitat may be excluded from protection if the 2018; Supreme Court 2018). Even if the lower court decided
economic analysis determines that the cost of protection to affirm the parcel’s status as habitat, Roberts further
poses too great a hardship in economic or other forms of instructed the court to review the government’s assessment
loss. These amendments made the ESA more flexible in of whether the habitat designation had imposed a dispropor-
resolving conflicts, but, in the eyes of many conservationists, tionate economic burden on the landowners.

The ESA has recognized from the beginning the impor-
tance of protecting critical habitat along with the species
itself. How might changing the order by which habitat is
identified and prioritized for protection (i.e., currently
occupied vs. unoccupied) affect species recovery?
12.2.3.5 Preventing “Take” Through Habitat

Conservation Planning
Struggles arising from conflicts of interest between private
individuals and conservation efforts have repeatedly caused
what former US Interior Secretary Bruce Babbitt has called
“environmental train wrecks” (Kaiser 1997). As noted in the
previous section, conflicts of this sort have occurred because
early versions of the ESA did not define the concept of Fig. 12.8 The red-cockaded woodpecker (Leuconotopicus borealis),
critical habitat well and did little to develop the idea of saving an endangered species that has been the subject of intense management
through habitat conservation plans. (Photo courtesy of US Fish and
species through preserving habitats (Noss et al. 1997). How- Wildlife Service)
ever, landowner concerns arising from critical habitat desig-
nation (Section 4) have often paralleled those of agencies a condition that can be maintained only by recurrent fires and
who have witnessed intentional and adverse habitat modifi- active understory management. During the mid-1900s, the
cation by landowners with fears about the impact of species red-cockaded woodpecker declined in abundance to
presence on their land-use decisions. Recall that Section 7 of fragmented populations of only a few to several hundred
the ESA prevents landowners from “take” -- any act that individuals, with a total population of less than 15,000
harms or harasses the protected creature in any way, inten- birds. It was listed as endangered under the ESA in 1970
tional or not. Thus, as Bean et al. (1991) note in their analysis (Fig. 12.8).
of landowners’ conflicts with the ESA, “a landowner whose Most of the woodpecker’s historical habitat is on privately
bulldozers crush the larvae of an endangered butterfly on his owned land, and landowners feared that federal regulations
land commits just as much of a taking as a hunter who would be imposed on their land if the listed species was
deliberately shoots a bald eagle.” discovered on their property. As a result, landowners often
This view of “take” has significant implications for managed their land to make it unattractive to the
landowners. If a landowner inadvertently harms a member woodpeckers by harvesting pines before they reached
of the endangered species through normal land-use activities old-growth stages, replacing longleaf pine with shortleaf
such as farming, logging, or development, criminal prosecu- pine, suppressing fires, and letting the understory grow. For
tion can result, although such prosecution is rarely pursued. example, in the town of Boiling Spring Lakes, North
Since the courts do require harm to be foreseeable, not every Carolina, red-cockaded woodpeckers were beginning to
inadvertent action would count as violation. However, the move back into an area of longleaf pine woodland that was
risk of prosecution is a discouraging prospect that leads many being reviewed by the Fish and Wildlife Service for designa-
landowners to deliberately alter habitat on their land. If an tion as critical habitat. During the review period, many of the
endangered species inhabits their property, private individual pines that had been identified as “candidate trees”
landowners may resort to the strategy of the Three S’s -- for red-cockaded woodpecker nesting mysteriously
“shoot, shovel, and shut up.” The long-term effect of the disappeared (Rawlins 2006). Actions like those in Boiling
resulting behavior is a reduction in available habitat for Spring Lakes can arise from rational economic behavior and
already endangered animals. As Myron Ebell, a property- from the landowners’ fear of the ESA’s prohibition against
rights advocate, has said, “. . .if there is an endangered spe- the “take” of any endangered species.
cies on your land, the last thing in the world you want to do is To prevent continued loss of habitat for endangered species
provide habitat for it” (Cooper 1999). and reduce conflicts with private landowners, agencies have
A perfect example of Ebell’s quote in action is the case of resorted to a mechanism known as the habitat conservation
the red-cockaded Woodpecker (Leuconotopicus borealis). plan (HCP). HCPs arose out of a 1982 amendment to the ESA
The woodpecker inhabits the southeastern United States that allowed the issuance of “incidental take” permits for
where it usually lives in stands of mature longleaf pine endangered species. “Incidental take” was defined as take
woodlands and prefers open forests with minimal understory, that is “incidental to, and not the purpose of, carrying out an
otherwise lawful activity.” To be granted such a take permit, The no-surprises policy was intended to increase land-
the applicant, whether corporate or individual, must first pre- owner cooperation and make the protection of endangered
pare and submit a conservation plan. The plan must explain species more effective, but critics were quick to attack it. One
what the effects of the taking will be on the endangered hundred sixty-four scientists, including many of the world’s
species, how the effects will be mitigated, how implementation leading conservation biologists, wrote letters protesting the
of the plan will be paid for, and how the species will benefit. policy to members of the US House Committee on Resources
Now called habitat conservation planning, this procedure was (Walley 1996). Their greatest concern was that there will be
patterned after the resolution of an environmental/economic many surprises, rather than no surprises, in conservation
conflict over the proposed development of San Bruno Moun- planning. Because uncertainty and change are intrinsic to
tain near San Francisco, California (USA). ecological systems, the policy unreasonably and unfairly
San Bruno Mountain, attractive as a site for upper-class restricts the ability of agencies to change conservation plans
residential and commercial development, also represented and adapt to changing conditions. The policy also has been
some of the last undisturbed mountain habitat in the San criticized because it guarantees no surprises to the landowner
Francisco Bay area and was the home of two endangered as an inherent right, rather than as a privilege earned through
species of butterflies (Lehman 1995). Rather than resorting to proper conservation planning. According to the policy, the
litigation, the parties involved in the controversy devised a no-surprises assurance must be given to all landowners
series of agreements that allowed for development on whether or not they make conservation commitments
one-fifth of the mountain but protected the remaining 80% (Walley 1996).
and 90% of the butterflies’ respective habitats. Congress was Limitations of HCPs have led to attempts to improve this
so impressed with the San Bruno example that it codified it in approach. Increasingly, HCPs are supplemented with “no-
a 1982 ESA amendment so that HCPs would “encourage take” management plans implemented via a memorandum of
creative partnerships between public and private sectors and agreement (MOA) and so-called safe harbor agreement
among government agencies in the interests of species and (Costa 1997). MOAs are agreements between a federal
habitat conservation” (Lehman 1995). The process was agency (usually the USFWS) and a corporate landowner
intended to foster resolution through negotiation, compro- outlining conservation actions that the landowner can take
mise, and recognition of the interests of all participants. to meet or exceed requirements of the ESA for habitat pro-
Supporters of HCPs maintain that this approach involves tection. For example, landowners can satisfy their ESA
all vested interests and focuses on protecting the highest- obligations by monitoring populations, managing and
quality and most productive habitats (Lehman 1995; Evans retaining current and future nesting habitat, producing and
et al. 2016). Critics claim that the plans have inadequate maintaining foraging habitat, conducting cooperative
scientific guidance, permit landowners to destroy habitat research, education and outreach, and letting the managed
later if they enhance it initially (Kaiser 1997), provide few population provide donors for other populations (Costa
or no opportunities for public participation in formulating the 1997). One of the first agreements was signed in 1992 by
plans, and have ineffective management provisions and poor the Georgia-Pacific Corporation (a lumber company) and the
oversight of plan implementation (O’Connell 1997). Further- USFWS to preserve habitat for the previously discussed
more, most HCPs are for single areas or species, and red-cockaded woodpecker. Within five years, this MOA
landowners and critics argue that this approach is overly was protecting more than 66,000 acres of forest for the
narrow, restricted, and fragmented (O’Connell 1997). woodpecker (Costa 1997).
Despite these criticisms, officials in the Clinton adminis- Safe harbor agreements are contracts under which a
tration continued to work to make HCPs more attractive to landowner agrees to actively maintain suitable habitat (“safe
landowners. In 1994, the US Department of Interior and the harbor”) for a predetermined number of a species equal to the
Department of Commerce issued a new policy entitled “No number present on the site when the agreement was
Surprises: Assuring Certainty for Private Landowners in formulated. In return, the landowner receives an incidental
Endangered Species Act Conservation Planning.” This revi- take permit that authorizes future land-use changes or man-
sion, known as the “No-Surprises” policy, requires the agement on other parts of the site that may be occupied by
responsible federal agency to provide landowners with additional individuals of the endangered species. The major
assurances that they are not responsible for species protection benefit of the safe harbor agreement is that it provides direct
if unforeseen circumstances arise (Walley 1996; Schilling habitat improvement and maintenance for all the individuals
1997). Under this policy, after an HCP is approved, federal or population subunits that are enrolled in the original con-
agencies cannot require any additional mitigation measures servation agreement. Once again, the first example of the use
from a landowner to conserve an endangered species unless of a safe harbor agreement was for protection of the
the agencies demonstrate “extraordinary circumstances” that red-cockaded woodpecker (Leuconotopicus borealis). An
warrant increased protection. initial agreement in 1995 in the Sandhills Region of south-
central North Carolina succeeded in enrolling 24 landowners found that a landowners’ enrollment in the program led to
and more than 21,000 acres of habitat to be actively managed some potential benefits for population recruitment. While the
for the woodpecker. This acreage originally supported probability of cluster (egg) abandonment by female
46 woodpecker groups but was estimated to be able to support woodpeckers increased ~14% on control properties over
up to 107 groups (Costa 1997). The agreement was endorsed the 19-year period, it remained constant on Safe harbor
by the landowners because it was based on initial numbers of properties (Smith et al. 2018). This difference was likely due
woodpeckers present on a landowner’s property at the time to the creation of artificial cavities (used by the woodpecker
of enrollment. The landowner agrees to manage and monitor to lay and protect egg clusters) on enrolled lands (Fig. 12.9).
the habitat to maintain those numbers, but additional However, despite this finding, enrollment in Safe Harbor did
woodpeckers moving onto the property may be “taken.” not have a significant effect on most measures of breeding
In a recent review of the efficacy of the Safe Harbor success or foraging habitat quality. The authors suggest that
program towards the protection of the woodpecker (between because the “program permits habitat management to vary
1980 and 2014), biologist Jennifer Smith and her colleagues substantially across properties, it may have contributed to the
Fig. 12.9 Location of the study

area within (A) North Carolina,
USA, and (B) Hoke and Moore
counties, North Carolina, where
Smith et al. evaluated the
effectiveness of the Safe Harbor
program for the conservation of
red-cockaded woodpeckers.
Oxford University Press, from
Smith, J.A., Brust, K., Skelton, J.,
Walters, J.R. 2018. How effective
is the Safe Harbor program for the
conservation of Red-cockaded
Woodpeckers? The Condor 120,
223–233. # 2018 American
Ornithological Society)
absence of an overall Safe Harbor effect” (Smith et al. 2018: The ESA has performed well at the functional level and
231). Furthermore, a lack of consistent funding for habitat operated in the courts more efficiently than many other legal
management through federal and state programs, such as the attempts to preserve biodiversity because it contains easily
Wildlife Habitat Incentive Program, Landowner Incentive defined concepts and goals. In particular, the “species” con-
Program, and Private Stewardship Grant Program, may cept, the cornerstone of the ESA’s validity, has proved more
have been a factor, especially as these programs are often definable and defensible in legal circles than have concepts
limited to large landowners. such as “biodiversity,” “habitat,” or “ecosystem” (Karr
Despite their imperfections, conservation approaches like 1995). Perhaps most importantly, the ESA remains an impor-
HCPs, MOAs, and safe harbor agreements acknowledge tant legislative model for efforts to save species worldwide.
fundamental truths about the future of conservation. First, Not only has the ESA been used in tandem with international
habitats must be conserved if species are to be conserved, and law (i.e., CITES) to protect foreign endangered species with
secondly, habitat and species conservation cannot be success- prohibitions of import, export, take and commercial activity,
ful in the long run if they are restricted entirely to public land but has been replicated successfully in other countries in the
or to private reserves established by conservation European Union as well as Canada and Australia.
organizations. Habitat and species conservation can be suc- The law has also faced its share of critiques. Complaints
cessful in a landscape context only if private landowners are from private business and development interests are chronic
involved and motivated partners. These realities reveal that and predictable, with much of the focus targeted at land use
efforts of greater landscape scale are needed to preserve concerns. The ESA also has been increasingly subjected to
populations and their habitats. It is far easier and more cost substantive criticisms from conservation biologists. Perhaps
effective to protect intact landscapes and the species they the most substantive biological criticism is that it is reactive
contain than to initiate emergency measures for critically rather than proactive, responding only to the needs of species
endangered populations on degraded habitat. on the brink of extinction rather than focusing on the conser-
vation of overall biodiversity and the management and pro-
12.2.3.6 Strengths and Weaknesses of the ESA tection of critical habitats and ecosystems (Rohlf 1991). One
The ESA has become “the pit bull of environmental laws. . . constructive response to this criticism has been habitat- and
It is short, compact, and has a hell of a set of teeth. Because of regional-level analysis of endangered species’ distributions,
its teeth, the act can force people to make the kind of tough and development of strategies to promote the recovery of
political decisions they wouldn’t normally make” (quoted in multiple species in the same habitat or region (Flather et al.
Rosenbaum 1995:334). More than any other statute, the ESA 1998).
affirms that species have intrinsic value, and US courts have Some biologists have also expressed concern about the
interpreted the Act to give protection to any species listed as overall effectiveness and scope of the recovery planning
“endangered” or “threatened” regardless of the economic cost process (e.g. Glick 2005; Gregory et al. 2012; Himes Boor
of protection (Rohlf 1995). The ESA has also clearly and 2014; Neel et al. 2012). For example, there is a clear gap
explicitly extended legal protections to non-human species between available resources (e.g. funding, staff, etc.) and the
(Karr 1995). The US environmental historian Joseph Petulla needs of recovery plan programs (Restani and Marzluff
described the ESA as one of the most remarkable, radical, 2002). This gap is even more apparent among particular
and original laws ever passed because, through its protection, taxonomic groups, with large charismatic megafauna receiv-
“a listed non-human resident of the United States is ing the most funding (Miller et al. 2002). Furthermore,
guaranteed, in a special sense, life and liberty” (Petulla 1977). recovery plans themselves can be tedious and unwieldy
The ESA has been instrumental in preserving many spe- documents that vary greatly in quality (Foin et al. 1998;
cies, albeit often at small population sizes. The USFWS notes Boersma et al. 2001). Poorly developed recovery plans
that the ESA has prevented 99% of extinctions, that of often exhibit limited nonfederal participation and are typi-
roughly 291 species, with only 4 species having ever cally linked to a lack of dynamic and explicit science within
been confirmed extinct after being listed (Greenwald et al. the recovery planning process (Boersma et al. 2001; Lamb
2019). Furthermore, although there has been criticism leveled et al. 2014; Weijerman et al. 2014). One review showed that
at the low number of species delisted (39 species, or ~1.3% of although recovery goals for approximately 90% of
all listed species since 1967), this statistic masks progess invertebrates required a specified number of populations for
made on species recovery (Greenwald et al. 2019). Many downlisting or delisting the species, the current number of
argue that the very listing of endangered species strongly populations was not known for 35% of these species (Tear
contributes to the species’ survival. Without such protection, et al. 1995). Furthermore, even when revised recovery plans
many species would have likely become further endangered make efforts to include more information regarding species
or extinct. biology, status, and threats, they simultaneously fail to
clearly link this information to management actions, moni- law, and (4) judgments of an international court or tribunal
toring protocols, or recovery criteria (Harvey et al. 2002). (Sands 1999:122). Most bilateral (between two countries)
Other biologically based criticisms of the ESA include and multilateral (three or more countries) environmental
complaints that the law lacks defined thresholds to delineate agreements take the form of a treaty. The starting point for
endangered, threatened, and recovered species; that it does determining what constitutes an international treaty is the
not adequately protect patchily distributed populations 1969 Vienna Convention on the Law of Treaties, with
(“metapopulations”); that it does not protect habitat reserves many of its provisions considered binding on all nation states.
sufficiently to sustain recovered populations; and that uncer- The Vienna Convention defines a treaty as “an international
tain or long-term threats to endangered populations are agreement concluded between states in written form and
discounted (Rohlf 1991). As one potential response to these governed by international law, whether embodied in a single
concerns, a recently proposed rule by USFWS (January instrument or in two or more related instruments and what-
2019) seeks to amend up to 182 recovery plans, covering ever its particular designation” (Article 2(1)(a)). There can be
205 plant and animal species, to include “quantitative recov- confusing terminology around international treaties, given
ery criteria” which would improve the roadmap for listed that the term also encompasses other terms such as conven-
species’ recovery and provide guidance to conservation tion, agreement, pact, protocol, charter, statute, covenant,
partners on how to minimize threats to the species (WGA engagement, accord, and Memorandum of Understanding.
2017; USFWS and NOAA 2019). In most cases, the term used to describe each specific
treaty indicates a differing degree of political and practical
significance. A “charter” is a particularly formal designation,
12.3 International Conservation Law often signifying a constituent (establishing) treaty such as the
1945 Charter of the United Nations and 1963 Charter of the
12.3.1 Understanding Key Terms Organization of African Unity. The designations of “conven-
tion” and “agreement” are often used with multilateral legal
Today conservation is an international effort involving all instruments, rather than bilateral ones. While the term “agree-
modern nation states to varying degrees. Although interna- ment” can be used to describe the collective range of interna-
tional conservation law does, in some cases, implicitly rec- tional legal instruments, this term is also frequently used to
ognize the intrinsic value of the species and habitats it describe a specific restricted multilateral treaty with a
preserves, it is primarily driven by utilitarian interests and narrower range of subject-matter and is considered to be
equalization of risks, usually in the form of increased mutual less formal than those legal instruments with “treaty” in
international interdependence and concern for intergenera- their title (InforMEA 2019). “Conventions” are considered
tional equity. The primary agent that brokers international to be open to participation by the entire international com-
conservation initiatives and multinational agreements is the munity. Most international instruments regarding environ-
United Nations (UN) and, within the UN, its Environmental mental protection and sustainable development are given
Programme (UNEP). Largely through the impetus of UNEP this designation by the United Nations. One prominent exam-
and other UN environmental programs, nation states have ple is the 1992 UN Framework Convention on Climate
entered into over 325 treaties, conventions, and agreements Change (UNFCCC). Protocols are legally binding
focusing on international conservation during the last agreements entered under the authority of specific treaties
40 years, and today, over 1000 international legal (think of the Kyoto Protocol and the Paris Agreement as
instruments, most of them binding, contain at least one sec- subsidiaries to the UNFCCC). They contain specific substan-
tion or provision that addresses environmental conservation tive obligations to implement the more general text in a
(InforMEA 2019; Table 12.2). At the heart of advisement and framework convention through what is often called the
enforcement is the International Court of Justice (ICJ). All “framework-protocol approach.” Finally, the term “declara-
members of the United Nations are ipso facto (automatic) tion” is often used for international instruments that are not
parties to the ICJ, whose work is to render advisory opinions legally binding but instead declare a set of global aspirations,
on any legal questions as requested by the UN General such as the 1992 Rio Declaration on Environment and
Assembly. The ICJ plays a role in both developing interna- Development.
tional law, and handling disputes among contesting nation Treaties undergo multiple rounds of negotiation and pro-
states, though as a practical matter, very few conservation cedure before becoming enforceable. When major multilat-
cases are decided in the Court. eral treaties are first adopted and open for signatures, all
Modern efforts in international conservation law arise countries are invited to participate and show their support
from one or more of the following sources: (1) bilateral or visibly by becoming a “signatory.” Although all countries
multilateral treaties among nations, (2) binding acts of inter- who have signed a treaty should immediately refrain from
national organizations, (3) rules of customary international acting contrary to its objective and purpose, the treaty is often
12.3 International Conservation Law 511
Table 12.2 Examples of multilateral treaties regarding biological conservation and enviornmental protection
No.
Convention or treaty namea Yearb of Parties c Primary objective
Bamako Convention 1998 25 Prohibits the import into Africa of any hazardous (including
radioactive) waste. Covers more wastes than those included under
the Basel Convention.
Basel Convention 1992 186 Protects human health and the environment against the adverse
effects of hazardous wastes. Its scope of application covers a wide
range of wastes defined as hazardous based on their origin and/or
composition and their characteristics, as well as household waste
and incinerator ash.
Carpathian Convention 2006 7 Provides a framework to pursue comprehensive policy and
cooperation in order to guarantee protection and sustainable
development of the Carpathians, one of Europe’s largest mountain
ranges.
Convention on Biological Diversity (CBD) 1993 196 Promotes the conservation of biological diversity, including
through the sustainable use of its components, and fair
and equitable sharing of genetic resources.
Convention on International Trade in Endangered 1975 183 Ensures international trade of wild animals and plants do not
Species of Wild Fauna and Flora (CITES) threaten their survival. Degree of protection afforded depends on
Appendix level designation (I, II or III).
Convention on the Conservation of Migratory 1983 130 Provides a legal foundation for internationally coordinated
Species of Wild Animals (CMS or the Bonn conservation measures throughout a migratory range, including the
Convention) conservation and sustainable use of those animals and their habitats.
Minamata Convention on Mercury 2017 118 Protects the environment and humans from the damaging effects of
mercury pollution by controlling anthropogenic releases throughout
its lifecycle.
Rotterdam Convention 2004 161 Promotes cooperation and shared responsibility in the international
trade of hazardous chemicals to protect the environment and human
health.
Stockholm Convention on Persistent Organic 2004 182 Aims to restrict or eliminate the production, release and use of
Pollutants persistent organic pollutants (POPs) in order to protect human
health and the environment.
Tehran Convention 2006 5 Provides a framework for the sustainable use and environmental
protection of the Caspian Sea region and with it the livelihoods,
health and well-being of present and future generations.
Convention on Nature Protection and Wildlife 1942 22 Parties to establish parks, wilderness reserves and national
Preservation in the Western Hemisphere monuments to protect and preserve native fauna and flora species in
their natural habitat, and preserve geological, aesthetic, historic and
scientific areas of extraordinary, unusual and striking significance.
Convention on Wetlands of International 1975 170 Framework for national action and international cooperation for the
Importance, Especially as Waterfowl Habitat conservation and wise use of wetlands and their resources.
(Ramsar Convention)
Convention to Combat Desertification 1996 197 Aims to limit desertification and mitigate the effects of drought
through international cooperation by enacting national programs
and strategies focused on sustainble land management.
World Heritage Convention (WHC) 1972 193 Establishes duties of parties to identify potential World Heritage
Sites, and their role in protecting and preserving them.
International Plant Protection Convention (IPPC) 1952 183 Protects sustainable agriculture and enhances global food security
by preventing the spread of invasive pests and pathogens.
Trilateral Committee for Wildlife and Ecosystem 1995 3 Seeks cooperation between the wildlife agencies of Mexico, the
Conservation and Management United States, and Canada to conserve wildlife, biological diversity,
and ecosystems of mutual interest.
Convention on the Law of the Sea 1994 168 Establishes a comprehensive regime of law and order in the world’s
oceans and seas establishing rules governing all uses of the oceans
and their resources.
a
Many conventions also have specific protocols that form the basis for action, with their own sets of Parties and Signatories. Although many
international agreements are governed through the United Nations, others may be the result of regional cooperatives such as the Organization of
American States
b
Year the convention or treaty entered into force; each convention has a minimum number of Parties that are required before it can be enforced
c
As of November 2019. Parties are countries that have ratified the treaty or convention and make it legally binding on themselves
not enforceable (or in full effect) with signatures alone. The efforts were often ad hoc and largely uncoordinated. A turn-
next crucial step in the process is to ratify the treaty. This ing point in international conservation came in 1972, with the
requires each nation state to follow its domestic procedures convening of the United Nations Conference on the Global
for ratification, which could include approval by the legisla- Environment in Stockholm, Sweden, better known as the
ture, parliament, or Head of State. For instance, in the United Stockholm Conference.
States, a treaty can be negotiated by the President, but must
be approved by two-thirds of the Senate to be legally ratified.
In some cases, the President can negotiate an executive 12.3.2 Stockholm: The Beginnings of Modern
agreement between two heads of state, such as the well- International Conservation Law
publicized 2014 joint announcement between former
US President Barack Obama and Chinese President Xi Most international legal scholars today mark the beginnings
Jinping on their targets for reducing greenhouse gas of coordinated environmental and conservation law with the
emissions. However, these executive agreements are consid- convening of the Stockholm Conference of 1972 (DiMento
ered to be politically binding rather than legally binding. 2003:18). The expressed purpose of the Stockholm Confer-
With ratification, a signatory state expresses its consent to ence was to provide a framework within which the UN could
be bound by the treaty. comprehensively assess the problems of the human environ-
A treaty becomes legally binding internationally when it ment and place the focus of national governments and the
“enters into force.” For this to occur, each treaty has its own public on such problems. Its most significant achievement
set of conditions, as outlined in the document, that must be was the production of the Declaration on the Human Envi-
satisfied. In most cases, a treaty will enter into force when a ronment, a document containing 26 principles and
minimum number of nation states have ratified it. It is conse- 109 recommendations related to environmental protection
quently important to distinguish between the number of and conservation (Fig. 12.10). Perhaps even more impor-
signatories and the number of countries that have ratified tantly, it was at Stockholm that the UN created its first
the agreement, also referred to as “Parties.” Some countries, specifically environmental agency, the aforementioned
notably the United States, will sign treaties in an expression United Nations Environmental Programme. UNEP was
of support from the executive branch, the chief diplomatic charged with the responsibility for creating new international
arm of the government. But when it comes to making the conventions to foster conservation and protect the environ-
country subject to international oversight through ratification, ment, as well as for their enforcement.
some Senators consider the treaty to be a potential intrusion The Stockholm Conference was significant in that the
on their country’s sovereignty and ultimately do not vote to United Nations became involved in world conservation in
ratify. The effectiveness and legitimacy of international envi- comprehensive and systematic ways, something it had rarely
ronmental agreements has long been a subject of debate.
Legal scholars Andresen and Hey suggest that “given the
resources they contribute to addressing international environ-
mental concerns,” so-called developed countries, “tend to be
most concerned with the effectiveness” of international gov-
ernance regimes (2005). In contrast, developing countries,
feel they have “a limited say [in negotiation processes], and
have tended to be more concerned with the legitimacy of
these same regimes (Andresen and Hey 2005).
Despite these challenges, the UNEP and other UN
programs such as the United Nations Development
Programme (UNDP), have directly stimulated the develop-
ment of new international organizations and encouraged
global conservation priorities. This work has also led to the
development of international conservation agreements
among nations in the same region, serving as a catalyst for
more coordinated regional action for environmental conser- Fig. 12.10 Mr. Maurice F. Strong, Secretary-General of the United
vation. For example, the Carpathian Convention, a subre- Nations Conference on the Human Environment (right), shows United
gional treaty across seven European states within the Nations Secretary-General U Thant a design for the official Conference
poster. To the left is Mr. Keith Johnson (Jamaica), Chairman of the
Carpathian mountain range, provides a common vision for Preparatory Committee for the Conference. (Photo taken by Teddy Chen
protection and sustainable development of the region (UNEP on 15 September 1971 at the United Nations Headquarters, New York.
2019). During the first two decades of its existence, such UN Reprinted by permission from UN Photo)
done before. UNEP made environmental concerns and the most important international conservation agreements
programs a permanent fixture of the United Nations agenda. operating today because it specifically regulates or prohibits
For the first time, a global institution created a series of global commercial trade in globally endangered species or their
programs designed to address environmental and conserva- products. Most wildlife trade occurs at local and national
tion concerns. Although nations in the global south did not levels, but large volumes of international trade also take
play a major part in establishing this institution, “the fact that place each year, worth an estimated USD20 billion a year
the UNEP’s headquarters were located in Nairobi [Kenya] globally (Challender et al. 2015).
strongly contributed in it being perceived as a more legiti- The origins of CITES begin in 1950 with a tiny grant
mate institution on the part of the developing states” (USD$2500) from the United Nations Educational, Scien-
(Andresen and Hey 2005). tific, and Cultural Organization (UNESCO) to the Interna-
The creation of UNEP had an almost immediate impact on tional Union for Conservation of Nature (IUCN). IUCN used
world conservation. In 1973, just one year after Stockholm, these funds to begin a program it called the Survival Service,
the UNEP Governing Board declared regional seas to be an a unit within the organization that began making lists and
important conservation priority. This emphasis led directly to short status reports of endangered and threatened species
the development of the Barcelona Convention of 1976 for the worldwide (Chap. 1). These reports, which by the 1960s
Protection of the Mediterranean Sea Against Pollution, an were known as the famous “Red Data Books,” ultimately
agreement developed by Mediterranean nations that sought to became the most important source and most respected author-
reduce pollution and preserve native Mediterranean Sea spe- ity for identifying the world’s endangered species and their
cies (DiMento 2003:28). The Barcelona Convention status. Using relatively simple decision rules as criteria, the
provided the incentive and model for regional environmental Red List categorizes species according to their relative
and conservation treaties that would follow during the next endangerment (Chap. 2). By drawing the attention of the
30 years, with regional treaties developed for most of the international community to the plight of endangered species,
world’s oceans from 1972 to 1986. By 1988, more than IUCN’s reports began to spur debate in the UN. As early as
100 nations and 50 international organizations were 1963, the directors of UNEP called for “an international
cooperating in regional seas programs (Sand 1988) and the Convention on regulations of export, transit, and import of
number has continued to increase. In addition to regional rare or threatened wildlife species or their skins or trophies”
conventions and protocols, the 1982 Montego Bay Conven- (Holdgate 1999:114). Various drafts circulated from 1964 to
tion, developed in association with the Third UN Conference 1972, but none could bring consensus. Finally, in 1973, an
on the Law of the Sea, addressed major issues of ocean intergovernmental negotiating conference was convened in
conservation on a worldwide basis. As such conventions Washington, DC. A formal agreement was reached, and the
have developed, the most important trend has been a shift text of CITES was prepared and circulated in three languages
from use-oriented to resource-oriented approaches. The (Holdgate 1999:115). In fact, the 1973 Endangered Species
use-oriented approach emphasized navigation and fishing. Act was intended to provide the US with legal authority to
The resource-oriented approach emphasizes sustainable implement CITES.
development and harvest of ocean resources, focusing on Support for CITES built steadily, and the Convention
defining and enforcing standards of “protection,” “conserva- entered into force in 1975. In the Preamble to the Convention,
tion,” “management,” and “development” (Sand 1988). the “Contracting States” or the “Parties” outline their broad
motivation before presenting the specifics of their
approach (993 U.N.T.S. 243):
12.3.3 Protection of Endangered Species: The
Convention on International Trade Recognizing that wild fauna and flora in their many beautiful and
varied forms are an irreplaceable part of the natural systems
in Endangered Species of Wild Fauna of the earth which must be protected for this and the
and Flora (CITES) generations to come;
Conscious of the ever-growing value of wild fauna and flora from
Although the programs, treaties, and conventions that devel- aesthetic, scientific, cultural, recreational and economic
points of view;
oped out of the Stockholm Conference were critical to world
Recognizing that peoples and States are and should be the best
conservation, most did not deal directly with the problems of protectors of their own wild fauna and flora;
endangered species or the preservation of world biodiversity. Recognizing, in addition, that international co-operation is essen-
One of the most important international agreements on this tial for the protection of certain species of wild fauna and flora
against over-exploitation through international trade;
issue, The Convention on International Trade in Endangered
Convinced of the urgency of taking appropriate measures to this
Species of Wild Flora and Fauna of 1973 (CITES), arose end; Have agreed as follows. . .
from the combined efforts of the International Union for the
Conservation of Nature (IUCN), and UNEP. CITES is one of
The heart of the CITES treaty is then described via three previously sold on the world market as raw material for
appendices that list three categories of species regulated medicines, aphrodisiacs and, in some Middle Eastern
under the terms of the treaty. Appendix I lists species that countries, as handles for ceremonial daggers. Although the
are endangered and vulnerable to existing or potential trade. original mandate was justified as a means to eliminate
Commercial trade in Appendix I species is prohibited and incentives for national governments to trade in rhino horn
permits from both the importing and exporting country must products and thus discourage poaching, more recent delegate
be obtained even for non-commercial transport. Appendix II opinion was that destruction of stockpiles would cause the
species are those that either could be threatened by large price of rhino horn to increase, escalating poaching pressure
volumes of trade or that cannot be distinguished from a (Kelso 1995). Practically, this repeal meant that governments
threatened species. Trade involving species in these must now “identify, mark, and secure” their rhino horns in
categories requires a permit from the exporting country. national stockpiles that have, ironically, grown because of
Appendix III species are not globally endangered but may increasingly effective enforcement of conservation laws,
be listed at the initiative of an individual state seeking inter- leading to seizures of rhino horns taken by poachers.
national cooperation for that species’ protection. For Appen- Since these changes to policy in the 1990s, South Africa
dix III species, nations are asked not to permit importation of has witnessed a second wave of illegal poaching.
the species without an export permit from the listing country. South Africa is home to more than 90% of the world’s
As of January 2017, there were 35,801 species and 68 sub- 20,000 white rhino, and 40% (more than 80% together with
species in the Appendices comprising 1045 in Appendix I its neighbor Namibia), of the 5000 remaining black rhino
(3%), 34,608 in Appendix II (97%) and 216 in Appendix III (Diceros bicornis) (Emslie 2012). Poaching in South Africa
(<1%) respectively (CITES 2017). has, on average, more than doubled each year from 2008 to
Parties to the treaty meet every two years to make 2012 (Biggs et al. 2013, Fig. 12.11). The rapid rise in
amendments to the appendices and develop new species poaching levels appear to be driven by tremendous growth
and animal product lists and identification manuals to in the retail price of rhino horn, from around $4700 per
improve enforcement (Slocombe 1989). CITES has proved kilogram in 1993 (Loh and Loh 1994) to around $65,000
to be an evolving document, and amendments to original per kilogram in 2012 (Marshall 2012).
provisions are not uncommon, reflecting changes in The South African government has investigated a range of
perceptions among delegates about the best way to achieve potential policy options to combat this activity, including
conservation of wildlife. There has always been disagree- possible submission of a proposal to the 2016 CITES Con-
ment, fueled in part by the treaty’s own ambiguous language, ference of Parties to re-establish an international commercial
as to whether CITES is an instrument for wildlife protection trade in rhino horn, supplied from existing legal stockpiles
or a means to regulate wildlife trade. Although originally a and established private breeding operations that practice rou-
treaty that equated conservation with strict protectionism in tine dehorning (Sas-Rolfes 2017). However, this proposal
international trade, more recent meetings of CITES received criticism domestically, especially from private
participants have shown a growing tendency to permit some rhino owners, and as of 2019 has not been officially pursued.
trade in formerly protected species if it can be shown that Although the 1994 meeting of CITES delegates did not
such trade actually enhances species conservation. Thus, actually approve the sale or trade of horns from stockpiles,
attempts to apply CITES to specific conservation dilemmas it paved the way to do so at a later time, under strict controls,
have proven problematic. if current inventory can be carefully marked.
For example, in November 1994, CITES delegates agreed
to allow trade in live southern white rhinos (Ceratotherium
simum) from the Republic of South Africa, an action based
Some economists and biologists have argued for a legal
primarily on the success of rhino conservation programs in
trade in rhino horn in part because all current income
that country that had restored a population of 20 individuals
from illegal trade practice is going to criminals and
(all that remained in the country by 1920) to about 6300, the
none is invested back into conservation efforts. Why
largest national population in Africa (Kelso 1995). Sales of
might others be hesitant about legalizing the trade?
white rhino were actually expected to improve the status of
the species in South Africa because proceeds would be spent
on further rhino conservation efforts. The rhinos sold to other
governments were expected to aid in restoring rhino Outside South Africa, other countries such as the United
populations currently in decline in other countries. Through States have tried to reduce wildlife poaching through govern-
this effort, the country’s southern white rhino population was mental action. In April 1994, the Clinton administration of
down-listed from CITES Appendix I to Appendix II. the US government imposed trade sanctions on Taiwan after
At this same meeting, CITES delegates also repealed the its government failed to curtail trade in rhinoceros and tiger
1987 mandate to destroy existing stockpiles of rhino horns, parts despite warnings from the United States. Some scholars
mark this action as the first time in history that international the new rules will go a long way in helping law enforcement
trade sanctions had been used directly to protect wildlife more easily distinguish legal from illegal ivory” (Actman
(Coggins 2003:5). More recently, in 2016, the Obama admin- 2016).
istration proposed a near-total ban on the commercial trade of
African elephant ivory. The law had previously allowed for
the sale of ivory and ivory products in limited cases where the 12.3.4 Combining Conservation
seller could prove the ivory was old and lawfully imported. and Development in International
However, the new rules further restricted exports and sales Agreements
across state lines, as well as limit ivory trophy imports to two
per year per hunter. The US Fish and Wildlife Service, the In June 1992, a major global environmental gathering
agency charged with the enforcement of CITES within occurred in Rio De Janiero, Brazil, to integrate efforts to
US borders, argued that “wildlife traffickers have exploited protect planetary ecosystems with economic development
previous regulations allowing for a legal trade in ivory and of the poor nations of the world. The United Nations
Fig. 12.11 The current distribution of white rhinoceros by permission from the IUCN from The IUCN Red List of Threatened
(Ceratotherium simum) populations in Africa, including areas where Species. Version 2019-2. https://www.iucnredlist.org/species/4185/
the rhinoceros is now extinct, possibly extinct, and extant due to either 16980466; Graph reprinted by permission from the American Associa-
introduction or re-introduction (a). As an indication of the threat tion for the Advancement of Science from Biggs, D., Courchamp, F.,
poaching poses to white rhino populations, the number of white rhinos Martin, R., Possingham, H.P., 2013. Legal Trade of Africa’s Rhino
poached in South Africa more than doubled each year over a ten year Horns. Science 339, 1038–1039. https://doi.org/10.1126/science.
period, reaching 668 individuals in 2012 (b). (Distribution map reprinted 1229998, 2017 # The Authors)
Fig. 12.11 (continued)
Conference on Environment and Development (UNCED), protection of land resources, halting deforestation, conserv-
popularly referred to as the Rio Summit or Earth Summit, ing biodiversity and protecting freshwater and saltwater
was a formal conference of official government delegations. resources, and (2) the problems of human industry and tech-
Simultaneously, a large gathering of non-governmental nology that pose particular threats to the environment, includ-
organizations gathered for the Global Forum, a mixture of ing threats posed from biotechnology, toxic, hazardous
NGO networking, street shows, trade fairs, and environmen- wastes, sewage and agriculture (Sands 1999:130–131).
tal demonstrations (Parson et al. 1992). The result of this In its social and economic dimensions, Agenda 21
convening was five major environmental documents, signed affirmed the need to eradicate poverty and hunger, to manage
by most or, in some cases, all of the participating nations. The resources sustainably, to link human health to environmental
best known of these is the Rio Declaration, originally and socioeconomic improvements, and to integrate environ-
conceived as a kind of “Earth Charter” that contained mental factors into policymaking, law, economics and
27 principles for sustainable development (more on the national accounting. In addressing conservation and manage-
origins of this term in Chap. 11). The Rio Declaration, signed ment of resources for development, Agenda 21 supported
by all participating nations, affirms environmental protection allocation of land that provided the greatest sustainable
as an integral part of development. benefits, affirmed the need for worldwide conservation of
The most comprehensive document signed at the Rio biodiversity, the need for proper management of mountain
Summit was Agenda 21, an 800-page “work plan” addressing resources, more information on mountain ecosystems, and
social and economic dimensions of environment and devel- integrated development of mountain watersheds. In this sec-
opment, conservation and management of resources, and tion, Agenda 21 also affirmed the importance of freshwater
means of implementation. Agenda 21’s structure was based resources, provision of safe drinking water, and the need for
on key environmental and conservation issues, including the safe management of various kinds of toxic chemicals and
problems of desertification, protecting the atmosphere, and hazardous wastes.
managing toxic wastes. It also addressed social issues with In its final section on implementation, Agenda 21
environmental dimensions such as poverty and technology supported promoting public awareness, establishing a new
transfer (Greene 1994). Overall, Agenda 21 identified prior- UN body, the Commission on Sustainable Development
ity environmental issues and divided them into two (CSD), to coordinate pursuit of sustainable development
categories: (1) the priority needs for environmental protec- among international organizations and monitor progress by
tion, including atmospheric protection and climate change, governments and international organizations toward reaching
Fig. 12.12 The eight Millennium Development Goals (2000–2015) contrasted with the 17 Sustainable Development Goals (2016–2030). The
Sustainable Development Goals each include a series of specific targets with measurable data indicators to track progress relative to each target.
(Graphics courtesy of the United Nations)
the goals set out in the Agenda. It concluded with a discus- these limitations, conservation targets outlined in the SDGs
sion of the importance of collecting and using information for and other global agreements, have spurred new partnerships,
sustainable development and for implementing Agenda raised awareness and promoted investment (UN 2015, 2018;
21 (Parson et al. 1992). Doherty et al. 2018).
Over the last couple of decades, the implementation of In addition to the Rio Declaration and Agenda 21
Agenda 21 has subsequently resulted in the establishment of documents, two major conventions were opened for signature
several high-profile global goals organized as the Millennium at the Rio Summit. One of these, the Framework Convention
Development Goals (MDGs) (2000–2015) and the Sustain- on Climate Change (UNFCCC) primarily addresses
able Development Goals (SDGs) (2016–2030) (Fig. 12.12). emissions limits and standards of “greenhouse gases”
Although an integrated human-nature paradigm is being associated with fossil fuels. Although the convention does
practiced by disciplines like conservation biology, the imple- not set specific targets, its ambitious objective was the “sta-
mentation of integrated social-ecological decision-making bilization of greenhouse gas concentrations in the atmosphere
through international law and policy is harder to achieve that would prevent dangerous anthropogenic interference
(Anderson et al. 2019). There are currently 17 SDGs, with with the climate system. . . .within a time frame sufficient to
at least four being overtly environmental in focus and several allow ecosystems to adapt naturally” (UNFCCC 1992). This
others having strong connections between nature and people Convention, with 197 Parties as of August 2018, has resulted
from a systems view. Each of these broad goals has a series of in the subsequent negotiation of the Kyoto Protocol (1997)
targets and a supporting set of indicators to measure success and Paris Agreement (2015) (Chap. 4).
relative to these targets. However, as Doherty et al. note, “to The other treaty opened for signature in Rio was the
account for the competing political, socioeconomic, and Convention on Biological Diversity (CBD), which addressed
environmental interests that are common in conservation conservation and sustainable use of biodiversity along with
policy, compromises are typically made during target devel- fair and equitable sharing of benefits arising from use of
opment and implementation, which can weaken their primary genetic resources. The signatories pledged to develop plans
objectives” (2018). Furthermore, with limited regulatory and to protect habitats and species, provide funds and technology
enforcement power, countries face few to no legal to assist developing countries to provide protection, ensure
consequences for failing to meet identified targets. Despite commercial access to biological resources for development,
Fig. 12.13 The response

hierarchy to Land Degradation
Neutrality (LDN) encourages
broad adoption of measures to
avoid and reduce land
degradation, combined with
localized action to reverse
degradation, to achieve LDN
across each land type. (Reprinted
by permission from Elsevier, from
Cowie, A.L., Orr, B.J., Castillo
Sanchez, V.M., Chasek, P.,
Crossman, N.D., Erlewein, A.,
Louwagie, G., Maron, M.,
Metternicht, G.I., Minelli, S.,
Tengberg, A.E., Walter, S.,
Welton, S., 2018. Land in
balance: The scientific conceptual
framework for Land Degradation
Neutrality. Environmental
Science & Policy 79, 25–35.
# 2017 The Authors)
share revenues fairly among sources and developers, estab- of differences between industrialized countries that wanted a
lish safety regulations, and accept liability for risks associated treaty focusing on tropical forests and developing countries
with biotechnology development (Parson et al. 1992). Enter- that wanted a treaty including boreal and temperate forests.
ing into force only 18 months after it was introduced, the Over the next decade, forest concerns were integrated into the
Convention currently has 196 Parties, with subsequent work of the Commission on Sustainable Development, UN
negotiations of the Cartagena Protocol (governing the Food and Agriculture Organization’s Committee of Forestry
movements of living modified organisms (LMOs) resulting and the UNFCCC, CBD, and UNCCD. However, some
from modern biotechnology) and Nagoya Protocol argue that even with potential synergies among these efforts,
(a transparent legal framework for the effective implementa- there remains a need for a single treaty which covers all the
tion of one of the three objectives of the CBD: the fair and gaps in regulation and ensures sustainable conservation and
equitable sharing of benefits arising out of the utilization of development of the world’s forests.
genetic resources).
Although not open for signature until 1993, The Conven-
tion to Combat Desertification (UNCCD) was established as 12.3.5 The Process of Creating and Enforcing
a direct result of the Rio Summit, intending to slow the International Conservation Law
process of desertification by adopting measures to protect
dryland environments and improve the living standards of If international conservation law consisted merely of value-
people who use them through enhanced livestock and for- neutral rules, its most important element would be hard law,
estry practices, land use reform, soil and water conservation, formal conventions and treaties adopted by many nations,
and wildlife protection. To date, 197 countries have ratified with explicit mechanisms for enforcement. However, the
this convention and have submitted plans on how they plan to actual behavior of the modern international community has
combat desertification within their own borders. The Parties demonstrated the growing importance of soft law,
recently launched their 2018–2030 Strategic Framework to non-binding agreements that, although having no official
achieve global Land Degradation Neutrality in order to means of enforcement, eventually come to define the norms
restore the productivity of vast expanses of degraded land and standards for international behavior. The reality of this
and improve the livelihoods of more than 1.3 billion people concept can be seen in the way in which international laws on
(UNCCD 2017; Fig. 12.13). environmental conservation actually come into being.
The final outcome of Rio was a non-binding declaration, In 1977, UNEP established a Working Group of Experts
the Statement on Forest Principles, under which its signers on Environmental Law, whose recommendations were
pledged to keep 17 principles “under assessment for their endorsed by the UNEP governing council and, in 1982, by
adequacy with regard to further international cooperation on the UN General Assembly. Although individual nations were
forest issues” (Parson et al. 1992). Progress toward a formal not legally bound to use of these guidelines, much of the
treaty on forests at the Rio Summit failed primarily because so-called “soft law” recommendations from this panel of
experts and other sources has become, over time, an increas- smaller the number of participants involved in the activity,
ingly recognized international standard (Sand 1988). Such the easier the activity is to regulate internationally. Likewise,
soft law agreements are often the sources for developing the participants in an activity that deals with large, global
actual wording of “hard law” agreements in more formal markets are also easier to regulate than participants in smaller
conventions, and generally create a climate of compliance firms and more local markets because global corporations
by establishing a normative standard that makes them as and businesses are far more concerned about international
effective as hard law. As legal scholar Jane Roberts observed, image.
these agreements often create such a spirit of shared values According to this model, the most important characteristic
and goals that in terms of international behavior, they “have a of the treaty or convention itself is equitability. Agreements
predictive value similar to those expressed in hard law” perceived by all parties to provide for fair treatment had much
(Roberts 2004:103). The Rio Declaration, Agenda 21, the higher compliance than those that were perceived to favor
Millennium Development Goals, and the Sustainable Devel- some participants over others. International reception also
opment Goals are all examples of soft law. plays an important role in compliance. The more persistently
Even soft laws in conservation must have a catalyst. and publicly the international community focuses on a con-
Although every international convention, treaty, or protocol servation problem, the more compliance increases with inter-
is a product of unique circumstances, the development of national conservation agreements related to that problem. In
international instruments in conservation usually follows a addition, the clear support of a “leader” country or group of
four-step process: (1) issue definition; (2) fact finding; (3) cre- countries, such as the United States or the European Union,
ation of an international body or regime to address the prob- for a particular accord also is a critical factor in the level of
lem; and (4) consolidation and strengthening the regime. compliance. Where such leadership is present, international
For increased clarity and understanding, certain concepts compliance is high.
repeatedly invoked in international conservation law require Compliance is affected by both intent and capacity. Intent,
careful definition. Once conceived and defined, laws must or political will, can be judged from the behavior of national
find a mechanism of implementation, that is, nations must leaders and political bodies, and is a necessary but insuffi-
take specific actions to make international treaties operational cient condition for compliance. With intent, the country also
in their own national legal system. The purpose of creating must possess the capacity to comply, requiring an efficient
mechanisms of implementation is to increase compliance, and honest environmental bureaucracy, economic resources,
that is, to increase the extent to which the behavior of a technical expertise and public support. Weiss and Jacobson
state, as a party to an international treaty, actually conforms have suggested three strategies for strengthening interna-
to the conditions of the treaty (Faure and Lefevere 1999: tional compliance. The first of these is the sunshine
139). Methods used to force states to first implement and approach which focuses on mechanisms to bring the behav-
then comply with international agreements are mechanisms ior of key parties into the open for public scrutiny, including
of enforcement, and vary with individual agreements and such actions as regular reporting, peer scrutiny, on site moni-
conditions. The goal of such enforcement is ultimately effec- toring and media access and coverage. In this area, NGOs in
tiveness, a measure, not simply of whether the nation lives up conservation often play a critical role. In countries where
to the conditions of the treaty, but of whether such behavior NGOs are active in publicizing examples of
actually achieves the objectives stated in the treaty. Thus, an non-compliance, the more likely they are to strengthen their
ideal international conservation agreement is one in which government’s intention to comply. Echoing Weiss and
there are clear and feasible mechanisms of implementation, Jacobson’s conclusions, international legal scholars Michael
high levels of compliance, and workable methods of enforce- Faure and Jürgen Lefevere note that “the stronger and more
ment, all leading to accomplishing the goals for which the active NGOs are with respect to the issue area of the treaty,
agreement was formed in the first place (high effectiveness). the larger the probability of compliance” (Faure and Lefevere
Regrettably, not every international conservation treaty or 1999:138). And, the more actively both government and
convention gains high marks in all areas. NGOs are engaged in reporting information relevant to the
Weiss and Jacobson (1999) developed a conceptual agreement, the more compliance increases.
model, based on the actual success of a variety of interna- Secondly, a pattern of reporting behavior instigates its
tional environmental agreements, to show how various own reinforcement because it encourages the development
factors affect implementation, compliance and effectiveness of compliance information systems that are built into gov-
of international conservation treaties (Fig. 12.14). Compli- ernment structures, systems whose aim is to ensure compli-
ance with international treaties is affected by the ance and report non-compliance (Faure and Lefevere
characteristics of the activity (for example, numbers of 1999:143). At the international level, one coordinating body
participants, characteristics of markets, location of the activ- for such compliance information systems is The Global
ity), characteristics of the agreement, and the state of the Environmental Facility (GEF), established by The World
international environment. A general trend has been that the Bank in 1991 in cooperation with UNEP and the UNDP.
Fig. 12.14 A model of factors that affect implementation, compliance Strengthening Compliance with International Environmental
and effectiveness of international treaties and conventions in conserva- Accords, MIT Press, Cambridge. # 1998 Massachusetts Institute of
tion. (Reprinted by permission from The MIT Press, from Weiss, Edith Technology)
Brown, and Harold K. Jacobson, eds. Engaging Countries:
To encourage compliance, GEF provides funding for the landscapes and seascapes and ensuring that the impact of
implementation of treaties that target various aspects of envi- productive sectors on biodiversity is avoided, or substantially
ronmental quality and conservation. In particular, the GEF reduced or minimized; (2) enhancing the effectiveness and
makes funds available to developing countries and countries sustainability of protected area systems; (3) supporting the
with economies in transition in an effort to aid compliance. complete and effective implementation of the Cartagena and
For example, with respect to the implementation of the Con- Nagoya Protocols; and (4) improving biodiversity policy,
vention on Biological Diversity, the GEF is making planning, and review.
investments in countries that can demonstrate improvements Finally, positive incentives work where a country has
in (1) sustainably managing biodiversity in productive compliance intention but not capacity. Here, inputs of
12.4 The Challenge of Interdependence on a Global Stage 521
money, technical expertise, capital, training, or special and work best when they are synergistic in design and appli-
considerations from other countries can increase compliance. cation. Although the worldwide trend in response to conser-
UNESCO has instituted a number of programs to provide vation treaties and conventions has been one of increasing
such incentives toward compliance, such as the World Heri- compliance, factors affecting compliance are complex and
tage List (WHL) of sites of cultural and natural heritage. national responses to international conservation efforts are
Administered by the World Heritage Centre in Paris, France, not uniform.
the WHL, a program created by the Convention Concerning Increasingly, conservation efforts at international levels
the Protection of the World Cultural and Natural Heritage of are guided, as well as constrained, by two overriding
1972, is designed to identify and protect sites of outstanding principles that often pull in opposite directions. The first is
cultural and natural value in every nation. In this case, the the increasing awareness and consensus that every nation has
positive incentives take the form of providing help with a responsibility to conserve its natural resources and must not
administrative oversight, technical expertise, financial and damage them for use by future generations. This first princi-
material resources, and international influence for the desig- ple is rooted in an axiom that has become even more founda-
nation and protection of listed sites. As of 2019, the WHL has tional to international conservation: the commitment to
recognized (technically, “inscribed”) 1092 individual sites intergenerational equity, which is itself supported by three
identified for their particular cultural or natural value core ideas. First, each generation should be required to con-
(UNESCO 2019). Although this designation, and the serve the natural and cultural resource base of its own nation
accompanying support can increase capacity, there are so that it does not restrict the options available to future
always challenges on ongoing protection. There are 57 sites generations in addressing their problems and achieving their
currently listed in danger, due to ongoing threats to their goals. Second, each generation should maintain their envi-
protection ranging from internal conflict war, rapid popula- ronmental quality in such a state that it is in no worse
tion growth, and lack of domestic legal protection. condition than that which they received. Finally, members
If the above mechanisms fail, coercive measures can be of every generation have comparable rights of access to the
effective against parties that have capacity to comply but lack legacy of past generations and should conserve this access for
intention. Sanctions, penalties, loss of membership in inter- future generations (Weiss 1999). These are not empty
national organizations or of privileges in international platitudes, but rather foundational principles that increasingly
dealings can be effective in motivating unwilling parties to influence how international law is expressed and
comply with agreements (Weiss and Jacobson 1999). Coer- implemented. For example, in the Philippines, the Supreme
cive measures are often most clear and most effective at the Court recognized intergenerational equality by granting con-
domestic level, such as the presence of a heavy fine or stitutional standing to a group of children to represent the
criminal charge for an illegal “take” of an endangered species interests of future generations in their efforts to stop the
under the US Endangered Species Act. It can be harder to leasing of biologically diverse forests for development
implement at international levels and is one reason why many (Axelrod et al. 2011).
scholars view international treaties as “morally-binding” in The second foundational principle guiding international
terms of compliance, since countries themselves are often the conservation is that every nation has sovereign rights over its
ones reporting their own state of compliance or lack-there-of own national resources, and these rights are not to be
(Samaan 2011). In response to this challenge, there is increas- infringed by other nations (see also Principle 2 of the Rio
ing desire for a neutral, independent organization, one that Declaration). Thus, competing claims of responsible conser-
could act under its own authority without answering to any vation stewardship for the sake of intergenerational equity
specific government, to encourage uniform compliance, and and national resource sovereignty by different nations can
establish a clear mechanism for reprisal in the case of create problems. In a world of increasing global connection
violations (Samaan 2011). and dependence, any nation taking unilateral initiatives for
global conservation must consider the effects of such
initiatives upon other nations in order to create positive
12.4 The Challenge of Interdependence outcomes. Environmental policy scholar Edith Brown
on a Global Stage Weiss has noted, “In international environmental law, the
most important development for the next century may be
12.4.1 The Nature of Legal Interdependence the emerging interaction of intergovernmental environmental
Among Nation-States law with transnational law. . .” (Weiss 1999:102). Further,
Weiss perceives that “International law has always been
Both international conservation law and the national laws linked with national law, for it is implemented through
of individual states have increased in breadth and matured national, provincial, and local laws. . . national laws, inde-
in application over the last five decades. National laws and pendent of any treaty, provide protection to other countries or
international conventions are often aimed at the same goal their citizens for harm that occurs within the country but
injures those outside” (Weiss 1999:104). To better under-

stand and appreciate the fascinating complexity of and
connections between the claims of environmental protection
and national sovereignty, national and international conser-
vation law, and governments and non-governmental
organizations, we consider the following examples of legis-
lation designed to protect dolphins from tuna fishermen, sea
turtles from shrimp trawlers, and Brazilian biodiversity and
indigenous communities from global companies.
12.4.2 Case History I: Tuna and Dolphins
In 1972, just two years after passage of NEPA and only a year
before passage of the amended Endangered Species Act, the
US Congress enacted the Marine Mammal Protection Act Fig. 12.15 The Dall’s porpoise (Phocoenides dalli) is one of many
(MMPA). The MMPA was a relatively minor and species of porpoises and dolphins often killed as “bycatch” (Chap. 8) in
non-controversial piece of legislation that enjoyed broad the process of tuna fishing. The “incidental kill” of dolphins in associa-
bipartisan support. The Act’s clear and simple goal was to tion with tuna fishing has resulted in the deaths of millions of dolphins
worldwide. (Photo courtesy of Texas Parks & Wildlife Department)
protect “certain species and population stocks of marine
mammals that are, or may be, in danger of extinction or
depletion as a result of man’s activities.” One of the the MMPA was altered to require an embargo on tuna imports
MMPA’s mechanisms to achieve this goal was to reduce from any country whose commercial fleets killed more
“incidental kill or serious injury of marine mammals.. . .to dolphins than US fleets. In 1988, Congress added additional
insignificant levels approaching a zero mortality and serious requirements for all tuna-exporting nations attempting to mar-
injury rate.” ket tuna in the United States. Tuna-exporting countries were
The deaths of marine mammals associated with “inciden- required to reduce incidental kill of non-tuna species to the
tal kill” had increasingly become a cause for scandal and level of US fishing fleets, and were prohibited from using
condemnation by the public and the press, particularly large-scale drift nets, encircling marine mammals without
regarding the killing of dolphins by tuna fishers. The problem direct evidence of the presence of tuna, or using purse-seine
had been developing since the 1950s, when tuna fishers nets after sundown. The amendment also specified that failure
began to employ purse-seine nets for capturing tuna. Such to comply would lead the US Secretary of Commerce to ban
nets captured tuna in large schools when they fed near the imports of tuna from countries violating the regulation as well
surface. After tuna were surrounded by the purse-seine net, as from countries to which the offending nations sold tuna
the bottom of the net was pulled together, trapping the tuna (to prevent trans-national shipments as a way of getting
and all other organisms inside (Joyner and Tyler 2000). around the regulation). In effect, this placed a US embargo
Dolphins often travel directly above schools of tuna, so on tuna products of the offending nation, not only in its direct
tuna fishers began to track dolphins as indicators of tuna dealings with the United States, but indirectly throughout the
presence. Thus, it was not surprising, or even “incidental,” world (Miller and Croston 1998; Joyner and Tyler 2000;
that dolphins were killed with tuna, either by drowning in the Salzman and Thompson 2013).
net or being crushed by the harvesting machinery. Since the In 2016, the US increased pressure against violators
1960s, an estimated 6 million dolphins have perished in this by creating stricter standards for any company placing a
manner (NOAA 2016) (Fig. 12.15). “dolphin safe label” on its tuna products. The rules required
By the late 1980s, US environmental and conservation fishing boat captains to complete training and certifications in
NGOs persuaded the US Congress to add an amendment fishing techniques that would ensure that dolphins would not
to the MMPA which established stringent guidelines for all be killed as tuna bycatch, and enhanced chain of custody
tuna fishing in the US waters to assure protection for tracking requirements for tuna and tuna products (NOAA
dolphins and other species. It soon became apparent, how- 2016). Although the “dolphin safe label” had been around
ever, that other countries, including those harvesting the for years, these new restrictions were the result of an ongoing
majority of tuna, were not following standards set by the trade dispute with Mexico, which had long complained (suc-
MMPA. To encourage adoption of such standards on an cessfully; see below) to the World Trade Organization that
international level and to protect dolphin populations world- US rules disporportionately discriminated against its tuna
wide, the US Congress twice amended the MMPA. In 1984, fishers. The World Trade Organization sided with Mexico
Fig. 12.16 A Turtle Excluder

Device (TED) that can be installed
in a shrimp net to release sea
turtles from the net. TEDs,
properly installed, can reduce
sea-turtle mortality associated
with shrimp fishing by up to 97%.
(Figure courtesy of the Marine
Research Foundation)
and ordered the US to make changes. The US responded by turtles that could delay or halt ratification of GATT, and
elevating restrictions for every other country in the world US officials delayed enforcement of the Act against its
(Actman 2016). most important shrimp suppliers. Such reticence eventually
led to a federal lawsuit by the Earth Island Institute, a
US NGO. Earth Island Institute demanded that the provisions
12.4.3 Case History II: Shrimp and Sea Turtles of the Sea Turtle Act be enforced uniformly against all
nations exporting shrimp to the US. After a series of appeals,
In 1989, the US Congress added a provision (Section 609) the Earth Island Institute won the case in the US Court of
to Public Law 101–162 that became known as the “Sea Turtle International Trade, forcing the US to ban imports from
Act” (Joyner and Tyler 2000). The Sea Turtle Act was nations that had not complied with the Sea Turtle Act,
motivated by concern over worldwide declines in the including the largest Asian shrimp exporters, and effectively
populations of all seven species of sea turtles and by scientific creating a worldwide shrimp embargo by the US against
studies that implicated nets used in shrimp fishing in sea non-compliant nations.
turtle mortality. One of the world’s largest consumers of The tuna and shrimp embargoes, now in full force, led to
shrimp, the US also was one of the first nations to employ legal challenges by the sanctioned nations before the World
the turtle excluder device (TED). A TED is a grid trapdoor Trade Organization (WTO). In separate but similar cases, the
installed inside a trawling net that keeps shrimp in the net but tuna and shrimp-exporting nations argued that the MMPA
directs other, larger objects or animals out (Fig. 12.16). By and Sea Turtle Act were violations of the free trade
the 1980s, TED technology had reached the point that, prop- provisions guaranteed by GATT. The European Community
erly installed, 97% of sea turtles caught in shrimp nets could joined in challenging the MMPA because the embargoes it
be released alive and unharmed without loss of shrimp functionally created prevented them from selling to the
(Joyner and Tyler 2000). US tuna that they had purchased from Asian nations that
Earlier legislation had already required TEDs for all did not comply with the MMPA. The plaintiffs argued that,
shrimp trawlers operating in the Gulf of Mexico and in the under the terms of GATT, an individual nation could not
Atlantic Ocean off the southeast coast of the United States. impose restrictions on imports from other nations, even for
The Sea Turtle Act went further. It prohibited fish imports conservation reasons, that those nations had not been party to
from any nation that failed to adopt sea turtle conservation developing. Further, the US could not impose sanctions
measures comparable to those in the United States. Initially based on the processing and production of a product, but
such sanctions were applied only to western Atlantic and only on the product itself. That is, what mattered was the tuna
Caribbean nations, which eventually complied. However, in the can, not how the tuna got in the can (Salzman and
the largest shrimp importers to the US were Asian nations Thompson 2013). In addition, the bottle-nosed dolphin
that did not use TEDs. As a result, the prohibitions of the Sea (Tursiops spp.), the principal species affected by the tuna-
Turtle Act were largely symbolic and did little to protect fishing methods in question, was not an endangered species,
turtles from shrimpers on a global scale. and therefore not subject to international protection. Finally,
As these events were taking place, the US was engaged the plaintiff nations charged that the entire embargo was only
in negotiations to ratify the General Agreement on Tariffs a ruse to protect US tuna fishers to give them an unfair
and Free Trade (GATT). The Clinton administration was competitive advantage in US markets, a form of protection-
reluctant to create controversy with Asian nations over sea ism wearing green clothing (Salzman and Thompson 2013).
The WTO ultimately agreed and ruled against the United decades, large foreign companies, particularly the pharmaceu-
States in the cases of both dolphins and sea turtles, agreeing tical industry, have sought access to these resources in order to
with the plaintiffs that US laws constituted unfair barriers to inform and bolster their own product development
free trade. The world conservation community condemned the (Neergheen-Bhujun et al. 2017). For example, more than half
GATT panel of the WTO for deliberately excluding environ- of all prescriptions filled in the United States contain at least
mental issues from consideration in its decision (Salzman and one major active compound now or once derived or patterned
Thompson 2013), and the US appealed the decisions, but its after compounds derived from biological diversity (Grifo et al.
appeals were not successful (Joyner and Tyler 2000). 1997). However, most of these so-called resources are “discov-
Although pledged to follow the rules of international law, ered” by western scientists at the expense of indigenous
the US continued its advocacy for conservation of both peoples and local communities without their approval or
marine mammals and sea turtles. In the former case, the involvement as the holders of such knowledge (Aguilar
US played a leading role in developing a new international 2001). Brazil’s efforts to combat this exploitation through
agreement, the La Jolla (California) agreement of 1992, a national legislation follows global efforts, codified by the Con-
ten-nation agreement that established a voluntary program to vention on Biological Diversity, and subsequently through the
limit dolphin mortality, as well as the Panama Declaration, an Nagoya Protocol, to protect and maintain local communities
agreement signed by 12 nations in 1995. The Panama Decla- embodying traditional lifestyles relevant for the conservation
ration went beyond the La Jolla agreement in establishing a and sustainable use of biological diversity (Fig. 12.17).
“permanent” mortality limit for dolphins along with stricter Despite its ambition, the law’s vague definition of “access
enforcement systems. The purpose of the agreements was to and shipment of genetic heritage” created frustration among
foster better methods of harvesting tuna through a voluntary members of the scientific academy, who asserted that
program of setting standards and procedures for dolphin research involving access to genetic heritage would likely
protection. Their outcome was the establishment of the Inter- generate economic benefits that could be shared (Do Amaral
national Dolphin Conservation Program. To implement the Axevedo 2005). Brazil responded with multiple attempts at
La Jolla agreement, the US Congress enacted the Interna- reform, but, due to the issue’s complexity, it was not until
tional Dolphin Conservation Act of 1992. To implement the 2015 that the new “Law on Biodiversity” (Law 13123) was
terms of the Panama Declaration and nationalize the intent of passed and the Provisional Act 2186-16 revoked. The Law on
the International Dolphin Conservation Program, Congress Biodiversity attempts to encompass all research and
passed the International Dolphin Conservation Act of 1997 use-related genetic information encountered anywhere within
(Miller and Croston 1998). the national territory, continental shelf, territorial sea, or
In the United States, all sea turtles are protected under the exclusive economic zone. Foreign researchers will only be
Endangered Species Act, and the US maintains its commit- able to access native biodiversity if associated with a public
ment to sea turtle conservation by continuing to sponsor or private Brazilian scientific and technological research
already existing TED certification programs for other nations. institution, which must ultimately take responsibility for reg-
In addition, the US has pledged to assist any government istering the research activity (da Silva and de Oliveira 2018).
seeking help in developing a TED sea turtle protection pro- Ethnobotanist James Welch writes that this move was
gram of its own (Joyner and Tyler 2000). In 2004, the “applauded by representatives of some important public
US negotiated with the Food and Agriculture Organization institutions in Brazil as a victory for researchers and for
(FAO) to develop guidelines to reduce sea turtle mortality in Indigenous and traditional peoples” (2015). However, Indige-
fishing operations in international waters. Because the nous and traditional peoples, with the support of many aca-
US imports more than 90 percent of its seafood, the demic societies and international organizations issued a “letter
US National Marine Fisheries Service works to make sure of repudiation arguing that they were not consulted, and their
US demand is met through sustainable and legally-obtained interests were insufficiently addressed” (Welch 2015).
sources, reporting on its progress via a federal task force Although many have come to respect international and
created for combating illegal, unreported, and unregulated domestic laws as agreements operating within and between
fishing and seafood fraud. sovereign nation states, it is equally important to respect legal
rights afforded to indigenous communities, which reside in
nation states but are often recognized as sovereign nations
12.4.4 Case History III: Brazilian Biodiversity with their own legal and governance structures and subsis-
and Genetic Resources tence rights. Arguably, efforts to protect biodiversity, and the
genetic resources it contains, should also protect the peoples
In 2001, Brazil passed Provisional Act No. 2186-16 to ensure and traditional wisdom associated with it.
the security and sustainable use of its biodiversity by regulating The 2007 United Nations Declaration on the Rights of
access to its genetic resources and derived products. For Indigenous Peoples was one recent effort to reverse the
Fig. 12.17 The Nagoya Protocol of the Convention on Biological their utilization. This is particularly important for indigenous peoples,
Diversity outlines the process by which countries can exercise sovereign who are often “providers” and entitled to rights within international
rights over genetic resources to ensure that providers of resources negotiations. (Graphic courtesy of the UN/CBD)
(or associated knowledge) receive a share of any benefit arising from
historical exclusion of Indigenous people from the interna- industry, and government bureaucracies, NGOs and indige-
tional legal system, and recognize their right to the “recogni- nous communities. They illustrate the fine line between con-
tion, observance and enforcement of treaties, agreements and servation leadership and (in the eyes of some) conservation
other constructive arrangements” (UNHR 2013; Sherpa et al. imperialism or economic protectionism disguised as conser-
2014). There has been intense debate for nearly two decades vation. Conservation laws of individual countries can no
regarding group vs. individual rights as well as issues longer be enacted or enforced without first considering the
concerning lands and resources, but countries that originally interests of other nations or the likely international response.
voted against the Declaration have since reversed their posi- For example, although trade sanctions might be justified
tion, making it a powerful tool for eliminating human rights against processing and production methods, as well as
violations against the over 370 million indigenous people products, of other countries that violate international conser-
worldwide (Sherpa et al. 2014). vation interests, an individual country cannot be confident
that it will win in the international courts unless: (1) the
measure is not unilaterally imposed and (2) the harm done
12.4.5 Outcomes and Future Prospects is local (within the jurisdiction of the country imposing the
sanctions) (Salzman and Thompson 2013).
These difficult cases of conservation conflict offer insight Today’s worldwide commitment to global free trade has
into a world of complex interactions between national and created international bodies, such as the WTO, whose
international conservation law, public interest and private decisions have the force of law. Such decisions may override
the laws passed by a single nation in matters of international

In the past, much of the activity associated with
commerce. In the tuna-dolphin and shrimp-turtle decisions,
conservation was focused on outcomes that were pre-
the WTO displayed its own preference for multilateral and
dictable effects of management actions. Goals such as
international agreements to reach conservation objectives as
sustained yield were based on an expectation of certain
opposed to unilateral, national initiatives (Joyner and Tyler
return. Today, conservationists are less concerned
2000). However, such decisions by the WTO appear to
about certainty of return than about managing risk.
sacrifice conservation to commerce, and its Dispute Settle-
Historically, environmental law has favored policies
ment Body (DSB) rarely selects panel members and experts
consistent with our past understanding of the rule of
for their environmental expertise. Although the DSB is
law (i.e., the consistent application of fixed rules that
authorized to seek expert advice on environmental issues, it
will yield a final, single decision representing an abso-
rarely does so (Miller and Croston 1998). The perception that
lute, moral ideal) (Tarlock 1994). As a result, individ-
the WTO favors trade at the expense of conservation is part of
ual environmental laws have been based on individual
the motive behind the anger and violence displayed toward
scientific premises, and have then continued the appli-
the WTO by conservation and environmental organizations,
cation of those premises regardless of what new studies
among others, in the large public, and sometimes violent,
uncovered. Today such legal certainties are inconsis-
demonstrations at the 1999 WTO meetings in Seattle,
tent with the uncertainty associated with the state of our
Washington and the 2000 WTO meetings in Washington,
knowledge of ecosystems. Conservation biologists’
D.C.
best estimates of genetic diversity, population persis-
US laws like the MMPA and the Sea Turtle Act helped
tence, and community ordination are also uncertain
provoke the international community to higher standards on
estimates. Modern conservation law and policy must
these conservation issues than would have been achieved
mature to the point that they can deal with such uncer-
without such initiatives. It is clear, however, that in an
tainty, rather than simply ignore or reject it.
increasingly global community, the US will have to improve
The development of conservation law and policy
its efforts to involve and support other nations in international
demonstrates repeated themes. First, the scrutiny of a
conservation efforts, particularly conservation efforts that
free press and the involvement of an educated populace
affect international trade. Furthermore, the rise of “next gen-
enables private organizations and citizens to make a
eration” disputes, particularly those pertaining to climate
difference in how things turn out. Second, even imper-
change, has upended classic assumptions about “good” and
fect attempts at international legislation may produce
“bad” actors within trade wars, with developing countries
positive results, and should be pursued toward the
adopting the pro-environmental policy, and so-called devel-
eventual goal of a comprehensive and coordinated sys-
oped countries seeking to have such policy declared illegal
tem of international conservation legislation. Third,
under WTO law (Wu and Salzman 2014). Finally, meaning-
programs of lasting effectiveness in conservation are
ful engagement of indigenous communities in the protection
strongly affected by economic incentives, as evidenced
of biodiversity will require enhanced sensitivity to other ways
by the efforts to save dolphins from tuna fishing, sea
of “knowing” ecosystems and their resources. Honoring tra-
turtles from shrimp netting, and genetic biodiversity
ditional values of biodiversity in the formation of conserva-
from unscrupulous pharmaceutical prospecting.
tion law will be as much about respecting people as it has
The future offers two challenges. Conservation
been about conserving the non-human species on which they
biologists must become more astute in their under-
depend.
standing of law and policy to make their research
effective in achieving conservation goals. And, they
Synthesis must become more sophisticated in learning how to
Environmental regulations and demands of conserva- change laws and policies, and formulate new ones,
tion law press scientists to address and answer that will make conservation law more consistent with
questions they may consider “unscientific.” Likewise, scientific findings. Failure on the first front would make
law and policy require an integrated, interdisciplinary conservation biology an interesting but irrelevant dis-
approach that conservation biologists may publicly cipline. Failure on the second would lead to irreconcil-
endorse, but are personally unprepared to fulfill. Envi- able conflicts between scientific and political
ronmental problems on a worldwide scale may now communities, and the eventual disconnection of con-
require a greater level of coordination than has histori- servation science from conservation law. Understand-
cally been characteristic of the independent nature of ing the transformative power of both NEPA and the
science and scientists. ESA is a helpful place to start when considering the
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Conservation as Vocation
13
It is never wise to seek prominence in a field whose routine chores do not interest you.
Eugene P. Wigner 1992
Keywords 13.1 Conservation as Vocation – First Steps

Professional development · Conservation education ·
Conservation mission · Conservation employment · Inter- 13.1.1 Articulating Your Personal Mission
disciplinary conservation · Conservation programs · in Conservation
Conservation social science · Workforce diversity · Con-
servation advocacy · Conservation leadership Almost everyone, everywhere, in most periods of their adult
life will have a “job,” a contractual arrangement of work done
to receive payment in money or benefits. But people who
become truly satisfied with the work of their life, and who
Overview change the lives of others through their work, are people who
In this chapter you will learn about: pursue and develop a vocation, derived from the Latin verb,
vocare, “to call.” In its oldest (Medieval) use, “vocation”
1. The value of a personal mission statement in con- referred to a calling culminating in a consecration for a
servation biology and the principles for writing one. particular task. The modern definition would describe voca-
2. Elements in educational experiences and profes- tion as a “summons” or strong inclination to a particular
sional relationships that lay the foundation for course of action or state. A vocation is a “life’s work,”
opportunity and service in conservation biology. integrating livelihood with purpose and immediate benefit
3. Criteria for selecting educational programs, with long term legacy. It is a concept far bigger than a
mentors, and jobs in conservation biology. “job,” or even a “career.” It is a path of work and service
4. The skills and practices that will lead to effective seen as a fundamental expression of one’s own identity to the
conservation leadership. world and to other people. And the majority of people active
5. Real problems and potential solutions associated in conservation careers see their work as an expression of
with creating a professional conservation commu- “vocation,” not simply “occupational interest.”
nity inclusive of women and people of color. Having a clear understanding of one’s own mission and
6. Approaches to effective advocacy for conservation. purpose is an essential first step toward the kind of self-
mastery that produces meaningful success and professional
effectiveness. Clarifying a personal mission makes your
motivations clear and explicit, enabling internal restatement

532 13 Conservation as Vocation
of motivations or explanation to others at any time and under 13.1.2 Foundational Elements of Conservation
any circumstances. Clarity of mission produces perseverance Education
through discouraging circumstances and events. The act of
writing a personal mission statement, or statement of pur- In March 2018, the Society for Conservation Biology’s
pose, also helps define not only one’s mission, but also one’s (SCB) official website (https://conbio.org/professional-devel
sense of self – an identity that exists independently of perfor- opment/academic-programs) listed 599 academic programs
mance or external evaluation. An individual’s personal mis- in conservation biology being offered at 525 colleges and
sion statement will change, grow, and mature through time in universities (SCB 2018). This is still a relatively small num-
the same ways that they do. But developing a personal ber of institutions in the global community of college educa-
mission statement, even, in fact, especially, at an early stage tion, but students are not excluded from the conservation field
of your career is an immensely valuable exercise. It not only if they fail to graduate from one of these programs. People
helps you know what you are looking for, but enables you to engaged in conservation today have earned degrees in a
avoid what you are not looking for. This kind of self- variety of disciplines. When considering the curriculum that
knowledge provides the confidence to try new, unorthodox would best prepare you to be a conservation biologist, do
ways of accomplishing meaningful goals, the courage to say what business management guru Stephen Covey said, “Begin
“no” to opportunities that, however prestigious, are not a with the end in mind” (Covey 1989). Start with a clear idea of
reflection of your own ideals, the resilience to cope with the goal. Then form the best plan to reach it.
setbacks and loss, and the freedom to fail and learn from Conservation scientists seek to maintain three important
failure. aspects of life on Earth: (1) the natural diversity found in
An effective, personal understanding of purpose should be living systems (biological diversity); (2) the structure, com-
simple, clear, and memorable. Many students confuse a per- position, and function of those systems (ecological integrity);
sonal mission statement with the answer to the question, and (3) and the resilience of these systems and their ability to
“What do I want to do with my life?” A personal mission endure over time (ecological health) (Trombulak et al. 2004).
statement does not tell you what to do. It points you in a To complement this knowledge, a student must also be able
productive direction to discover what you might be capable to identify individual components of these systems because
of doing, and why it might be meaningful to do it. Not these represent the specific elements to be preserved. Thus,
everyone wants a vocation. Vocations can take on a life of courses in various aspects of biological taxonomy rank high
their own, and make more demands than an ordinary “job.” in their educational importance in a curriculum of conserva-
But many students do enter college searching for a vocational tion biology. One needs to know how to tell the plants and
calling, with questions about what vocation to choose. Others animals apart.
think they already know, and move past the question of With this kind of emphasis, the core of a conservation
“What shall I do with my life?” to “How can I be most biology curriculum needs to provide a clear understanding of
effective in the vocation I have chosen?” In conservation the biological and physical sciences and their ecological
biology, many students who know that they want to be integration. The name of the program is less important than
conservation professionals may want to frame their personal the courses taken to earn the degree. Attempts to define ideal
mission statements in more specific ways. To do so, identify undergraduate preparation for a career in conservation biol-
qualities of your work in conservation without which a career ogy result in irresolvable debate; however, six components
in the field would not be satisfying. One student’s motivations are essential: (1) examination of basic biological processes
might lead to a statement like “I want to work for an organi- and entities at cellular/genetic, organismal, population, and
zation that deals directly with the management and conserva- ecosystem levels; (2) training in mathematical analysis, inter-
tion of endangered species.” That statement is broad enough pretation, and presentation of complex quantitative informa-
to allow the individual to consider both government and tion, and practice in designing experiments to generate such
non-governmental organizations at national and international information; (3) studies of physical and chemical processes
levels, but specific enough to limit and discipline inquiries governing biological processes; (4) use of technologies
and preparations for such a career. Without that specificity, applied in conservation; (5) consideration of social, political,
one could easily be overwhelmed by the variety of possible and cultural forces that shape the practice of conservation in
preparations they could undertake, or of organizations in human society; and (6) an understanding of management
which they might work. If you are serious enough to consider practices to enhance biodiversity and mitigate threats against
pursuing a career in conservation biology, be just as serious it at genetic, population, and landscape scales.
in defining your mission in conservation biology. Take the A curriculum that encompasses all of these elements
time to write a personal mission statement. Reflect on it, would be commendably ambitious, but hopelessly unrealis-
revise it, and use it as a decision-making guide as you tic. Choose a particular emphasis, such as mathematical anal-
continue your work in conservation. ysis, modeling, management application, or conservation
13.1 Conservation as Vocation – First Steps 533
policy and regulation, even as an undergraduate, to gain and biodiversity conservation (Chap. 2). Attempts to deliver
specific knowledge of one or more particular dimensions of conservation mission and conservation outcomes by science
conservation science. The current emphasis on interdisciplin- alone will fail. Conservation mission must be informed by
ary education should not obscure the need for intensive science, but will only be implemented if it is translated
training in at least one of these six key elements. through relationships that understand the ethical, social,
It is hard to do all this in a four-year undergraduate political and economic values of the people you are trying
experience. Most conservationists working today would to influence, and whose cooperation you need to succeed.
admit to being “under-educated” in at least some areas of Because success in conservation mission is tied directly to
knowledge and skill they now use daily. Yet they function interpersonal skills, every student must, at some point, recog-
effectively because they acquired the missing components nize the need for these skills and begin to practice them.
through graduate education, on-the-job experience, and by Through a series of intentional steps, you can develop skills
working with teams of individuals with diverse expertise. If, essential to shift from student to practitioner. With similar
upon inspecting your curriculum, you discover some areas intention, you can acquire experiences and credentials more
that are not covered as thoroughly as others, take electives substantive than grades in courses. But first you must recog-
that develop your competence in under-represented areas. nize what stands in the way.
But choosing the right courses is not enough. For an under- The best undergraduate programs facilitate the beginnings
graduate education to be truly effective, you must overcome a of this transition from student to colleague by requiring
hidden hurdle. undergraduates to participate in at least one intensive
research or professional experience, often in the form of an
internship. Formal requirements for such experiences are
13.1.3 Making the Transition from Student helpful (and a sign that you are in a good academic program),
to Colleague but there is also abundant opportunity to volunteer with
conservation efforts in non-academic settings, where, as a
13.1.3.1 The Hidden Hurdle volunteer, you, the student, can spend lengthy and intensive
Explicit hurdles for undergraduate students are passing time with conservation practitioners in a private conservation
courses and gaining a degree. For graduate students, it is organization or a government agency (Pietri et al. 2013).
the preparation of a thesis. For the new employee in the Research experiences are the most common requirement,
agency, it is meeting goals and targets. But beneath the but whether research experience is an explicit requirement
surface of all three cultures is the same goal – to attain the or not, undergraduates should actively pursue it, beginning in
status of a colleague among one’s associates. You cannot their freshman year. Many opportunities for research
begin this effort too soon. In teaching, the lecture format is an experiences exist—research-intensive courses, research with
efficient way to communicate large amounts of information institutional faculty and graduate students, research with
quickly to a large audience of students. Unfortunately, faculty at other institutions, and research with independent
learning from lectures rewards all the wrong behaviors for agencies or research organizations.
success in the student’s eventual working environment, a Where available, students should take research-intensive
community of fellow professionals in which two-way com- courses built into the undergraduate program of study. Many
munication is essential. As a student, you will not be able to programs include and require a course with a title such as
avoid lectures. But although the knowledge base of conser- Senior Thesis or Independent Research Study. Students who
vation biology is rooted in the natural sciences (and you can enroll in such courses should invest diligently to make this
learn a lot about this from lectures), it is effective human effort of the highest quality, with a final product that is as near
interaction with both other conservation scientists and other to publishable journal standards as possible. If there is no
people in human communities and societies that is the pri- such course in the curriculum, there may be advanced courses
mary means of delivering conservation mission. And deliv- that require a literature review paper in a particular subject.
ering conservation mission is not only about being able to Although neither a literature review paper nor a senior
have effective human interactions. It is also about under- research thesis may ever be published, if its quality
standing how human values, beliefs, attitudes and behaviors demonstrates that the student can do publishable work, it
really work themselves out in organizations and sociopoliti- can make a favorable and distinctive impression that grades
cal structures. These elements will form the foundation of alone cannot accomplish. When evaluators ask for a state-
what makes human interaction effective. This is why we have ment of research interests, students who have prepared a
devoted entire chapters in this book to subjects like ethics senior thesis or professional literature review can speak
(Chap. 10), economics (Chap. 11), and law and policy with greater power and precision than students who express
(Chap. 12), as well as significant sections of material in our their ideas only as vague preferences. Even if you do not
examination of things like ecosystem management (Chap. 9) encounter opportunities to prepare such a work as a course
requirement, you can take on such a project personally. For which you strive to become an asset to others’ work and a
example, you might consult with a professor to design a solution to their problems. As you begin to understand what
literature review that the professor would find useful for his your professors or fellow workers are doing, you will start to
or her own research, and offer to provide the literature review appreciate the difficulty of the problems they face. When you
for them. This approach creates an accountability structure, see the opportunity, offer yourself and your skills as a part of
provides ongoing input from a professional perspective, and the solution to one of their problems. Assist with the collec-
can build a long-term relationship of trust and respect. This tion of data in the field or with transporting equipment to a
kind of practice is good, but look for opportunities that are study site. Help with the preparations for a class or lab
more than practice, the kinds of studies that will be published. exercise. Look for the chance to work for faculty as a teach-
As an undergraduate or beginning graduate student, you can ing aide or research assistant. Graduate students should look
find opportunities to publish if you look for them. The world for opportunities for collaborative research, presentations,
is full of people, many of them scientists, for whom research and publication with faculty and fellow graduate students.
and resulting publication is an exercise in ego-boosting and And new employees in an agency should seek cooperative
self-centered career advancement, but you do not have to be efforts among others that pursue common goals important to
another one of those. Correctly understood, research is a form the agency’s mission.
of service to others, especially to other scientists and students
of science. Producing published work extends the value of 13.1.3.2 The Role of Vocational Experience
your efforts to conservation students and scientists all over In academic cultures, some students see professional prepa-
the world by ensuring that your work is credible (through ration in terms of fall and spring semester classes and sum-
peer review), accessible (through the journal that provides the mers as “vacations” somewhere else. The high cost of college
platform to present it) and “permanent” (within the limits of often demands summers of hard work at the best-paying job
career lifespans). But publication also has positive you can get, but summers are also the best opportunity to gain
implications for your professional and career development. experience and credentials that will facilitate the transforma-
Because publication provides a permanent and accessible tion from being a student of an academic subject to becoming
example of the quality of work that you do, it can be used a citizen of a professional culture. Some conservation
by others to determine your merits for other opportunities. opportunities are poorly paid, but many pay competitive
Faculty and graduate students at most colleges are salaries while providing experiences and opportunities that
conducting individual research efforts. They often seek can radically transform your vision, ability, and influence in
assistants. Wise undergraduates should respond enthusiasti- conservation. Summers offer opportunities to escape the
cally, even if the positions are unpaid. Many such positions boundaries of the local campus, and those opportunities will
are not advertised, but offered privately to students who have appear as posters on departmental bulletin boards,
performed well in classes and fostered positive relationships announcements in newsletters of professional organizations,
with faculty and graduate assistants. Some positions may be and notices on websites. Some types of grants in the United
initiated by students themselves by offering their services as States, such as those administered by the US National Sci-
volunteers to assist in ongoing research efforts. People who ence Foundation (NSF), may have required procedures and
prove themselves reliable in these roles invariably find them- protocols for notification and be posted at various types of
selves being given greater responsibilities and opportunities internet “clearinghouses” such as the Research Experiences
because they gain the trust of those with whom they work. for Undergraduates (REU) program. These and similar
Attaining the status of a colleague begins with understand- programs are targeted at undergraduates and funded for
ing that you must manage an array of relationships, not just long-term research, permitting applications to be made year
an array of courses. A first step toward learning how to be a after year. Some conservation organizations, such as The
colleague is the deliberate initiation of relationships with Nature Conservancy, have year-round programs specifically
your associates. For students, this means engaging professors designed to employ students, (https://www.nature.org/en-us/
in conversations that are not always about you, or your need connect/careers/internships/) while others, like the Student
to clarify (or complain about) a particular assignment, but Conservation Association, focus on placing students in con-
rather discussions of issues of mutual interest and concern. In servation work via both volunteer opportunities (https://
the environment of a conservation organization or agency, it www.thesca.org/serve/young-adult-programs) and more per-
means initiating the same kinds of conversations with other manent positions (www.thesca.org/jobresources). Research
workers, including superiors. Initiate relationships with and vocational training also may come through direct
others by venturing outside your own peer group, educational employment with a government conservation agency or
status, or employment level and addressing issues with them non-governmental conservation organization. In the United
that are about their needs, not yours. Genuine interest that States, agencies such as the National Park Service, Forest
you show in others lays the foundation for the second step, in Service, Bureau of Land Management, Fish and Wildlife
13.1 Conservation as Vocation – First Steps 535
Service, National Resource Conservation Service, and others relationships with her professors, of ongoing research on the
hire thousands of undergraduates each summer for agency effects of burning and mowing on native prairie communities
work. In many cases, agencies have specific programs for the and their associated communities of birds. With strong
professional development and training of undergraduates, recommendations and a good academic record, Sarah gained
often with the intention of grooming participants for future a paid position as a research assistant on such a study at the
employment with the agency. Most state and local govern- US Fish and Wildlife Service (USFWS) DeSoto National
ment agencies have similar programs. In the United States, Wildlife Refuge in Iowa. The work was demanding. Sarah
many private and public conservation organizations, such as started her bird surveys before sunrise on different experi-
Mount Desert Island National Laboratory, Savannah River mental treatments. When bird counts were finished, Sarah
Ecology Laboratory, Konza Prairie, and many others, also began intensive plant sampling that often kept her at work
offer programs of summer research experiences designed for until sunset.
undergraduates. Students can distinguish themselves based Sarah’s dedication, diligence, and increasing knowledge
on what they do professionally much more effectively than of the prairies at DeSoto did not go unnoticed by the USFWS
by their grade point averages. If summers are used wisely, staff. Increasingly they consulted Sarah about plant and bird
and complemented with experiences gained during the aca- responses to the management treatments, and began to make
demic year, students can build an impressive array of use of her knowledge of specific sites. Sarah was asked to
accomplishments and credentials that mark them as distinc- continue as an assistant for a second summer. During that
tive applicants for jobs and graduate research. Consider a year she co-authored progress reports, wrote grant proposals
case history that illustrates how to apply these principles. with her professor, and learned some of the nuances of
working with and gaining grant support from agency
13.1.3.3 An Example of a Student-to-Colleague administrators. By the end of her second summer, the quality
Transition of her work and management recommendations had so
Sarah Bowdish (Fig. 13.1) grew up on a farm in northeastern impressed the DeSoto staff that they arranged for Sarah to
Iowa, and went to college at a small liberal arts institution in speak to members of the Society for Range Management
another part of the same state. During her freshman year, she (SRM) at their annual meeting in nearby Omaha, Nebraska,
developed an interest in conservation, but her college had no and brought the SRM members to the refuge to do it. Sarah’s
major in conservation biology. Sarah chose a closely related presentation to the SRM provided her with many new profes-
major, biology/environmental science, to pursue her goals. sional contacts, as well as many compliments from the
By her sophomore year, she had learned, through building Society’s leaders and members.
Fig. 13.1 Sarah Bowdish, an

undergraduate student whose
involvement in tallgrass prairie
research and management,
combined with an interest in
environmental and conservation
policy, created a record of
accomplishment and a network of
contacts that prepared her for
graduate research. (Photo courtesy
of S. Bowdish)
Sarah then submitted her work for presentation at The conservation goals and pursued them through personal ini-
Wildlife Society’s Annual Conference, to be held that year tiative. Fourth, by making constructive contributions of value
in Austin, Texas (USA). After review, Sarah’s abstract was to her colleagues, Sarah gained an increasingly wide net-
accepted. Impressed by her accomplishments, her college work of contacts that expanded her influence, added to her
paid all expenses for her attendance. Her presentation was credentials, and increased her effectiveness.
not only an important contribution to the meeting but Sarah did not have the advantage of an explicit curriculum
provided an opportunity to establish further professional in conservation biology, nor the benefits of specialized
contacts. During the academic year, Sarah attended two stu- courses or extensive institutional resources typical of state
dent conferences in Washington, DC on issues of environ- universities. Sarah eventually produced a strong, but not
mental law and policy. She used her interpersonal skills to exceptional, academic record. But Sarah was not “lucky.”
become a leader among student caucuses, and an effective Her achievements were consequences of consistent, profes-
advocate for conservation with her own elected sional behavior, rightly applied to well-chosen ends through
representatives. intelligent means.
By her junior year, Sarah’s contributions to conservation
management and policy began to receive wider recognition.
She won one of 75 annual Morris K. Udall Scholarships in
What opportunities do you have in your current educa-
Environmental Policy; the premier US award given to under-
tional environment that can create the same kinds of
graduate students showing ability to make exceptional
outcomes that Sarah achieved? What actions should
contributions to environmental policy issues. Upon complet-
you begin to take to transform these opportunities
ing her undergraduate education, Sarah accepted a graduate
into tangible accomplishments and credentials?
assistantship at the University of Oklahoma to study the
effects of global climate change on plant communities in
tallgrass prairies. Along with fieldwork in Oklahoma, she
traveled to the Czech Republic for more specialized studies
in computer modeling related to carbon reactions in plants
and global climate change. Her interests in politics and her 13.2 Reaching a Wider Audience
strong interpersonal skills were soon having a pronounced
effect on her colleagues at Oklahoma. Within a year she had 13.2.1 A Professional Network of Contacts
been elected a Senator to the Graduate Student Senate of the and References
University and President of the University of Oklahoma
Botanical Society. Over the next 2 years, Sarah earned a In the total national employment of the United States, it is
Master’s of Science in Botany at the University of Oklahoma. estimated that approximately 80% of all new positions are
Sarah describes the value of her experiences this way: filled without being advertised. It has also been estimated that
My experiences at DeSoto were extremely valuable for preparing
the US national average percent success of an individual
me for graduate school. It gave me instant credibility with my applicant in getting an advertised position is less than 7%
professors and peers because I had real experience conducting (about one in 14). But when applicants have direct or second-
research, analyzing data, preparing documents for publication ary contact with the evaluator (i.e., one of the applicant’s
and presenting my work at professional scientific meetings. It
also gave me the confidence I wouldn’t have normally had
references knows the evaluator), likelihood of success has
because I had been through the entire process before. As an been estimated at 86% (Hart 1996), more than a 12-fold
undergraduate, I knew I loved the subject area I was studying increase! These statistics highlight the importance of net-
but I didn’t have a good grasp of how I could use my knowledge working. Why do networks make such a difference in suc-
in a practical way. My research at DeSoto helped me see what
kind of impact I could have to the work of conservation biology
cess, and how does one use them effectively?
and it motivated me to continue my professional development. Every person seeks satisfying personal and professional
relationships. People tend to associate with others who are
Sarah made contributions to conservation entirely on her like them and share their values and goals. Through regular
own merits by following four practices that any student of and repeated actions with one another, cooperation grows,
conservation can emulate. First, as an undergraduate, Sarah and increases even more among groups of people sharing a
used her summers to become involved in conservation prac- common functional objective. Through regular interactions,
tice. Second, Sarah used interpersonal skills and technical and the consequent trust they establish, comes a foundation
knowledge to build trust, establish cooperative relationships, for permanent organizational success which benefits all
and persuade both scientists and the public to change individuals in the group.
practices and behaviors in ways that were of benefit to Educational programs in most colleges that claim to pre-
conservation goals. Third, Sarah set out clear personal pare their students for professional success often fail to
13.2 Reaching a Wider Audience 537
prepare them for these relational realities. The typical college through mutual interest in the subject matter of a course,
curriculum in conservation science or related subjects continue through common efforts in the discipline outside
rewards (through grades) student demonstrations of knowl- the course, and mature through the student’s desire for guid-
edge and technical ability. This tells students that if they ance from the faculty member and the faculty member’s
increase in knowledge and ability, they will gain greater desire to help the student. The circle of primary contacts
reward. And it leads students to believe that they will always will expand for students who actively pursue research and
be evaluated fairly. Such a belief system can survive only in vocational experiences. The greater the diversity of
an academic classroom. Life in the professional world is experiences and organizations, the greater the number of
different. contacts established.
Interpersonal skills are often of primary importance in Primary contacts can become “network partners” if they
conservation because social and political outcomes are nec- possess certain traits. A good network partner respects you,
essary for conservation to occur. Thus, effective interactions likes and understands you, and is involved and influential in
with other individuals are essential for success because such an area related to your mission. Individuals who lack the third
success will require sharing information, explaining ideas trait may be good and valued friends, but will be of little help
and values, listening to and understanding others, and work- in gaining employment or graduate education opportunities.
ing together to solve problems, resolve disputes, and take Individuals who lack the first two qualities are potentially
effective action (Cannon et al. 1996). Yet despite the per- powerful but usually unhelpful. A person with all three traits
ceived need for high levels of training in all of these areas, will be able to form effective networks that lead to more
few academic institutions and even fewer conservation opportunities in employment and education.
organizations have regularly offered, much less required,
courses in human interaction skills.
Relational skills are necessary in building effective pro- 13.2.2 Conservation as a Social Process:
fessional networks, and networks are essential for profes- Involvement in Professional Societies
sional success. As organizations grow in size and applicants
grow in number, more and more evaluators rely on their Like other humans, scientists are social. They form
personal relationships with other professionals, whom they communities of common purpose, not only to achieve their
do know, to make decisions about the selection of applicants, purposes, but to support and encourage one another to con-
whom they do not know. How does one form such contacts, tinue striving toward those purposes. As Michael Soulé wrote
and how does one judge whether or not a potential contact is about the origins of conservation biology, “Conservation
a good one? biology began when a critical mass of people agreed that
Studies of human interactions reveal that people who they were conservation biologists. There is something very
work together toward common goals tend to form stable, social and very human about this realization” (Soulé 1986:3).
positive, and mutually supportive relationships, even when Once a professional society is established, it soon has its
personality differences are extreme. Thus, a first step in own journals, conferences, bylaws, membership
forming networks is to return to your personal mission, use requirements, and certification standards. Although such
it to identify your goals of primary importance, find other societies are formed to advance a common mission shared
individuals (in person or electronically) who share these by their members, they are also formed to provide personal
goals, and join them in mutually effective efforts. Joining and professional benefits to those members. There are many
and working in an organization in which members share a scientific societies engaged in various aspects of conserva-
common mission is an effective beginning. This requires a tion, but the most identifiable and intentional of them is the
commitment to attend the organization’s meetings, partici- Society for Conservation Biology (SCB). What does belong-
pate in its discussions, hold membership in a committee or ing to this society do for you and your development as a
office, and contribute to work that defines the organization’s conservation biologist?
purposes. Discussion groups abound that are defined by The SCB publishes three journals, Conservation Biology,
common interests and concerns for specific issues or Conservation Letters, and Conservation Science and Prac-
problems. By joining such discussion groups and tice. These journals inform members, as well as the world-
participating constructively in the discussion, relationships wide community of scientists engaged in conservation, not
are formed that mature into effective contacts. only of ongoing research in the discipline, but also, through
Although participation in organizations and discussion their editorial policies that determine what will and will not
groups is helpful in building an effective network, the most be published, functionally define the issues of conservation
committed long-term relationships are likely to develop in biology. Through editorials and commentaries, journals also
more focused efforts, and often in employer-employee or serve to form, and inform, community-based views on con-
mentor-student relationships. Such relationships may begin servation issues and values. This is not to suggest that a
society’s journals can dictate what its members think. Any- social contact is natural and easy. For others, it is uncomfort-
one who has ever attended a professional conference knows able and difficult. People who are introverted by nature may
that there is no environment more likely to produce disagree- find it easier to relate to others in activities focused on a
ment than an auditorium full of experts. But despite common purpose, whereas gregarious individuals may prefer
appearances to the contrary, professional societies and their purely social or recreational settings. Choose social contexts
journals shape a collective view of what a discipline is. In this that you find most appropriate to your own temperament and
way they provide members with professional identity, a func- interests, but do not neglect this dimension of professional
tion that is of great value to each person in further refining his life.
or her own personal mission statement.
The SCB holds international, national and regional
meetings at which members present results of their research, 13.3 Graduate Education in Conservation
meet one another, work together in committees or working Biology
groups, form other associations of common interests, and
recognize significant accomplishments of individual 13.3.1 Independent Evaluation for Graduate
members and the Society. Such meetings also produce offi- School – The Graduate Record Exam
cial organizational statements on important issues that allow
the society to speak with a unified voice to the general public No matter how successful one might be in their undergradu-
or the political process. ate career, that career will come to an end, and one is then
Finally, the SCB maintains and makes available resources presented with the first major fork in the professional road –
on jobs and graduate programs in conservation. The more to enter the workforce or continue education in graduate
members that are gainfully employed in conservation and the school. The best choice is determined by what is best for
more satisfied they are with the value of their work, the more the individual who makes it. Graduate education is neither
likely they are to remain active and productive members of required nor essential for many kinds of conservation
the Society. If such satisfaction erodes and is replaced by vocations, and it is not the only or best path for every
professional discontent, the Society suffers. individual and every vocational mission. But graduate edu-
It is valuable to visit the SCB website (http://www.conbio. cation is a road that many do take, and we will travel it first
org) to see how an organization of scientists with common before returning to issues related to jobs and workplace
mission and purpose present themselves socially to one decisions that are likely to affect all graduates, whatever
another and the world. Compare this presentation with other degree or degrees they hold. For those who aspire to a
organizations that have related missions, such as The Wildlife graduate education, it is best to know what one is getting
Society or the Ecological Society of America, and note both into, and how to get into it. To that end, we examine a test
the similarities and differences (for a longer list of called the Graduate Record Exam, or GRE, and why it is
organizations see: https://environment.yale.edu/careers/ important in graduate school admission decisions.
resources/env-professional-associations/#ecosystem-and- Standards for grades vary among institutions, and high
wildlife-conservation) (Fig. 13.2). If you wish to make your grades are more common, less distinctive, and less valuable
contributions to conservation effective, you must express than they once were. Recommendations, while important, are
them in a social context of other professionals. For some, viewed by evaluators with a measure of distrust. References
exaggerate. Sometimes they lie. Past cases of students suing
professors over poor recommendations have tarnished the
influence of letters of recommendations. Today students typ-
ically sign a waiver giving up the right to see letters of
reference, but many references are still reluctant to put nega-
tive comments in writing. This leads to applications that
contain unfailingly positive, but mostly meaningless,
recommendations. Records of employment and experience
are viewed favorably, but subjectively.
Evaluators have only one common measure for comparing
different candidates – the Graduate Record Examination, or
GRE. The GRE is a standardized, multiple-choice test in
three areas of ability: verbal reasoning, quantitative
Fig. 13.2 The logos of three prominent professional societies engaged
reasoning, and analytical writing. The analytical section
in wildlife conservation: Society for Conservation Biology, The Wild- tests the respondent’s ability to think logically and draw
life Society, and Ecological Society of America conclusions or inferences from given information.
13.3 Graduate Education in Conservation Biology 539
Quantitative reasoning tests ability to make inferences and interests, but their productivity and success. You might
draw conclusions using mathematical skills and knowledge. also look for how much collaboration there is within a lab
The verbal section tests the extent of the applicant’s vocabu- group or program based on shared authorship on peer
lary, skills in understanding the meaning of words and the publications.
ability to interpret written expression of ideas. 3. Do graduate students in the program receive adequate
Perfect scores are rare, but even the best programs do not financial support and other compensation? Annual
demand perfection. What constitutes an acceptable, or, at stipends vary from one institution to another, as well as
least, admissible score for an applicant to a graduate program with the type of graduate appointment. Many universities
or a job varies, but any student aspiring to graduate work in offer some Master’s degrees that are more narrowly
conservation biology must take the GRE. The most strategic defined professional programs that do not provide stipends
time is to do it is in the spring of the junior year, the following or other forms of financial support. Most research-oriented
summer, or, at the very latest, early in the fall of the senior Master’s programs, as well as most doctoral programs,
year. If the scores are high enough on the first try, the student however, not only provide a stipend, but waive or reduce
has this important credential in hand and can proceed with tuition, and many of these also provide health insurance as
preparing other application materials. If the scores are low, a benefit of graduate assistantships. Although salary is not
there is time to retake the test before most application the only factor, it is generally unwise to commit to a
deadlines pass. Although the GRE is promoted as an exami- research-oriented graduate program that does not fund its
nation that tests the breadth of educational experience, and graduate students. A lack of financial support means that
therefore not easily “studied for,” most individuals can raise the program has little commitment to or investment in
their scores on a second attempt. Performance on the GRE student success. If a specific program does not sponsor
will improve with preparation and practice. Various GRE test its own assistantships, it might be wise to consider well in
preparation services exist, ranging from free online prepara- advance of enrollment assistantship availability within
tion assistance to more intensive efforts via agencies that other units on campus.
charge for their aid. 4. Do graduates of the program enjoy a good record of
employment or placement in programs of more advanced
graduate study? Statistics on employment of graduates
13.3.2 Choosing a Program can usually be obtained from departmental or university
records. A poor record of placement suggests that the
Many students pursue graduate education to gain experience, program is not perceived as credible by outside reviewers.
knowledge, and credentials to better equip them to be effec- A strong record indicates the opposite and suggests that an
tive conservation scientists. The first decisions you make incoming graduate student can gain access to a
should happen in the application process itself because pre-existing network of supportive contacts who are suc-
applicants should ask specific questions about each and cessful products of the program. For those with an interest
every program they consider: in work outside of academia, it might also be valuable to
see what portion of graduate students find jobs among
1. Is its curriculum accredited by appropriate professional NGOs, industry or governmental agencies.
organizations? For example, people with a particular
interest in forest conservation can determine if their pro- Other program-specific factors are important, but some-
gram is accredited by the Society of American Foresters, times more subjective and difficult to measure. Many things
which maintains a list of schools with accredited curricula. can be learned only by on-site visits and interviews. These
Many other professional organizations with interests in personal contacts are essential to making a good decision.
conservation may accredit or in some way evaluate When meeting with faculty and students of the program in
curricula. person, visiting applicants should try to determine how well
2. Do the faculty have a successful record of publication and the faculty and graduate students work with one another. Do
grantsmanship? Because a student graduating with an they seem to enjoy one another, affirm one another’s efforts
M.S. or Ph.D. in conservation biology or a related field and accomplishments, and work cooperatively in joint
will be judged not only on the degree itself but on research and institutional efforts? An absence of these
publications associated with it, it is wise to choose a qualities, or, worse, evidence of personal hostilities, identifi-
program in which graduate theses and dissertations con- able “camps” or “followings,” or negative comments about
sistently lead to publications co-authored by the student. fellow faculty or students are symptoms of an unhealthy
In many programs, you can inspect the curriculum vitae of personal and professional environment that should be
faculty members online, determining not only their avoided.
13.3.3 Choosing a Project, Graduate Professor usually products of poor planning and weak underlying sup-
and Mentor port structures.
Many key leaders in conservation today can recall the
Although the research project, graduate professor, and men- critical role of inspiring and supportive mentors in their
tor are technically three different things, they are invariably development, and some would even say that the success of
related. Some graduate programs require students to develop current conservation leaders should be measured by their
their own research interests and questions, often independent success in advancing the careers of future leaders in the
of any existing project. However, it is also common for field (Dietz et al. 2004). But not every graduate professor
students to apply to work with a specific advisor because of makes a good mentor. Although it is fashionable for
the work this person is already doing. An ideal project is one academics and professionals to talk more about mentoring
the combines several traits. First, is the research of genuine today than in the past, good mentors remain rare. The word
interest to you as a student? Do you want to have your name mentor is derived from the Greek. It was not originally a
and reputation associated with this effort professionally and, word, but a name. When the Greek hero Odysseus departed
perhaps, publicly? Research interests will grow and expand for the war against Troy, he asked his most trusted friend,
in any healthy and developing career, but it is likely that a Mentor, to see to the education and training of his son,
student’s name will be associated with his or her first Telemachus. Out of this noble relationship has grown the
publications for a very long time. Students should consider definition of what a mentor ought to be: “a trusted counselor
carefully if a proposed project is one that they respond to with or guide, a tutor or coach.” Mentors are distinguished from
enthusiasm, apathy, or dread. Poor fit between student inter- teachers in that they are not merely interested in imparting
est and research effort leads to low levels of motivation, and knowledge, but concerned with the total welfare and growth
that, in turn, leads to poor research and low levels of of the student or younger colleague. The ideal mentor is one
subsequent professional success. Second, is the project of who takes a sincere interest in a student’s welfare, sees their
significant interest to the broader professional and scientific potential for growth, understands what they need to achieve
community? That is, does it matter? Does the research it, and has sufficient influence to create opportunities neces-
address questions of foundational interest to the discipline, sary for that growth to take place. Although every relation-
especially questions that, if answered, may provide general ship might be different, advisors who consistently make time
illumination on current theoretical or management for their students despite heavy workloads with teaching and
predictions? Third, does the research address significant cur- research are to be prized. Students should strive to develop
rent issues or problems in conservation? Often one reaffirms relationships with true mentors and pursue them actively. No
topic relevance later on by completing a literature review, but other influence will have greater effect on their future
taking account of what is being published on the topic in success.
peer-reviewed and public-facing media can provide a base-
line. Fourth, is the research doable within existing constraints
of time, expertise, and funding? Doing good research is 13.4 Innovative Educational Approaches
always challenging, but the probability of success should be
significantly greater than zero. The practicality of a research 13.4.1 The Need for Non-traditional Education
effort will usually be evident if one makes an analysis of
proposed experimental designs, past successes (and failures), Not even the best mentor, no matter how skilled and caring,
faculty expertise, existing team collaboration, and back- can fully make up for a bad educational curriculum. A grow-
ground technical and logistic support for the project within ing criticism of traditional graduate education in all sciences,
the department and the university. Talk to someone who including conservation biology, is that university curricula
knows about mathematical analysis and statistics to evaluate and professors teach students “how to know, not how to do”
the project’s overall design. What kind of data will it pro- (Knight et al. 2008), a problem that can be especially detri-
duce, and will the data be properly structured to answer the mental in conservation science. Many suggest that the reason
question of interest. Check the literature to see if there is there this problem occurs is because “. . .the structure of academia
a published “track record” indicating successful development is built around single disciplines and departments, whereas
and continuation of the research you are considering. Read conservation problems are inherently interdisciplinary. . .”
the faculty member’s vita and see if this professor has a (Muir and Schwartz 2009: 1358). One study of graduate
proven record of success in this line of research. Find out educational training in the sciences that surveyed 4114 doc-
who the project collaborators are, especially the ones respon- toral students in 11 disciplines across 27 institutions found
sible for special equipment or technology on which the that 7 out of 10 believed they were well prepared to become
research depends. Although some elements of a new project independent researchers (Pérez 2005), but, in another study,
are always uncertain, projects that are overly doubtful are less than 20% of doctoral students in science disciplines
13.4 Innovative Educational Approaches 541
thought they had sufficient training in workplace skills such training and experience in democratic discourse so that they
as teamwork, collaboration, organization, and management can participate effectively, responsibly, and reliably in public
(Gaff 2002). Two classic studies, which compared proceedings; and (3) Take courses that intentionally produce
expectations of employers regarding the kinds of skills finely honed critical thinking skills. Critical thinking skills
needed to be a successful professional conservation scientist allow students to analyze problems and recommend solutions
with the kinds of technical skills acquired during graduate in ways that are rational and comprehensive yet sensitive to
school, found that academically acquired skills accounted for context (Clark 2001). Success in conservation depends on
only 7% of the skill set employers desired (Jacobson and being able to effectively interact with people using real life
McDuff 1998; Stanley and Higley 2000). What do such factual information, including real life factual information
results really mean? about differences between people. As conservation policy
We have seen that the hidden hurdle of an undergraduate expert Tim Clark put it, “Scientific thinking seldom accounts
education is to begin to make the transition from a student to for cultural frames of reference, beliefs, or individual
a colleague. In graduate education, this hurdle becomes obvi- motivations that play a major role in problem solving at the
ous and specific. The question is no longer exclusively “How group or societal level. Thus, students should begin by being
do I become a colleague?” but “What kind of colleague do I explicit about their own frames of references – their values,
become?” There is nothing wrong with becoming a disciplin- why they hold them, how to justify them, and how to act on
ary researcher in conservation biology if that is what you them” (Clark 2001: 37). To be effective conservationists,
really want to be. But disciplinary researchers cannot solve you, as a student reading this book at this moment, need to
every kind of conservation problem. Ironically, many of the make sure you are gaining education and expertise in
same educators producing specialized researchers bemoan problem-focused analysis, the context of conservation
the failure of graduate education to produce more individuals problems, and how to use genuinely interdisciplinary
with better management and human interaction skills, more methods to solve them.
interdisciplinary approaches and cross-disciplinary thinking,
more creativity and problem-solution orientation, and more
sensitivity to the complex needs of the increasingly global 13.4.2 An Intentionally Interdisciplinary
workplace environment. Approach
Traditional graduate training in the sciences is designed to
produce researchers, who are intended to become like and The need for interdisciplinary thinking is becoming more
eventually replace the researchers (i.e., professors) under critical daily, especially for solving problems associated
whom they apprentice (Magner 2000; Kainer et al. 2006). with conservation policy that often requires economic, social
This problem, which has been described as academia’s mass and political considerations. Conservation biologist Karen
production of idiot savants (individuals highly skilled in one Kainer and her colleagues noted, in reference to real needs
particular area but inept in everything else), leads to produc- in conservation jobs, “As the workplace has become more
ing people who do not possess sufficient knowledge of interdisciplinary, global, and collaborative, graduates are
human interactions and policy processes to solve policy- required to be technically proficient, broadly trained, and
based problems so prevalent in conservation (Jacobson and capable of working in teams. More than ever, there is also
McDuff 1998; Clark 2001). an emphasis on working toward a more humanistic and
All practicing conservation biologists are, at some level, sustainable society, one in which the “academy” is obligated
policy advocates, even if only in their general desire to to generate knowledge and apply it to concrete problems. . .”
preserve biodiversity on Earth. But most conservation (Kainer et al. 2006: 4–5).
biologists take, intentionally or inadvertently, more active Given the state of things, it is not surprising that there are
roles. Not only do conservation scientists directly or indi- increasing calls both within and outside the community of
rectly inform conservation policy through their research, but conservation biology to not merely make minor adjustments
many are often asked to evaluate different policy options in in the educational process of training future conservation
conservation in light of scientific information and associated biologists, but completely overhaul it. Such overhaul is still
scientific uncertainty (Scott et al. 2008). If students want to in progress, but radical innovations are being attempted in
influence actual practice and policy in conservation, they some schools. Every student considering graduate education
must be intentional about acquiring three particular skill in conservation needs to be aware of these innovations, and to
sets. Specifically, they must (1) Get academic, professional, consider whether traditional or non-traditional approaches to
and experiential training in the science of human interactions, graduate education in conservation biology are the right fit
particularly the ability to understand such interactions as for their skills and goals. That is, every student must take
transactions in values, because such understanding is the personal responsibility to determine the kind of education,
essence of understanding the formation of policy; (2) Gain and the attendant projected educational outcomes, that are
most consistent with their own personal mission and motiva- ways? Be part of a group like this, and you will learn a
tion. Otherwise, as a student, you may find yourself living out great deal of (new) things about how to become more effec-
a career script written by someone else, and you may not like tive in conservation.
the part you have been assigned to play in it. To overcome Be part of a class, group, or organization that takes risks
this problem and avoid its consequences, you must practice with ideas and learns to manage the risks they take. Do you
and master habits that will increase your ability to think have a “safe place” where you can express, listen to, or
creatively. The following sections will provide some respond to new, non-traditional ideas, especially ideas about
examples of non-traditional approaches to graduate education conservation? If you don’t, find that place and participate. In
and training that are intended to help you do this. doing so, you will be intentionally cultivating your own
creativity, and you may begin to see answers to long-standing
problems in conservation that have never been considered
13.4.3 Intentionally Creative Thinking – New before. Social scientist Brené Brown described this as
Paths Out of Old Ruts “leaning into discomfort” (Brown 2000). With practice and
experience, you can then begin to move intelligent risk-
Creative thinking begins with intentionally surrounding taking behavior into your actual work in research, manage-
yourself with people who think differently than you, come ment, or advocacy in conservation, but in ways, and with
from different educational, national, ethnic, and cultural personal and professional support, that will not imperil a
backgrounds than you, and have been trained, or are now project or its stakeholders (Aslan et al. 2014).
being trained, to solve conservation problems differently than Recognize that fear of risk is an expression of fear of
you (Aslan et al. 2014). For most conservation students, failure, but failure is a normal part of professional experience
pursuing a major related to conservation will locate them in and growth, and an important way to find out how to increase
a department centered around biological concepts. The first effectiveness, not only at personal and professional levels,
step is to be intentionally multidisciplinary, cross- but in conservation. For example, the International Smart
disciplinary, and interdisciplinary. The distinctions are well Gear Competition (http://www.smartgear.org) provides a
summarized by Margles et al. who note that “. . .cross-disci- venue for small-scale trials (replete with many failures) in
plinary approaches view one discipline from the perspective pursuit of large-scale success in achieving the conservation
of another, while multidisciplinary approaches involve sev- goal of reducing bycatch in commercial fishing (Chap. 8).
eral disciplines focused on one problem or issue; each disci- Fishers submit ideas to reduce fisheries bycatch (incidental
pline attempts to contribute its own knowledge or approach capture of non-target species) for a relatively modest prize of
to the issue with little attempt to integrate. $30,000. But winning the prize is a better deal than paying
[In] interdisciplinary approaches an attempt is made to inte- the fines and penalties that come with violating bycatch
grate the approaches of different disciplines...” (Margles et al. regulations. For example, the 2011 winner was a Japanese
2010: 2). Many times programs will claim to be interdisci- fisher whose invention reduced seabird bycatch by nearly
plinary while in practice remain at best cross-disciplinary. 90% (Aslan et al. 2014)! The story of the black-footed ferret
Whether your major requires it or not, whether your faculty (Mustela nigripes, Chap. 6) is likewise a story of conserva-
advisor encourages it or not, take courses, attend seminars, tion success borne out of extreme risk. Recall that managers
and get to know faculty and students in other disciplines. captured the last 18 wild ferrets to begin a captive breeding
Learn to think like someone in that discipline. Because con- program in 1987. Had they failed, the black-footed ferret
servation problems are invariably people problems, direct would now be extinct.
your explorations in new disciplines toward scholars and Take time to reflect in a relaxed environment. Some peo-
students who learn how to work with people. Disciplines ple (including some students) never work hard enough to get
like sociology, political science, international relations or anything done. But many, knowing how important it is to
social and cognitive psychology may be very good places achieve solutions in conservation, work without ceasing, yet
to places to start. are frustrated in not finding a solution. In every advancement
Take an active role in a group or organization that takes of your own creativity, you will go through cycles of
problems to “where people are.” It is ideal if you can find a learning, struggle (experiencing difficulty applying new
group that does this in conservation, but it is not essential that skills and concepts) and reflection (Aslan et al. 2014). In
you restrict it to that discipline. Take a look around and ask every such cycle, it will be important to allow for periods of
yourself, “What group takes its message into the local com- stepping back from the pressures of the immediate problem,
munity? What group knows and understands its audience, especially in apparent defeat, and engaging in a relaxing
and frames its messages in ways its audience can understand? activity that takes the mind away from the problem. It is
What group consistently creates interactions between itself often in these “stepping back” periods that the most creative
and local residents in productive and mutually beneficial and inventive ideas emerge. It is also a necessary part of
coping with setbacks, disappointments, and apparent failures. collaboration with scientists from other countries, particu-
One of the most important qualities that define creative and larly in Central and South America, revealed a growing
lasting leaders in any field is their resilience, the ability to frustration and disconnect between the formal graduate edu-
“bounce back” from disappointment and apparent defeat. As cation students were receiving and the kinds of conservation
conservation scientist Clare Aslan of Northern Arizona Uni- problems they were actually being called upon to solve.
versity (USA) and her colleagues put it, “Regardless of one’s Conservation problems, especially in developing countries,
work environment, one must intentionally reframe challenges require engagement with economic motivators, social and
and value time for relaxed thought. In turn, the delight and cultural customs and practices, effective communication to
surprise inherent in creative discovery provide motivation to non-science audiences, and the ability to lead and work with
stay engaged in the long run” (Aslan et al. 2014: 350). others in interdisciplinary teams. These skills were not being
Although creative thinking is critical to conservation suc- effectively addressed by traditional approaches to graduate
cess, the narrowness and lack of creativity found in many education in conservation biology. “Conventional graduate
graduate programs remains a problem, and programs in con- training,” noted Karen Kainer and her colleagues at UF,
servation biology are not exempt. Efforts are now underway “related to tropical conservation and development has typi-
to increase creativity in graduate education programs because, cally separated the two fields, with students focusing on
as conservation scientist Amy Duchelle of the University of either conservation from the perspective of the biophysical
Florida (USA) and her colleagues have noted, “Graduate sciences or development as an extension of the social
programs. . .have an important role in preparing the next sciences. . .Many graduates, however, find that on entering
generation of scientists to build partnerships and exchange the workforce they are required to work beyond the
knowledge with local stakeholders during the research pro- boundaries of the discipline in which they were trained,
cess. Such preparation during formative graduate years can addressing the complex interconnectivity between biological
establish the attitudes, skills, and knowledge that build confi- conservation and human well-being. Fundamentally, devel-
dence and allow conservation scientists and practitioners to oping strong leadership from and for tropical regions is
embrace a more collaborative approach. This requires work- crucial for addressing this monumental challenge” (Kainer
ing in teams, and facilitating knowledge exchange and social et al. 2006: 4).
learning among diverse societal actors . . .” (Duchelle et al. To do this, Kainer and other faculty used a new approach,
2009: 578). forming the Tropical Conservation and Development (TCD)
When graduate studies and programs become too nar- Program “to bridge theory and practice to advance biodiver-
rowly focused, there is a disconnect between research study sity conservation, sustainable resource use, and human well-
and research implementation to solve real problems, the being in the tropics” (TCD 2018). “Rather than creating a
so-called Research-Implementation Gap (Courter 2012). formal degree program,” said Kainer et al., “TCD
The problem of professional narrowness in graduate science concentrates on developing a complementary set of activities
education is now so widespread that the US National Science (courses, workshops, conferences, fellowships, research
Foundation (NSF) developed a program devoted exclusively grants, and visitors) open to students from throughout the
to support more innovative educational approaches. This university who share an interest in tropical conservation and
NSF program, known by the unwieldy name as the Integrated development. The three central goals (training, research, and
Graduate Education and Research Traineeship (IGERT), was promotion of a learning and action network) are blended
conceived to “catalyze a cultural change in graduate educa- together in practice such that most programmatic decisions
tion, for students, faculty, and institutions, by establishing are based on consideration of how a particular decision might
innovative models for graduate education in a fertile environ- maximize gain in each of the three areas” (Kainer et al. 2006:
ment for collaborative research that transcends traditional 9). The TCD does not grant degrees but “. . .offers an inter-
disciplinary boundaries” (NSF 2005). IGERT funding has disciplinary certificate that functions much like a minor. It
supported numerous innovative approaches to graduate train- also provides a supportive learning environment and
ing in the sciences, including conservation biology. Here we fellowships and research grants for M.S. and Ph.D.
review one example. students. . . who are pursuing careers in tropical conservation
and development. . .” (Kainer et al. 2006: 6). Kainer et al.
explained the approach this way: “The University of Florida’s
13.4.4 Interdisciplinary Study Through Conservation and Development program has adopted a
Program-Level Innovation learning and action platform that blends theory, skills, and
praxis to create an intellectual, social, and professionally safe
In the United States, the University of Florida (UF) has been space where students, faculty, and other participants can crea-
long admired as an educational leader in conservation biol- tively address the complex challenges of tropical conservation
ogy. However, despite that reputation, increasing faculty and development” (Kainer et al. 2006: 3).
Unlike traditional approaches, which tend to focus on stool of theory, skills, and “praxis” (“practice with reflec-
independent research efforts, TCD provides grant support tion”) (Vella 1995) (Fig. 13.4). The required training (1) is
for students who want to engage in “practitioner experiences” problem centered, innovating across disciplines to focus on
as a component of their graduate work and education. real-world problems; (2) strengthens personal leadership,
Students supported by these grants form a partnership with building on student experience and enhancing communica-
a host organization, such as a government conservation tion and critical self-reflection skills; and (3) converges in
agency or national or international NGO. The student must field application, linking graduate training and research to a
learn about the host organization and its efforts and collaborative network of others involved in the policies and
objectives to reach the point of being able to complete tasks practice of tropical conservation and development (Kainer
for the organization that advance its mission. The exchange et al. 2006: 7).
of knowledge between the graduate student researcher and The emphasis in TCD is on developing skills essential for
local stakeholders begins with the process of sharing mutu- working at the interface of conservation and development.
ally valuable information as a foundation for building skills Students must learn outside their immediate disciplines, think
needed to solve the problem of interest and concern, in the in terms of linked sociological systems, work in teams, nego-
process generating new knowledge through the experience tiate among competing interests, communicate in nonaca-
(Fig. 13.3). At this stage, graduate students work with demic formats, and reflect critically on their own
stakeholders as partners to practice specific skills important perspectives and actions. One of the ways TCD accomplishes
for conservation, including data collection and analysis, this is through offering courses such as Conflict Manage-
grant-writing, or manuscript preparation. The final stage, ment, Facilitation Skills, Systematic Conservation Planning
knowledge generation, includes stakeholders as partners in and other classes that focus on relating to different and
the creation of knowledge, with the goal of creating more diverse stakeholder groups. All TCD students must take at
accurate information, appropriate decisions, and sustainable least one such skills course to receive their TCD certificate
policies and behaviors (Duchelle et al. 2009: 579–580). In the (http://uftcd.org/academics/certificate/curriculum/). TCD is
TCD Program, some examples of practitioner experiences intentional in its effort “to catalyze a cultural change in
include designing a forest inventory workshop for the local graduate education, for students, faculty, and institutions,
community, evaluating the environmental education program by establishing innovative new models for graduate educa-
of an NGO, developing a GIS framework to assist an NGO in tion and training in a fertile environment for collaborative
mapping an indigenous reserve, or organizing a workshop on research that transcends traditional disciplinary boundaries
conflict resolution for an NGO or agency staff. [emphasis added by authors]. . . (Kainer et al. 2006: 5).”
These kinds of practitioner experiences are designed to TCD projects often include elements of action research or
express the TCD learning and action platform, a three-legged multi-stakeholder processes, and sometimes both. Action
Fig. 13.3 Knowledge exchange

pyramid, a conceptual framework
of strategies for knowledge
exchange between graduate
student researchers and local
collaborators, as would be
employed in a nontraditional
graduate education emphasizing
skills in collaborative
relationships to achieve
conservation goals. (Reprinted
by permission from
Wiley, from Duchelle, A. E.,
Biedenweg, K., Lucas, C.,
Virapongse, A., Radachowsky, J.,
Wojcik, D. J., ... & Kainer, K. A.
2009. Graduate students and
knowledge exchange with local
stakeholders: possibilities and
preparation. Biotropica, 41(5),
578–585. # 2009 The
Association for Tropical Biology
and Conservation)
Fig. 13.4 A conceptual depiction

the graduate education
framework, based on theory,
skills, and practice, used in the
Tropical Conservation and
Development Program of the
University of Florida (USA).
Wiley, from K. A. Kainer, K. A.,
Schmink, M., Covert, H., Stepp,
J. R., Bruna, E. M., Dain, J. L.,
Espinosa, S., & Humphries,
S. 2006. A graduate education
framework for tropical
conservation and development.
Conservation Biology, 20(1),
3–13. # 2006 Society for
Conservation Biology)
research is an approach in which individuals who will use the worth the risks associated with setting up this type of gradu-
research (usually the stakeholders) are themselves involved ate training model” (Duchelle et al. 2009: 584).
in doing and directing research in which they have a strong One step beyond action research and multi-stakeholder
vested interest. Action research is effective in studies in collaboration is to employ demonstration-oriented research,
which the research questions are not rigidly predetermined the translation of scientific understanding into metrics of
and in which stakeholders may want to help shape research performance and cost of implementation under real-world
applications. A multi-stakeholder approach may be used to conditions (Hall and Fleishman 2010) (Table 13.1). For
address complex conservation problems in which there may example, basic research in stream conservation and ecology
be different and divergent interests among many different might identify changes in stream biota that occur downstream
stakeholder groups, and in situations where broad consensus of a dam after the dam has been removed. Demonstration
is needed before decisions can be made, especially when research design would include precise estimates of costs of
shared information may be controversial and appear to such dam removal and quantifiable measurements of
favor the interests of one stakeholder group over others outcomes and benefits associated with changes in down-
(Duchelle et al. 2009). stream biota. From the very beginning, demonstration
research would include “end users” and managers who
would want to apply the results of the research in their own
13.4.5 Systemic Pathways to Creative Education use and management. If end users are engaged proactively,
they often feel co-ownership over the project and become
Different as they are from traditional design in scientific champions of both the science and its products. A well-
investigation, approaches like action research and multi- designed demonstration research study will ensure that the
stakeholder processes may better prepare graduate students performance measures selected are credible and intelligible
for real world conservation problems because of their empha- so that they can be easily translated into direct management
sis on collaboration between researchers and stakeholders. action. When research is planned this way, the results and
Duchelle et al. (2009) summarized the value of such methods innovations it produces are more likely to be adopted by
well in saying, “Graduate programs in which collaborative actual managers of the system or resource. Graduate students
research is the norm provide important platforms from which who are especially interested in working in a specific and
students can experiment with knowledge exchange in their practical area of conservation may benefit from designing and
research. . . enhancing local scientific capacity for improved developing their investigations as demonstration research,
conservation and generating research that is better grounded, especially if they can include people and agencies they
more robust, and more responsive to local needs may be want to work for. Demonstration-oriented research can also
Table 13.1 Examples of hypothetical performance parameters for practical demonstration projects in areas of interest to conservation science. By
using clear metrics and criteria for “success” established in advance, demonstration-oriented research is likely to be more readily and rapidly applied
by managers and practitioners
Performance objective Metric Data requirements Success criteria Source
Improve translocation Number of recruits to the adult Number of new recruits to >20% increase in the number of new Germano and
success through use of population the adult population with recruits to the adult population Bishop
soft release and without soft release (2009)
Improve control of Sustained reduction in rubber Rubber vine cover before >80% mean reduction in rubber vine Valentine and
rubber vine vine cover and 3 years after treatment cover 3 years after treatment across Schwarzkopf
with fire all plots (2009)
Maintain diversity of Species richness and Number of individuals of No significant change (a 0.05) in Valentine and
native reptiles abundance of native reptiles each native reptile species species richness or total abundance Schwarzkopf
in treated and control plots between treated and control plots for (2009)
for 3 years after treatment 3 years after treatment or an increase
in species richness or abundance
within treated plots
Optimize species Number of parasitoid wasp Number of species of trees Parasitoid wasp species richness and Fraser et al.
richness of parasitoid species and rarity weighting and parasitoid wasps for rarity score significantly greater (2009)
wasps and capture of across vegetation types and all possible reserve sites (a < 0.05) than random for each
rare species in reserve number of sites in reserve vegetation type and number of sites
network network
Maintain species Species richness of birds Number of species of trees No significant (a 0.05) decrease in Fraser et al.
richness of birds across vegetation types and and birds for all possible species richness of birds as (2009)
number of sites in reserve reserve sites compared to random site selection
network
Optimize cost versus Cost of reserve establishment Data on cost, such as After five sites are include in reserve Fraser et al.
maintenance of species and management purchase of land or network, cost per additional site does (2009)
richness in reserve ongoing cost of not exceed a specified value per
selection controlling non-native additional species captured
invasive species
Reprinted by permission from Wiley, from Hall, J. A., & Fleishman, E. 2010. Demonstration as a means to translate conservation science into
practice. Conservation Biology, 24(1), 120–127. # 2009 Society for Conservation Biology
be effective in a multi-stakeholder process because the out- rapidly. Suffice it to say here that, even in traditional graduate
come of the research will be more likely to be understood and programs, students who increase their level of knowledge of
implemented by stakeholders if its outcomes are framed in and relational contact with social institutions (such as gov-
measurements they already use in their own management ernment agencies or boards, grass-roots conservation
decisions and interactions with the resource. organizations, or public education systems) will be better
All of these examples are cases of systemic, formal, or equipped, in their current research and beyond, to gain a
programmatic approaches to solving the problem of the hearing for their findings and a platform for their application.
Research-Implementation Gap, but students themselves, (3) Find ways to reward societal engagement and research
especially at the graduate level, can be actors in creating implementation for yourself, other grad students, and even
their own, program-specific solutions. Conservation biologist your own professors. In student clubs and organizations,
Jason Courter identified four of the most effective initiatives create awards like ‘Outstanding Community Achievement
students can take on their own behalf. (1) Facilitate commu- Award’ or “Outstanding Research Application Award” that
nication between researchers and practitioners in designing could be given to faculty or students, perhaps with the sup-
and developing conservation assessments. We have already port and help of local citizens and stakeholders who care
seen this described formally as demonstration-oriented about conservation goals. And make it a point to speak to
research, but even in more theoretical studies, such commu- and work with local conservation organizations yourself, a
nication, especially if initiated from the start, will increase the practice that you will find increasingly rewarding as you
chances that the research, in its very design, will be perceived grow in influence and respect with specific groups and
as applicable and useful by managers. (2) Expand the social communities of people. (4) Increase the availability and
dimension of conservation assessments and support conser- usefulness of research information to practitioners. Graduate
vation plans with transdisciplinary social learning students typically focus their writing on theses, dissertations,
institutions. We will have much to say about this shortly, as and manuscripts to be submitted for publication. These are
the inclusion of social science in conservation is expanding read by other academics. Increase your influence by also
creating, with faculty help and approval, traditional media scheduling time devoted to thinking creatively about current
such as pamphlets, booklets or brochures, or social media work, rather than doing those things immediately demanded
such as webinars, website blogs or live Q&A sessions on by current work); (3) doing things that get and retain the
Twitter, that provide clear applications or best practices attention of others; (4) combining strengths of other leaders
derived from your research or that of other students. These in their own efforts; (5) extending their influence through
kinds of products also exemplify what NSF refers to as networks of relationships; (6) strategically timing their
“broader impacts,” which they evaluate as an important com- efforts for maximum effect; (7) nurturing productive conflict
ponent of grant applications. Graduate students who take toward problem-solving in their conservation efforts and
these initiatives will not only discover that both their profes- (8) cultivating diversity (i.e., having people around them
sional and personal influence increase with people who can who have different backgrounds, training and perspectives)
actually use their findings to achieve conservation goals, and (Manolis et al. 2009).
sooner than they might expect (Courter 2012), but may also In the University of Maryland, College Park’s (USA)
make them more competitive in gaining additional funding to Graduate Program in Sustainable Development and Conser-
continue and expand their efforts. vation Biology (CONS), graduate students under the guid-
Important as both these systemic and individual ance of faculty member James Dietz took personal
approaches are, it is not only how research is done that responsibility for their own education and coordinated
influences the success and effectiveness of conservation in-depth, structured interviews with 10 conservation leaders
outcomes, but also who is doing the research, and whether in federal and state government, academia, and conservation
or not they understand and represent the diversity of NGOs (Dietz et al. 2004). Their purpose was to attempt to
stakeholders with whom they are engaged. define and understand common elements of experience,
background, and perspective in these individuals that had
contributed to their effectiveness and influence in conserva-
13.4.6 Relational Skills in Conservation: tion. Each leader was asked the same questions (Table 13.2),
Learning How to Lead and their answers were recorded, studied, and analyzed. All
10 leaders “acknowledged the value of interdisciplinary train-
Leadership skill in conservation has been called “the most ing in their career development. Even those who emphasized
important attribute in the toolkit of a conservation biologist” the need for grounding in a traditional discipline also
(Dietz et al. 2004: 274). It is believed by many that leaders identified the need for leaders to avoid ‘tunnel vision’ and
are born, not made. This belief is false. Traits of leaders can think about a variety of perspectives that different disciplines
be identified, learned and taught. An equally false myth is bring to a problem. Respondents also identified field experi-
that leadership is conferred by position. Therefore, most ence in both ecology and policy as helpful in their profes-
people do not attain the skills of leadership early in their sional development. Field work in biology was seen by some
careers because they do not hold positions of leadership as the best way to understand both the ecological reality and
until late in their careers. The truth is, a person can be a the challenges facing conservation practitioners on the
leader at any stage of their career from any organizational ground” (Dietz et al. 2004: 275). Interviewees noted that
position if they follow practices and principles that create key qualities of leadership in conservation to be taught or
meaningful influence. Early-career professionals should learn modeled by leaders to leaders included passion or intensity of
and practice leadership through active involvement in hands- character, love for nature, ability to influence, inspire, and
on projects, including volunteer experience, fellowships, and motivate others, persuasiveness, a knowledge base in science
mentoring that complement coursework and training or policy, diplomacy, persistence, willingness to take risks,
programs. Likewise, people who aspire to leadership in con- resilience, knowing when to compromise, integrity, consis-
servation should develop a broad range of leadership skills, tency, and hope (Dietz et al. 2004).
including listening, communication, team building, collabo- Believing, like Dietz et al., that leadership is a skill that
ration, self-management, and strategic thinking. can be taught, Jeremy Martinich, Susan Solarz, and James
Forest conservation scientist Jim Manolis of the Lyons of Washington, D.C.’s American University designed
Minnesota (USA) Department of Natural Resources and his an innovative educational experience for their students in
colleagues identified eight specific traits leaders consistently conservation biology that stressed three elements: (1) review
display that enable them to influence others and change and examination of the discipline; (2) exposure to and inter-
situations for good. These are (1) recognition of the social action with leaders in the field; and (3) a restoration project
dimension of the conservation problem (i.e., recognizing (Martinich et al. 2006: 1579). The first element was covered
what is keeping people from working together and determin- in a traditional classroom format. The second involved
ing how to get them to work together); (2) practicing a numerous guest lectures by political leaders, executives of
personal activity cycle of action followed by reflection (i.e., conservation NGOs, and heads of major federal
Table 13.2 Questions asked of 10 national and international leaders in conservation on issues of leadership by graduate students of the University
of Maryland, College Park’s (USA) Graduate Program in Sustainable Development and Conservation Biology
Questions asked of ten national and international leaders in conservation
Why did you become involved in conservation?
What steps in your career were most important for developing your leadership skills?
Is a Ph.D. necessary for conservation leaders?
Were there specific role models who provided direction in your conservation thinking or activities?
What skills or characteristics are useful to leaders in conservation?
What leadership skills are necessary to direct a conservation organization or agency?
Are coalition building skills important to conservation leaders?
How did you deal with experiences that tested your skills as a leader?
Is it the task of a leader to pursue his or her own vision or the vision of the organization?
Does the conservation movement need a global leader, a Gandhi of conservation?
Based on questions from Dietz, J. M., Aviram, R., Bickford, S., Douthwaite, K., Goodstine, A., Izursa, J., McCarthy, K., O'Herron, M., & Parker,
K. 2004. Defining leadership in conservation: a view from the top. Conservation Biology, 18(1), 274–278. Table format by M. J. Bigelow
Table 13.3 A description of team responsibilities in a restoration project assigned within a project-based learning course at American University,
Washington, DC, USA
Team name Team responsibilities
Problem Identify current biological, ecological, and socioeconomic problems at the site
identification Prioritize and select problems to be addressed in the management plan
Design Collect work products from other teams and determine how to craft a single plan from the variety of work
stakeholders Identify stakeholders such as residents, industry, government, and advocacy organizations
Secure media coverage and financial support, equipment, labor, and expert advice
Continuity Secure commitments for labor (e.g,. scout troops and church groups), infrastructure (i.e., support from local organizations),
and funding (e.g., businesses and government) to continue project in the future
Implementation Design implementation procedures
Manage work events and budget time
Acquire materials, tools, and any necessary permits
Evaluation Set standard of success and determine if project was successful in meeting goals
Create experimental design and manage all data collection/analysis
Reprinted by permission from Wiley, from Martinich, J. A., Solarz, S. L., & Lyons, J. R. 2006. Preparing students for conservation careers through
project-based learning. Conservation Biology, 1579–1583. # 2006 Society for Conservation Biology
environmental agencies, providing a diversity of perspectives et al. 2004: 277). The skills described in these leaders are
beyond traditional academic and scientific research. The third the same skills that students should acquire for themselves
element, the restoration project, required the students to form today.
five teams (Table 13.3), each with a different role in the
project’s completion. Each team had to succeed in order for
the restoration to be successful. Some teams had a harder
Suppose your own college offers no courses within
time completing their assignments than others. Take for
your major that focus on learning managerial and rela-
example, the continuity team, whose mission was to secure
tional skills important to being an effective conserva-
commitments of labor, infrastructure, and funding to bring
tion professional and leader. Based on what you have
the project to completion: “After repeated trial and error the
learned from the preceding examples, what steps would
continuity team eventually learned not to ask for help, but
you take to add these skills to your education?
rather to make clear how joining the effort would be benefi-
cial to the stakeholder” (Martinich et al. 2006: 1582). This is
a valuable lesson because it is one of the keys to understand- 13.4.7 A Career in Conservation Social Sciences
ing how to appeal to stakeholders in building conservation
coalitions and broad-based support. It has been noted by past conservation biologists that conser-
Dietz et al. noted that “Conservation leaders. . .possess the vation problems often start out as biological problems but
interpersonal skills necessary to garner support within and eventually turn into people problems (Teague 1979). Conser-
outside their organization. They have a realistic view of what vation biologists Susan Jacobson and Malory McDuff put the
can and cannot be accomplished, and they strike compromise matter more plainly. “In reality, people are in the beginning,
as necessary to keep the organization moving forward” (Dietz middle, and end of all management issues. Recognition of
this central role will improve our ability to conserve nature. active agents for conservation, interacting with others in a
Conservation is a human endeavor driven by people’s values social context that, if successful, leads to greater social and
toward the management of land and resources. Preservation community commitments to conservation goals. But what
of biodiversity depends upon public commitment to its prosocial sciences could most inform and enhance conservation
tection” (Jacobson and McDuff 1998: 263). And all this education, as well as conservation itself?
means that an understanding of social science, both in theory As Nathan Bennett of the University of British
and application, matter very much in conservation. Columbia’s (Canada) Institute for Resources, Environment
We have already seen (Chap. 1) how conservation is and Sustainability and his colleagues note, “The conservation
increasingly integrating elements of social science into con- social sciences draw on the classic disciplines, such as
servation research and policy, precisely because a social anthropology, sociology, political science, economics, psy-
understanding of the conservation context in any particular chology, human geography, and on applied disciplines such
effort may be more important to the success of conservation as education, development studies, marketing, communica-
effort than scientific understanding. For example, in conflicts tion studies, and law. . .” (Fig. 13.6) (Bennett et al. 2016: 58).
between wildlife and humans – which are often foundational And their increasing use and application in conservation is no
to many conservation problems – both environmental risk passing fad. Bennett et al. observe, perhaps wryly, that
factors and social risk factors must be understood and “Everyone working in conservation, it seems, recognizes
evaluated in order to achieve a solution (Fig. 13.5) (Dickman that natural science alone cannot solve conservation
2010). An appreciation of social factors in conservation also problems. . .” and, as a result “The conservation social sci-
raises the importance of conservation education in solving ence fields have grown significantly over the last few
conservation problems, because, as conservation educators decades. . .” (Bennett et al. 2016: 57). Bennett and his
Nicole Ardoin and Joe Heimlich argue, “building a caring colleagues note, for example, that the SCB’s Social Science
and educated population that can holistically address envi- Working Group, formed in 2003, had become the Society’s
ronmental threats and connect people with nature is critical to second largest such group by 2011. There are an increasing
the success of future conservation efforts. . .” (Ardoin and number of textbooks dedicated to conservation social sci-
Heimlich 2013: 98). And, in the most successful conservation ence, and the SCB itself elected its first President from the
efforts, the learners who benefit from such education become field of conservation social science, Mike Mascia, Senior
Fig. 13.5 Conceptual framework and relationship of environmental permission from Wiley, from Dickman, A. J. 2010. Complexities of
and social risk factors associated with human-wildlife conflict, which conflict: the importance of considering social factors for effectively
provide a pattern for evaluating intensity of interactions in human- resolving human–wildlife conflict. Animal Conservation, 13(5),
wildlife conflicts and in conservation more generally. (Reprinted by 458–466. # 2010 The Zoological Society of London)
Fig. 13.6 Social science and humanities disciplines that interact with science: Understanding and integrating human dimensions to improve
and inform conservation science and practice. (Reprinted by permission conservation. Biological Conservation 205: 93–108. # 2016 The
from Elsevier, from Bennett, N. J., Roth, R., Klain, S. C., Chan, K., Authors)
Christie, P., Clark, D. A., ... & A. Greenberg. 2017. Conservation social
Director for Social Science at Conservation International, in and understanding of individual human behavior (Fig. 13.7).
2017. The growth of the social sciences in conservation is not The growing emphasis on applications of social science in
only certain to make conservation efforts more effective conservation is also creating an environment that fosters
worldwide, but opens new doors of opportunity for students increased multi- and interdisciplinary collaboration,
with social science interests to pursue active careers in con- providing opportunity for students pursuing interdisciplinary
servation. Today, when it comes to social scientists, conser- studies to be well-prepared for conservation careers and
vation organizations are hiring! Major international and contribute substantively to conservation efforts.
national organizations and government agencies are now The reason for the continuing growth and incorporation of
incorporating social science analysis from the very beginning social sciences in conservation is not simply that conserva-
of the project cycle, including The Nature Conservancy, tion biologists have decided to become more inclusive of
Conservation International, Wildlife Conservation Society, non-biologists, but that the social sciences have means to
Rare, Ecotrust, and, among federal government agencies in investigate problems that biology and other natural sciences
the United States, the Fish and Wildlife Service and the cannot, and so have specific and essential applications. Social
National Park Service (Bennett et al. 2016). Social science sciences can be used to understand and describe social phe-
analysis of conservation efforts now affects, and is evaluated nomena, processes or individual attributes under study by
by, units of government from local to international, asking why or how something is occurring. Like natural
applications of scale from individual to global, and subjects scientists, social scientists are also interested in developing
from those as foundational as conservation theory and phi- theory or testing pre-existing theories. Bennett et al. (2017)
losophy to applied decision-making management strategies give examples, such as “What factors are associated with
13.5 Entering a Vocational Setting: How Do I Get a Job? 551
Fig. 13.7 Examples of units and

topics of social science and
analysis applicable to different
scales of inquiry. (Reprinted by
permission from Elsevier, from
Bennett, N. J., Roth, R., Klain, S.
C., Chan, K., Christie, P., Clark,
D. A., ... & A. Greenberg. 2017.
Conservation social science:
Understanding and integrating
human dimensions to improve
conservation. Biological
Conservation 205: 93–108.
# 2016 The Authors)
illegal activity?”, “What governance arrangements lead to

effective conservation?’ or ‘When does money motivate peo- 13.5 Entering a Vocational Setting: How Do
ple to participate in conservation?’ – thus contributing to I Get a Job?
legal, governance, or behavioral economics theory (Bennett
et al. 2017: 95). And social scientists might also critically 13.5.1 Choose Courses for the Job, Not
examine and analyze an issue affecting conservation efforts the Degree
in order to construct more effective solutions, especially in
circumstances in which conservation effectiveness is being The demand for people to work in conservation is global, and
constrained by social behaviors, phenomena, or issues like for that reason, jobs in conservation are diverse. Recent
racism, environmental justice, or equity. Social scientists surveys of the global conservation job market have found
may be able to anticipate future social trends and processes that only about 10% of the available jobs are traditional
that influence conservation through modeling and forecasting academic positions (i.e., university faculty), consistent with
social and economic conditions. Finally, social science might recent findings that only one in six Ph.D. graduates in
be used to plan and identify actions that could improve biological sciences will find long-term employment in acade-
policies, programs or social outcomes in the long run mia (Lucas et al. 2017). The global job market is primarily
(Bennett et al. 2017: 95–96). Bennett et al. conclude, “The non-academic, and so are many of the skills employers are
conservation social sciences are not an optional complement looking for in the workplace (Table 13.4) (Pérez 2005; Muir
but rather a vital component, along with the natural sciences, and Schwartz 2009; Blickley et al. 2013). General disciplinary
for effective conservation decision-making during planning, knowledge and skills in science are important in all sectors,
implementation and management. Integrating the social and and two-thirds of all postings surveyed in recent research also
natural sciences will ensure that these processes are indeed want a person with a background in a specific scientific field
guided by the best available information” (Bennett et al. (Lucas et al. 2017). However, as we have already discovered,
2017: 104). If you are a student with an interest in conserva- other, non-academic skill sets consistently emerge as being of
tion, but also one with dedication to the social sciences, your primary importance in non-academic conservation positions.
time has come. There could be a great career ahead. These include excellent written communication skills,
Table 13.4 Some non-technical skills essential for effective practice and leadership in conservation biology
Professional skill Source
Problem solving and evaluation Noss (1997), Eriksson (1999), Clark (2001), Robertson and Hull (2001) and
Bonine et al. (2003)
Sociopolitical interaction Scott et al. (1995), Noss (1997), Jacobson and McDuff (1998) and Clark (2001)
Law, regulations, and policy analysis Noss (1997), Clark (2001) and Bonine et al. (2003)
Teamwork, conflict resolution, and negotiation Noss (1997), Robertson and Hull (2001) and Bonine et al. (2003)
Public speaking and communication Jacobson and McDuff (1998), Brewer (2001), Clark (2001) and Robertson and
Hull (2001)
Leadership, organizational management, and human resource Noss (1997), Jacobson and McDuff (1998) and Bonine et al. (2003)
management
Stakeholder and community relations Jacobson and McDuff (1998) and Robertson and Hull (2001)
Marketing and social psychology Jacobson and McDuff (1998), Brewer (2001) and Clark (2001)
Strategic planning and project design Scott et al. (1995) and Bonine et al. (2003)
Economics and fundraising Bonine et al. (2003)
Reprinted by permission from Wiley, from Pérez, H. E. 2005. What students can do to improve graduate education in conservation biology.
Conservation Biology, 19(6), 2033–2035. # 2005 Society for Conservation Biology
experience in project management, (Lucas et al. 2017), organizations that focus on translating and integrating scien-
decision-making skills, and policy implementation skills tific assessment into policy-relevant outcomes.
(Muir and Schwartz 2009), with government and private It is one thing to read a job announcement. It is another to
sector jobs also emphasizing specific and analytical disciplin- get the job. That requires not only that you have the skills, but
ary skills in particular areas of science (for example, wildlife that you can clearly signal to prospective employers that you
ecology, statistics, or aquatic biology) (Blickley et al. 2013). have the skills. Signals refer to information available from an
Government and nonprofit employers frequently mention applicant’s curriculum vitae or resume, or from what an
networking, fundraising, and outreach communication skills applicant says in an interview, that communicate to
as important in many positions. When the job posting data employers that this particular applicant has skills in the
were evaluated to identify the most important needed areas. In contrast to a signal, a skill can be defined
non-disciplinary skills, skill and experience in project man- as competency in a particular subject or technique (Blickley
agement was consistently at the top of the list (Blickley et al. et al. 2013). Effective applicants are those who determine in
2013; Lucas et al. 2017). advance what they will provide in written materials, such as a
For most employers, the most commonly identified train- cover letter or vita, and what they will say in an interview,
ing needs for students, in addition to foundational knowledge that speaks directly to the need of the prospective employer.
of conservation biology, were associated with project design In other words, they intentionally deliver clear signals that
and management, policy analysis, fundraising, and econom- they possess competency in the specific job requirements. By
ics and business management (Newing 2010). Overall, noted studying job announcements and then preparing responses
Muir and Schwartz, “. . .practitioners placed higher work- that speak to the employer in the same language used to
place importance on all skill sets compared with academics describe the position, a candidate will have an advantage
. . . Program management, decision making, conducting out- compared to those who do not. For example, a signal from
reach, and implementing policy. . .” were all documented to an applicant could be showing that they possess skill in
be more important to practitioners than academics project management, communicated by speaking of their
(Table 13.5) (Muir and Schwartz 2009: 1362). Multiple stud- experience in managing volunteers, organizing a departmen-
ies have shown that academics place higher importance on tal event, or completing a complex, specific, non-research
research-oriented skills including writing grant proposals, task for a faculty member.
experimental design and data analysis, and communicating Jessica Blickley and her colleagues at the University of
with other scientists (Fig. 13.8). In contrast, practitioners California (USA) offer nine recommendations to
looked for different skills, including conducting a broad undergraduates preparing to leave college. (1) Focus on
number of field methods, conflict resolution, decision- gaining command of key skills in the area you want to
making, and interpreting science for decision makers. But work, and especially focus on gaining competency in skills
even if such training is not offered in formal courses, students with multiple and transferrable applications (for example,
can form their own peer discussion groups where they can competency in using GIS). (2) Decide on a career track
learn about and practice non-academic skills important in before completing your degree and tailor your course work
conservation (Pietri et al. 2013). Students can also look for to that track. (3) Go beyond minimum requirements in your
opportunities to engage in research projects or with selection of courses. Stop thinking of taking courses to get
Table 13.5 Average scores given to 37 job activities by conservation practitioners and academics in order of importance in the workplace and in
graduate training (mean [SE])a
Workplace importance (n)b Training importance (n)b
practitioner academic practitioner academic
(n ¼ 104) (n ¼ 73) (n ¼ 104) (n ¼ 73)
Core competencies
Writing well 4.66 (0.05) 4.81 (0.06) 4.53 (0.06) 4.68 (0.07)
Critical thinking 4.59 (0.06) 4.84 (0.06) 4.48 (0.06) 4.80 (0.06)
Communicating science to decision makers 4.55 (0.08) 4.00 (0.11) 4.17 (0.07) 3.94 (0.09)
Working well within a group 4.43 (0.07) 4.25 (0.09) 3.87 (0.10) 3.84 (0.12)
Interpreting research and making recommendations or assessments 4.40 (0.09) 3.96 (0.11) 4.37 (0.07) 4.21 (0.10)
that decision makers can act on
Public speaking 4.36 (0.08) 4.55 (0.08) 4.13 (0.08) 4.40 (0.09)
Broad ecological understanding or fluency 4.27 (0.08) 4.36 (0.09) 4.36 (0.07) 4.44 (0.08)
Working in a group as a leader 4.21 (0.08) 4.05 (0.10) 3.55 (0.09) 3.59 (0.11)
Total 4.43 (0.04) 4.35 (0.05) 4.18 (0.04) 4.23 (0.05)
Program management
Planning and executing projects on time and in budget 4.35 (0.07) 4.13 (0.09) 3.63 (0.10) 3.65 (0.11)
Running a successful meeting or workshop 4.05 (0.08) 3.58 (0.10) 3.50 (0.10) 3.25 (0.10)
Managing and archiving data 3.95 (0.09) 4.08 (0.11) 3.70 (0.10) 3.92 (0.12)
Managing grants and contracts 3.88 (0.09) 3.79 (0.12) 3.29 (0.10) 3.40 (0.11)
Human resources issues (e.g., employee supervision, staff 3.83 (0.08) 3.47 (0.11) 3.03 (0.10) 2.94 (0.12)
motivation, workplace regulations)
Total 4.01 (0.06) 3.81 (0.08) 3.43 (0.08) 3.43 (0.09)
Decision making
Decision making with multiple competing priorities 4.15 (0.08) 3.39 (0.11) 3.80 (0.08) 3.60 (0.10)
Decision making with minimal information 4.00 (0.09) 3.18 (0.12) 3.67 (0.09) 3.49 (0.11)
Choosing between conservation scenarios or management 3.93 (0.10) 3.37 (0.12) 3.99 (0.08) 3.86 (0.09)
alternatives
Understanding risk assessment 3.78 (0.10) 3.29 (0.12) 3.70 (0.08) 3.56 (0.10)
Total 3.96 (0.08) 3.31 (0.10) 3.79 (0.07) 3.61 (0.09)
Conducting outreach
Communicating science to the public 4.19 (0.08) 4.01 (0.10) 3.92 (0.08) 3.93 (0.09)
Communicating science to private organizations 3.76 (0.10) 3.68 (0.12) 3.62 (0.09) 3.51 (0.10)
Using public relations 3.64 (0.10) 3.31 (0.12) 3.38 (0.11) 3.25 (0.12)
Communicating science to the media 3.62 (0.10) 3.51 (0.12) 3.45 (0.10) 3.54 (0.11)
Total 3.80 (0.08) 3.63 (0.10) 3.59 (0.08) 3.54 (0.08)
Implementing policy
Understanding stakeholder dynamics 3.91 (0.10) 3.46 (0.12) 3.64 (0.09) 3.44 (0.10)
Conflict resolution and negotiation 3.84 (0.10) 2.94 (0.13) 3.65 (0.09) 3.15 (0.11)
Understanding how science informs policy 3.83 (0.10) 3.59 (0.11) 3.76 (0.08) 3.67 (0.09)
Familiarity with environmental law and regulation 3.82 (0.10) 2.93 (0.11) 3.69 (0.09) 3.37 (0.09)
Understanding how to influence policy constructively 3.73 (0.10) 3.25 (0.12) 3.64 (0.09) 3.54 (0.09)
Understanding how policy is made and implemented 3.70 (0.10) 3.18 (0.12) 3.67 (0.08) 3.56 (0.09)
Total 3.80 (0.07) 3.23 (0.09) 3.67 (0.06) 3.45 (0.07)
Research fundamentals
Communicating science to other scientists 3.86 (0.09) 4.55 (0.10) 3.88 (0.09) 4.51 (0.07)
Experimental design and data analysis 3.71 (0.11) 4.62 (0.11) 4.17 (0.08) 4.75 (0.06)
Writing grant proposals 3.65 (0.12) 4.38 (0.13) 3.74 (0.10) 4.43 (0.09)
Total 3.73 (0.07) 4.52 (0.09) 3.93 (0.07) 4.56 (0.06)
(continued)

Workplace importance (n)b Training importance (n)b
practitioner academic practitioner academic
(n ¼ 104) (n ¼ 73) (n ¼ 104) (n ¼ 73)
Conservation research skills
Understanding how to conduct a broad number of field monitoring 3.66 (0.10) 3.75 (0.12) 3.76 (0.09) 3.88 (0.10)
methods
Spatial analysis (GIS) 3.55 (0.10) 3.52 (0.13) 3.74 (0.08) 3.93 (0.11)
Conservation science (e.g., landscape ecology, conservation 3.53 (0.09) 3.58 (0.12) 3.79 (0.08) 4.04 (0.10)
genetics, reserve design)
Designing and conducting research applicable to conservation 3.47 (0.12) 3.77 (0.13) 3.71 (0.10) 4.03 (0.11)
practitioners
Ecological modeling 3.24 (0.10) 3.32 (0.11) 3.46 (0.08) 3.61 (0.10)
Total 3.49 (0.07) 3.59 (0.09) 3.69 (0.06) 3.90 (0.07)
Using the social sciences
Familiarity with social sciences 3.12 (0.10) 3.21 (0.12) 3.00 (0.09) 3.45 (0.09)
Understanding economic models in conservation 3.03 (0.11) 2.88 (0.13) 3.25 (0.09) 3.42 (0.10)
Total 3.07 (0.09) 3.04 (0.10) 3.13 (0.08) 3.44 (0.09)
Grand total 3.90 (0.02) 3.75 (0.02) 3.75 (0.02) 3.80 (0.02)
Reprinted by permission from Wiley, from Muir, M. J., & Schwartz, M. W. 2009. Academic research training for a nonacademic workplace: a case
study of graduate student alumni who work in conservation. Conservation Biology, 23(6), 1357–1368. # 2009 Society for Conservation Biology
a
Range of scale 0-5
b
Significance Kruskal-Wallis test: p < 0.05; p < 0.01; p < 0.001; p < 0.0001; no asterisks, not significant
5.0
Experimental design Critical thinking
Core competencies
& data analyis Comm. science to
Program management other scientists
Decision making Writing well
4.5 Conducting outreach
Academic workplace importance
Implementing policy
Research fundamentals Comm. science to
Writing grant decision makers
proposals
Conservation research
4.0
Using social sciences
Interpreting research and

Ecological
making recommendations
modeling
3.5 that decision makers can act on
Decision making
with multiple priorities
Decision making
3.0 with minimal information
Conflict resolution
Conducting a & negotiation
broad no. of Familiarity with
field methods env. law/regs
2.5
2.5 3.0 3.5 4.0 4.5 5.0
Practitioner workplace importance
Fig. 13.8 Reported workplace importance for 37 job skills by conser- M. W. 2009. Academic research training for a nonacademic workplace:
vation practitioners compared with the workplace importance of the a case study of graduate student alumni who work in conservation.
same skills for academics (range: 0, not important, to 5, very important). Conservation Biology, 23(6), 1357–1368. # 2009 Society for Conser-
(Reprinted by permission from Wiley, from Muir, M. J., & Schwartz, vation Biology)
the degree; take courses to get the job! (4) Start collecting job not be the last stop in a career journey. You must consider
information early in your education, and use it to prepare what kind of vocational setting is appropriate to your goals.
application materials that send clear signals for the kinds of In any vocation, the work environment is an important deter-
jobs you want to get. (5) Identify and acquire job skills that minant of personal satisfaction and professional productivity.
make you different, and potentially more valuable, as a We sometimes have the luxury of choosing from several
match for specific kinds of job compared to other applicants. employment options. Location, salary, and benefits should
(6) Take personal initiative and responsibility for your edu- receive consideration, but most people find that these are not
cation. If the program you are in at your college is not the most satisfying factors. There are additional, and more
offering all the courses you want, identify the organization important, criteria to consider.
you want to work for and, in the local area, contact that
organization or one like it and volunteer. Learn to do the 1. Is there strong correspondence between the
work they do. (7) Keep a record of your experiences of all organization’s mission and your mission as a conserva-
kinds and, where relevant, integrate these experiences into tion biologist? Effective organizations place boundaries
your application material for specific jobs. One suggestion is on their goals and priorities. Their accomplishments are,
to always have a full CV or resume that you can draw from in large measure, the result of working within these
when you ultimately must submit a carefully collated 1–2 boundaries. By focusing on missions and targets for
page version. (8) Talk to conservation professionals doing which an organization is uniquely suited, it achieves suc-
the kind of work you want to do. Don’t focus all your cess. As a potential employee, an important consideration
interactions on academic relationships and take advantage is to look for alignment between your mission and the
of informational interviews with those who have a job you organization’s mission, between your interests and their
think you may want in the future. (9) Budget your time so that interests, and between your abilities and their needs. The
you aren’t spending all of it studying course material, but are closer the match in mission, interest, and need, the more
also investing time gaining skills in doing work you want to satisfying the work is likely to be, and the more you will
do (Blickley et al. 2013: 33). be valued and esteemed by your colleagues in the organi-
Because professional skills sets vary widely among con- zation. You must also consider the differences in reward
servation practitioners, students should interact with the systems associated with different professional cultures and
organizations they aspire to work for during their student determine, in advance, what you want to be rewarded for
career and deliberately consider their career goals to identify and what you are good at producing that will gain those
and get the training their desired profession needs (Muir and rewards. For example, the reward systems of academic
Schwartz 2009). But, having done all these things, how do and non-academic careers in conservation biology are
you choose the vocational setting where you want to work? very different (Table 13.6). Academic researchers are
rewarded for publication and grantsmanship. Conserva-
tion managers in non-academic settings, such as govern-
13.5.2 Choosing a Vocational Setting – Should ment, are rewarded for providing data to guide
I Take this Job? management actions and policies and doing it quickly,
and for getting results from their management actions
Graduate school is only one path to the goal of effective consistent with agency or organizational policies. These
conservation work. And for all but those who become life- different reward structures create different working
long college professors, academic life is something that will environments.
Table 13.6 Contrasting constraints, rewards, and goals of conservation managers and academic researchers
Job Component Conservation Manager Academic Researcher
Motivation Questions driven by need to answer specific Questions driven by theory and basic science
problems, eye toward application
Goal(s) Provide data to manager to guide management; Publish in high-quality journals; compete for research funding
derive guidelines for action
Service Explicit responsibilities to agency; realistic goals Work within context of publicly supported and idealistic goals
Time frame/ Work quickly to obtain data; long planning range Conform to class schedules and academic calendar; projects chosen to fit
work schedule of agency budget process thesis and dissertation schedules of graduate students
Staffing Cost-effective workers Train students in modern techniques; find students jobs; recruit and
support new students
Financial Accomplish as much as cost-effectively possible Support projects via grants; recover indirect costs for home institution
considerations
Reprinted by permission from Wiley, from Huenneke, L. F. 1995. Involving academic scientists in conservation research: perspectives of a plant
ecologist. Ecological Applications, 5(1), 209–214. # 1995 Ecological Society of America
2. Does the organization reach the same audience I want to First, effective conservation workers make work output
reach? Conservation organizations operate at interna- responsive to meeting the needs of others. Every task and
tional, national, regional and local levels. Is the sphere of output, whether written reports, oral presentations, manage-
influence in which you want to operate the organization’s ment decisions, or data analysis, interfaces with others in the
sphere of influence? If yes, then work is likely to be organization who might use such output in their own tasks.
satisfying. If no, it is likely to be frustrating. A biologist Before undertaking a task, determine: (1) Who will receive
who wants to address national issues of conservation is the output? (2) What questions should this output answer to
likely to be unhappy working for an organization that help them accomplish their goals? (3) In what format should
deals only with local or regional concerns. this output be presented so it can be adapted to other
3. Are the people I would be working with individuals I can contexts? Employees who consistently produce output with
respect and trust, and with whom I share common co-workers’ needs in mind and frame their work so that it is
interests? It is always uncertain, given limited contacts easy for others to use are employees who become influential
with potential co-workers, exactly how relationships will in organizational life and are appreciated by others.
develop in an organizational environment. A measure of Effective workers in conservation also make it a point to
optimism is appropriate. Blind faith is not. If potential determine the indispensable needs of their organization or
co-workers display obvious behaviors or attitudes you workgroup, and then make themselves the person who meets
cannot accept, practice patterns of work and activity these needs through their work output. Although all
showing a lack integrity, or display a philosophy of organizations manifest a diversity of work output, there are
work or management that you are opposed to, decline certain core objectives in work that must be accomplished or
the position. Never, under any circumstances, accept a the organization will cease to function. Given a measure of
position with individuals whom you know beforehand freedom in tasks, an employee who gives priority to complet-
you do not like or respect, regardless of their professional ing the organization’s indispensable tasks soon becomes the
prestige. Consideration of your personal “boss” will be organization’s indispensable person. You become that person
even more important. It is a common saying among by determining what the indispensable needs are, and then
professionals in human resources that people do not consistently fulfilling them.
leave jobs, they leave supervisors. Find out who your Finally, current research on organizational productivity
supervisor will be. Make a careful assessment of whether reveals that the most effective and productive organizations
or not you can work with that person. are those that manage through relationships, or, as it is some-
4. Is my job something that I can do effectively and with times expressed, those that manage networks rather than
satisfaction? Even in the best organizations, jobs vary. Is tasks. Effective leadership does not attempt to control others
the proposed job description one that offers an opportunity but to inspire them to pursue common goals and take owner-
for you to display your strengths, grow in your ship of the organizational tasks needed to accomplish them.
competencies, and contribute to the organization’s goals Such an approach requires a high quality of relationships in
in ways meaningful to the organization and to you? A the work environment. Quality of relationships can be
satisfying position should provide a clearly defined orga- maintained by pursuing certain principles in relationships
nizational role and involve you in decision-making appro- with fellow workers.
priate to that role and relevant to your expertise. If you do
not find satisfaction in your work, it becomes much harder 1. Seek to build relationships in a work environment based
to perform well and can hinder personal and professional on common goals, shared tasks, and shared credit for
growth. accomplishment. To the extent possible, expand your
own work tasks and objectives to include the work of
others. When this strategy is followed, it produces not
13.5.3 How Can I Excel in my Work only better work, but a work environment where people
and Nurture Professional share the fruits of accomplished tasks and enjoy their
Relationships? relationships with one another.
2. Take a genuine interest in the welfare of others and build
To experience satisfying work in any position in conservation trust by understanding their needs and being an active
science, your job will require creativity and imagination. part of meeting them. Organizational relationships without
Effective workers who build and develop strong careers trust, no matter how efficiently designed, simply do not
over time follow certain principles in approaching their work. Trust is established by deliberately being attuned to
assigned tasks and in making strategic decisions about both the professional and personal needs of fellow-
which organizational tasks to take on when they have the workers and, whenever possible, being a resource that
freedom to choose tasks themselves. helps such needs to be met.
3. Involve others in decision-making. People do not support become more representative of the communities and cultures
what they do not create. Every person who has a stake in they engage and collaborate with. When a group is composed
creating an organizational objective has a stake in achiev- of people from diverse backgrounds and perspectives who
ing it. Work at providing others the opportunity to share in also possess a diversity of skills, the decisions and
decision-making processes, especially in contexts where recommendations they make are more likely to approach
they can make constructive contributions arising from scientifically optimal and culturally acceptable solutions
their own strengths in experience, expertise, and (Page 2007).
education. Equity, inclusiveness, and diversity have “top down” and
“bottom up” dimensions of relevance. Greater racial and
ethnic diversity among conservationists increases the credi-
13.5.4 How Do I Overcome Barriers? Inclusion bility of conservation efforts with under-represented groups.
and Diversity in Conservation Inclusive and equitable treatment of all ethnic and racial
groups by the conservation profession, especially in the
13.5.4.1 Historic Barriers and Blindspots areas of education, employment, and mentoring, will be
Conservation work, and its workers, are active throughout the needed to sustain the profession itself. The contributions of
world and in every nation, yet the “workforce” of conserva- minorities and underrepresented groups to the next genera-
tion is not currently representative of that world or the tion of conservation scientists are essential to the future work
nationalities of its people. In fact, even conservation of conservation. Solutions to the problem will vary in differ-
workforces within individual countries are often unrepresen- ent cultural contexts, but we will focus here on the nature of
tative of their own populations. In the United States, for the problem and its particular solutions in the United States as
example, the general population, in 2008, was one in which an example of a nation with growing ethnic, national, and
African Americans comprised 12% but only 3.9% of the racial diversity in its population, yet little diversity in its
science and engineering workforce and 2.8% of the national conservation workforce. We will address the nature
biological and life science workforce. In that year, Hispanic of the problem in terms of the particular obstacles that minor-
Americans comprised 16% of the general population but only ity individuals might encounter in entering, continuing, and
4.9% of science and engineering professions and 5.6% of the succeeding in the profession, what is being done to remove
biological and life science occupations (summarized in those obstacles, and what individuals can do to overcome
Haynes et al. 2015). them through their own initiative.
In order for future conservation biologists to serve and Barriers to participation and success in conservation on
relate to international audiences about global conservation the basis of race, ethnicity, nationality, or gender have differ-
problems, they must themselves understand, include, and ent but systemically related causes that exert themselves in all
represent the international human community. There is stages of life, beginning with early childhood experiences. In
irony in the fact that a professional discipline and vocation – the leadership interviews described earlier for graduate
conservation biology – whose mission is the conservation students at the University of Maryland, College Park (USA)
and protection of biodiversity has not done well in fostering (Sect. 13.4.6), the first question asked of participating leaders
diversity among conservation biologists. For many reasons, was “Why did you become involved in conservation?”
the origins of conservation biology were primarily western (Table 13.2). Most leaders “identified childhood enjoyment
and North American (Chap. 1). Its founding leaders were of exploring nature as a primary factor influencing them to
predominantly male and Caucasian, even though the pursue a career in conservation biology” (Dietz et al. 2004:
problems it was attempting to address were global, and 275). These early, positive experiences with nature, so criti-
often centered in Majority World countries, especially in cal for career formation, are the first and perhaps most impor-
South America, Africa, and Asia. The predominance of tant barrier to US minorities in entering conservation and
North American perspectives in conservation biology has natural resource careers (Tanner 1980; Haynes et al. 2015).
often limited its effectiveness, as well as the appropriateness Such early exposure to nature is disproportionately lacking or
of its efforts and approaches, in non-North American settings, limited for US minorities, primarily because of lack of access
and the scarcity of women and people of color in the field has and opportunity to visit and explore natural areas (Haynes
limited its persuasiveness to a wider range of stakeholders, et al. 2015). In the absence of such experiences, youth form
activists, and agency officials from other backgrounds. To negative perceptions of and predispositions toward nature,
increase the chance of success in conservation initiatives and such negative perceptions create barriers to interest in
worldwide, the professional community of people working and exploration of nature at all stages of life and to interest in
on conservation (that is, conservationists) must themselves conservation professions (Haynes et al. 2015).
Minority students also demonstrate lower degree comple-

tion rates in science, technology, engineering, and mathemat- Information Box 13.1. Emerging Leader in Zoo
ics (STEM) degree programs which reduces recruitment to Conservation: Corina Newsome, Biology Graduate
conservation careers. Minority students also report increased Student, George Southern University
perception of current or expected future discrimination, as In high school, Corina Newsome knew she wanted to
well as a lack of mentorship, as barriers that reduce motiva- work with wildlife but did not know what specific
tion to complete these degrees. Such negative expectations vocational pathways were available to her beyond vet-
diminish hope for desirable outcomes in a conservation erinary medicine. During her senior year of high
career, further reducing recruitment to pursuing degrees in school, however, she earned an internship at the
conservation science (Haynes and Jacobson 2015; Haynes Philadelphia Zoo and was introduced to zookeepers,
et al. 2015). field biologists, conservation educators, and others who
These and other problems reveal that changes are needed helped clarify the range of opportunities for conserva-
in the conservation profession. For the conservation commu- tion science. In Corina’s own words: “The most impor-
nity, the most effective path to greater representation and tant experience during that internship was when a
inclusion of minorities is steadfast, long-term investment in zookeeper, who was a Black woman, reached out to
early conservation education, particularly in urban areas or me – one of two Black students in an internship group
otherwise highly modified human landscapes where there of about 80 individuals. She was a lead keeper in the
might be a lack of access to green space (anthromes, carnivore department and invited me to come behind
Chap. 3). Continued engagement with older students will the scenes with her to see, firsthand, what her job
require involving a diverse group of professionals who entailed. It was not until that moment that I realized
may be more effective in establishing mentoring relationships that such a career was a possibility. It was not only
with minority students who have never seen someone like important to receive exposure to the vast number of
themselves in the field. Perceptions of positive outcomes in a careers options, a broadening of my horizon, but to see
conservation career can be increased by highlighting the a reflection of myself in such career.” This internship
accomplishments of such professionals and involving them experience, and others she pursued at the Philadelphia
in educational experiences, internship development, and Zoo throughout college, also gave Corina an advantage
recruitment of minority students (Haynes et al. 2015). when securing work after she earned her Bachelor’s
Perceptions of conservation careers can also be improved degree in Zoo and Wildlife Biology.
by taking students to conservation organizations and With new skill sets in public speaking and animal
workplaces that display diversity in their workforce, equity training, Corina was able to secure the job of Ambassa-
and inclusion in organizational function and structure, and dor Animal Keeper at the Nashville Zoo. There she
providing examples of minorities in positions of influence engaged the public through animal shows, allowing
and leadership (Haynes et al. 2015). guests to experience wildlife and their natural behaviors
Personal initiative also can change perception, opportu- up-close. Had she not learned from her internship that
nity and career outcomes. If one has had limited exposure to keeper positions require at least 3–4 years of prior animal
nature in childhood, they can seek out that experience now at care experience, she would have needed to work for free
any age. They can choose to work with conservation for several years after graduation to even be considered
organizations that spend substantial time in natural areas. for hire as a zookeeper. Corina’s job at the Nashville Zoo
Neither books nor videos can provide the same transforma- pushed her to develop new and important skills, such as
tive effect on the human mind and spirit as extended, adjusting communication techniques to most effectively
immersive time in nature. If this has not been part of a reach a wide variety of audiences, honing her animal
student’s early life experience, they can choose to make it training skills with a thorough understanding of animal
part of their life now. Minority students can also seek out behavior, and evidencing an abundance of patience in
conservation professionals from minority backgrounds and high-stress situations.
learn from them what the future may hold in a conservation Since entering the field of wildlife conservation
career, and where to find opportunities to advance. Such through zoos, Corina has become increasingly inter-
relationships may also lead to long-term professional encour- ested in helping zoos better protect species facing
agement, guidance, and mentoring, among the greatest extinction. As a Graduate Student in Biology at
resources to success and satisfaction in any career. One Georgia Southern University, she has further devel-
such example of this kind of investment is evidenced through oped her field research skills and and work in avian
the story of graduate student Corina Newsome (Information conservation conservation (Information Box Fig. 13.1).
Box 13.1).
(continued)
Information Box Fig. 13.1 Corina Newsome, pictured here with a research in avian conservation as a graduate student. (Photo by
Harris’s Hawk (Parabuteo unicinctus), whose internship and work Quentin Thompson, courtesy of Corina Newsome)
experiences in zoo conservation led to opportunities for expanded
Birds have always captivated Corina’s attention, and “zoos and other wildlife conservation organizations
“were the singular reason that I began to pay closer need to fundraise and budget for programs which pay
attention to the natural world around me.” She hopes to interested students of color to complete internships.
use this training to help zoos better care for the species They must be intentional to articulate and craft
they have committed to conserving and to participate initiatives that promote a much needed diversification
directly in conservation science by garnering public of the field, through (1) opportunities for exposure, and
awareness of, and support for, these efforts. (2) opportunities for the necessary work experience. I
Corina’s story highlights both her own commitment have founded, directed and come across organizations
to personal and professional development, and an that promote diversity in conservation science in one or
ability to connect personal passion with vocational mis- both of those of those capacities. Such efforts include
sion. Her story also suggests that conservation biology non-profit organizations that fund a variety of different
would be strengthened if more young men and women kinds of work experience, government funded initiatives
of color were supported and equipped to become (e.g., through housing and urban development funds),
leaders and in turn, become an example for other and institution-led programs within conservation
young people of color who might have never seen a organizations. These opportunities must increase in
reflection of themselves in the discipline. Reflecting on abundance and funding in order to properly address the
opportunities for removing systemic barriers for lack of ethnic diversity in conservation biology.”
minorities entering the field, Corina notes the following, (C. Newsome, personal communication, August 2019)
(continued)
13.5.4.2 Initiatives of Inclusion accessible to women than symposia organized by male

Michael Foster of the Center for Biodiversity of the Ameri- leaders.
can Museum of Natural History and his colleagues, in their
ground-breaking article, “Increasing the Diversity of
US Conservation Science Professionals via the Society for 13.5.5 How Do I Learn to Recognize
Conservation Biology,” recommended that the Society form Opportunity?
a diversity taskforce or committee with specific plans, metrics
and goals for helping the conservation profession in general, Many people in conservation today work through established
and SCB in particular, to become a more diverse organization educational institutions, government agencies, or
more representative of conservationists and conservation non-governmental organizations. But there are increasing
stakeholders worldwide (Foster et al. 2014). This recommen- numbers of people who could accurately be described as
dation has been put into practice in SCB’s creation of the conservation entrepreneurs who create their own careers in
Equity, Inclusion, and Diversity Committee, whose stated conservation by recognizing opportunities that others over-
objective is to “engage a broader community of students look and responding in creative ways that others do not
and professionals in SCB’s activities, recognizing that imagine.
today’s complex biodiversity conservation challenges require Dr. Katie Moon, a professor in the School of Business at
input and analysis from a variety of voices, vantage points, the University of New South Wales Canberra (Australia) and
and expertise from different geographies, backgrounds, her colleagues have intensively studied the role and develop-
disciplines, and dimensions. . .” (SCB 2019a). This commitment of conservation entrepreneurs and how they contribute
tee is implementing diversity training in SCB sections and to achieving conservation goals. Moon and her colleagues
working groups, and the SCB itself has begun to make grants note that “conservation entrepreneurs. . .influence different
and support available for national and international confer- aspects of the social–ecological system by working collabo-
ence attendance by individuals from underrepresented ratively to generate support for conservation action” (Moon
groups. Addressing barriers to gender representation, new et al. 2014: 1489). Specifically, conservation entrepreneurs
research shows that greater inclusion of women in the disci- become skilled at exploiting three kinds of opportunities.
pline may be a function of helping women gain more promi- Potential opportunities are those capabilities within a bio-
nent leadership roles. Studies that identify systemic causes of physical system (whether natural or human-contrived) that
barriers to participation in conservation, and how to remove can be identified by the conservation entrepreneur as being
them, will help the conservation profession become represen- capable of being manipulated in ways that advance conserva-
tative of the world it is attempting to serve. tion actions, such as the managed release of water from a
Stephanie Sardelis and Joshua Drew of Columbia Univer- reservoir to create habitat for fish or waterfowl. Traction
sity have analyzed the role that leadership positions play in opportunities are created by external events, especially
increasing diversity, specifically in the case of women. They events that create change, or even shock, in the sociopolitical
found that, in the SCB, when women were in the position of environment, that remove previously existing barriers that
organizing a conference symposium, “there is a strong posi- prevented conservation initiative. For example, environmen-
tive relationship between the number of women involved in tal damage to marine life from the explosion of an offshore
organizing symposia and the number of women speaking in drilling rig and subsequent leakage of oil may galvanize
those symposia. Conference organizers should therefore public opinion and political will to pass legislation to reduce
encourage and actively recruit talented female scientists to or curtail such drilling near environmentally sensitive areas.
submit symposium proposals for theses conferences” Existing opportunities arise in circumstances in which no
(Sardelis and Drew 2016: 17). Sardelis and Drew concluded barriers exist, and the entrepreneur recognizes and takes
that, at SCB conferences, “as the organizer pool increased in advantage of such conditions to solve a conservation prob-
size, the opportunity for female organizers increases signifi- lem, such as when new legislative action funds the acquisi-
cantly” (Sardelis and Drew 2016: 6). “Each additional female tion of large areas of land as conservation reserves.
organizer per symposium,” they noted, “has a nearly one-to- Recognition and development of potential opportunities by
one increase in female speakers” (Sardelis and Drew 2016: a skilled conservation entrepreneur “primes the system” to
9). Overall, the strong positive relationship between the num- subsequently take advantage of traction and existing
ber of women involved in organizing symposia and the opportunities when they develop (Fig. 13.9), sometimes in
number of women speaking in symposia highlights the response to the earlier development of potential conservation
importance of mentoring for women from women. Women opportunities.
conference organizers provided a platform for other women Drawing on an example close to home, Moon et al.
to highlight their work which, for multiple reasons, was more identified how, over a 30-year time span, a progression,
Fig. 13.9 Mission and task

pathways in conservation include
(1) research (investigate and
discover), (2) management
(demonstrate and apply), (3)
teaching (educate and interpret),
(4) civic and political activism
(advocate and inspire), (5)
conservation law, policy, ethics
and social processes (legislate and
influence), and (6) organizational
supervision and guidance
(administrate and lead). Most
meaningful and influential roles in
conservation combine multiple
functions described by these
pathways. In everything,
however, there remains a need to
integrate and apply different skills
and abilities to achieve
conservation objectives. (Figure
concept by F. Van Dyke. Figure
design by K. DeVoss. People icon
under CC BY-SA 3.0)
development, and recognition of potential, traction, and

existing opportunities led to major advances in protecting Information Box 13.2 (continued)
Australia’s Great Barrier Reef, which we have examined in Conservationists effectively captured such opportunity
earlier chapters (Chap. 8, Sect. 8.8.2; Chap. 9, Sects. 9.5.4 in ways that led to the legal establishment of the Great
and 9.5.7) (Moon et al. 2014). The first was the rezoning of Barrier Reef Marine Park, the establishment of effective
Great Barrier Reef Marine Park and the second highlighted jurisdiction and oversight structures (the Great Barrier
the development of more intentional management structures Reef Marine Park Authority, GBRMPA), and the pas-
to directly influence the ecological drivers of the Park’s water sage of new legislation to ensure the Park’s protection.
quality. By effectively capturing and exploiting all three These traction opportunities in turn led to existing
types of these opportunities at the right time and in the right opportunity, culminating in the the development of a
sequence, critical conservation goals for the Great Barrier Park zoning plan that kept disturbance away from
Reef were eventually achieved (Information Box 13.2). highly sensitive areas and additional staff positions
that increased capacity to inform policy makers and
involve stakeholders. New staff were able to provide
Information Box 13.2. Recognizing Opportunity
scientific evidence for the effects of mining and other
in the Conservation of Australia’s Great
human activities, more effectively engage stakeholders
Barrier Reef
in the Park’s protection and management, and provide
In the 1970s in Australia, proposed coastal and off-
advances in technology for Park planning and compli-
shore mining around the Great Barrier Reef (GBR)
ance with new legislation and conservation goals. Such
raised public awareness and concern about its environ-
additions contributed to further advancement in legisla-
mental effects, effectively turning a potential opportu-
tion, scientific consensus, public awareness and engage-
nity for more effective conservation of the Reef into a
ment, further rezoning, and the development of
traction opportunity that removed many former
mechanisms for long-term conservation management
barriers from taking effective protective action.
and assessment (Information Box Fig. 13.2).
(continued)
(continued)
Information Box 13.2 (continued)
Information Box Fig. 13.2 A timeline of conservation opportu- opportunities (black) arose in which all circumstances and resources
nity and action on the Great Barrier Reef from the 1970s to 2014. were in place to take effective conservation action in legislation,
Potential conservation opportunities are shown in white, indicating management, public education and awareness, and other fronts to
areas in which conservationists identified aspects of the Great Barrier create a long-term protective environment for the Great Barrier Reef.
Reef ecosystem that would be responsive to management action. (Reprinted by permission from Wiley, from Moon, K., Adams, V.
Traction opportunities (light gray) represent circumstances in which M., Januchowski-Hartley, S.R., Polyakov, M., Mills, M., Biggs, D.,
conservationists identified windows of opportunity arising from Knight, A.T., Game, E.T., Raymond, C.M., 2014. A Multidisciplin-
“shock” events that changed public perceptions and concerns, ary Conceptualization of Conservation Opportunity. Conservation
including the perception that management was ineffective at Biology 28, 1484–1496. # 2014 Society for Conservation Biology)
protecting the Reef from activities like offshore mining. Existing
13.6 Becoming an Effective Advocate for Conservation 563
If you find that you have the knack of recognizing poten- began as an advocacy movement in science (Chap. 1). In its
tial, traction, and existing opportunities in conservation, you early years, members of the Society for Conservation Biol-
might have the makings of a conservation entrepreneur. For ogy were, for all their professional struggles, relatively
example, the development of conservation easements has led unified in the view that conservation biologists must speak
to an entirely new area of legal practice – lawyers who and stand against the threats to biodiversity, and work to
specialize in developing such easements (Chap. 11). In the prevent the extinction of species, not only in scientific stud-
United States, state and federal laws that require “no net loss” ies, but in their appeals to managers, policy makers, and the
of wetlands have created a burgeoning industry in private public at large. They saw themselves as advocates for biodi-
wetland restoration. And the complexity and rapid advances versity conservation. As conservation biology has grown
in statistical analysis and technology required to effectively more respected and established, complete with its own
design and evaluate conservation studies, from data collec- departments, graduate programs, endowed chairs, journals,
tion to publication, has spawned an array of private consult- and funding sources, a more traditional and more conserva-
ing firms that assist investigators in problems associated with tive approach to advocacy has, perhaps predictably, set in. As
technologies like Geographic Information Systems (GIS), a result, the controversy over the proper role and engagement
computation of detection and occupancy probabilities for of scientists in postures of advocacy for conservation has not
cryptic (and endangered) species, and the creation and appli- disappeared, but continued and expanded. As highlighted in
cation of predictive mathematical models for population and larger conversations about the relationships between science
habitat assessment and management. Of course, many and policy in Chap. 12, the issues of advocacy are complex,
individuals now operating as conservation entrepreneurs and must be considered carefully and reflectively by every-
learned their entrepreneurial skills in more traditional roles one who aspires to vocation as a conservation biologist.
in universities, government agencies, or well-established Some conservation biologists, like Peter Brussard and
conservation NGOs at an early stage in their career, and others, have argued that activism by conservation biologists
may have spent many years developing such abilities. If should occur “outside of our professional society” [i.e. SCB]
you find yourself drawn to the kind of creativity and freedom and that conservation biologists and conservation activists
(with its environment of both high risk and potentially high should, as groups, maintain “a strategic and measurable dis-
reward) that is typical of an entrepreneurial environment, tance” from one another (Brussard et al. 1994). This stance is
then seek a position in an organization that fosters develop- consistent with the view that conservation biology is a form
ment of entrepreneurial skills. You may learn the very things of regulatory science that should be understood as “the appli-
you need to know to start your own conservation cation of classical scientific methodology to the conservation
organization. of biological diversity” (Murphy 1990: 203). Conservation
Whatever your career path, you will inevitably discover biologists with this perspective argue that “Conservation
and experience the most important and chronic professional biology only exists because biological information is needed
tension that has always existed in conservation and still exists to guide policy decision-making.” Therefore “the practice of
today – the relationship between the objectivity of the infor- conservation biology ends where science ends and where
mation you provide and your advocacy for the fundamental advocacy begins” (Murphy 1990: 203). If this is a correct
goal of conservation – the protection of biodiversity. The way perspective, then conservation advocacy, specifically the act
in which you handle and manage this tension will influence of working toward a normative outcome or condition in
both your credibility and your effectiveness as a conservation conservation management or policy, is not conservation biol-
scientist. ogy. Rather, the role of conservation biology is to provide
scientific knowledge to resolve technical questions associated
with the formation of conservation policy. But this is not the
13.6 Becoming an Effective Advocate only view.
for Conservation
13.6.1 Professional Expressions of Advocacy 13.6.2 An Alternative View of Advocacy
The relationship between science and advocacy has tradition- Conservation biologist Graham Caughley asserted that “the
ally been one of strict separation in order to avoid any saving of a species from extinction has always been a para-
contamination of professional objectivity. But conservation mount responsibility within the field of biology” (Caughley
biology is not a traditional science, and anyone who pursues 1994). But to assert that science has a “responsibility” to save
the practice of conservation biology as a vocation will invari- species is a normative, value-laden statement of advocacy
ably be brought into contact with issues of conservation that makes no sense unless one appeals to a standard of what
advocacy. Remember that historically conservation biology “ought” to be the correct application of scientific knowledge.
The birth of conservation biology is rooted in such normative allegedly dictated by model results or other kinds of scientific
assumptions that formed the basis of its identify and mission facts. And if that is the case, would it not be appropriate to be
(Chap. 1), and the rapid growth of conservation biology has an advocate of the “better” management plan than the
been a testament to their appeal. In attempting to define the “worse” one?
discipline in its early years, Michael Soulé wrote, “conserva- This second problem leads to a third. If conservation
tion biology is a crisis discipline grounded in the recognition biologists do not consider normative values associated with
that humans are causing the death of life – the extinction of their research, they may find themselves addressing trivial
species and the disruption of evolution” and that conservation questions instead of significant issues. For example, Graham
biology is a response by “those scientists who feel compelled Caughley (1994) asserted that a primary query answered by
to devote themselves to the rescue effort” (Soulé 1991). population viability analysis – how long a population will
The proposition that the sole job of conservation persist – is a trivial question. Having a specific answer does
biologists is to provide information to managers and policy not make the question any more significant. The significant
makers could be called conservation positivism, or perhaps questions, according to Caughley, are, what is putting the
the “just study it” approach to conservation problems. This population’s persistence in jeopardy, and what can we do
view is problematic on three counts. First, conservation about it? But to answer the latter question implies a normative
biologists must make a priori decisions about what informa- end, that the population ought to persist, and its solution
tion to present and how to present it. This is not a question of requires not merely provision of information but changes in
unscrupulous manipulation but of legitimate scientific per- human behavior, as well as specific social and political
spective, and such perspective is determined, in part, by what outcomes that change conditions causing endangerment.
the biologist understands to be the normative uses and values Working towards these ends sounds suspiciously like
of the information. Second, managers and policy makers advocacy.
often interpret objective information, such as model results, If conservation biology is to be defined as information
as offering normative diagnoses and prescriptions (such as driven, then its research is to be dedicated to the needs
why a population is declining and what to do about it) instead defined by management and policy and to the consequences
of seeing model results as objective descriptions of “what of management decisions. It need seek no normative
would happen if” certain conditions prevail. Clouded by this outcomes. Further, it need not be overly concerned with
misconception, policy makers often present normative interdisciplinary study because specialization will remain
recommendations to the public as if such recommendations the most productive and efficient path to generate the greatest
were objective and inescapable conclusions dictated by the quantity and precision of information. On the other hand, if
model, not by their own normative values. For example, conservation biology is value driven and mission oriented, it
managers and policy makers sometimes use ecological and must not only pursue research defined by management need
population models to legitimize policy decisions rather than and by the consequences of management decisions, but it
inform the public (or, in some cases, themselves) about the must also engage the process of management itself, offering
consequences of possible decisions they could make. In the recommendation as well as information. In the latter case,
words of environmental modeler J. B. Robinson, “By interdisciplinary approaches become essential, because con-
cloaking a policy decision in the ostensibly neutral aura of servation biologists would now really need to speak
scientific forecasting, policy makers can deflect attention conversantly with the public and with other disciplines, to
from the normative nature of that decision. . .” (Robinson consider and take responsibility for societal outcomes, and to
1992). As noted earlier (Chap. 10), management decisions evaluate rigorously an array of such outcomes against differ-
are inherently normative, reflecting a desired outcome of ent, sometimes conflicting, normative standards.
what “ought” to occur in the managed system. Conservation
managers are constantly engaged in a process of trying to not
only understand and value ecological processes as scientists, 13.6.3 Examining Outcomes: Implications
but also to make discerning value judgments precisely of Alternative Views of Advocacy
because they must choose the point at which they intervene
in such processes, which is primarily an ethical decision. One’s perspective on what conservation biology is – or what
They must then communicate the reasons for their interven- it ought to be – will determine their view of advocacy in
tion to diverse public interests. Therefore, conservation conservation. Let us begin with the most conservative view,
biologists should ask themselves if they would be better off what is sometimes called professional advocacy or, more
to consider and evaluate normative (i.e., ethical) values in colorfully, the “trickle down” approach. “Professional advo-
their presentation of data and recommendations or to let cacy involves informing policy makers, managers, and the
managers and policy makers use their results to make policy public about issues that arise in one’s area of expertise”
decisions that are disguised as value-neutral statements (Brussard and Tull 2007: 21). Traditionally, adherents to this
view considered publication in books and journals as the only either. In fact, Brewer’s use of the phrase “we do not have
appropriate outlet for information transfer. This kind of advo- time” indicates an assumption of urgency driven by a moral
cacy is sometimes effective, but always slow. Even conserva- imperative: we must save these species! Let the imperative be
tion biologists Peter Brussard and John Tull admit that, in granted. If so, what is the best kind of advocacy to express
their own work, this kind of advocacy is often too slow to do and advance it?
any good. With reference to one study regarding pika The “trickle-down” method is ineffective, and has been
(Ochotona princeps) extinctions and climate change (Beever criticized on a global scale. For example, Erik Meijaard of
et al. 2003), Brussard and Tull note that “Because of the The Nature Conservancy in the province of East Kalimantan,
importance of these findings, we distributed reprints of this Indonesia, and his colleague, Douglas Sheil of Indonesia’s
article to agency heads at a Nevada Biodiversity Initiative Center for International Forestry Research, took on the prob-
meeting. By speeding up information transfer in this way the lem of the “trickle down” advocacy approach in their provoc-
Nevada Department of Wildlife incorporated the findings ative and now classic paper, “Is Wildlife Research Useful for
from this paper into their Comprehensive Wildlife Conserva- Wildlife Conservation in the Tropics?” (Meijaard and Sheil
tion plan for the state in 2005” (Brussard and Tull 2007: 21). 2007). After examining 284 recent publications on tropical
Although still framed as an expression of professional advo- wildlife studies, including 153 from peer-reviewed journals,
cacy, isn’t the assertion that these findings are “important,” Meijaard and Sheil concluded that few of these studies
and not merely “interesting” suggest a judgment about the “. . .address threats to species and fewer still provide input
value of pikas? And doesn’t this pattern of behavior, namely for or guidance to effective management. . . Research is sel-
handing out reprints of the pika study directly to the decision dom judged on its relevance to pragmatic problem solving.
makers at the decision-making meeting look at lot more like Furthermore, many research programs lack the necessary
direct advocacy for pikas than like the strict spirit of “profes- long-term vision and organizational structure for useful
sional advocacy” that only transfers information through applied research. We consulted conservation leaders about
books and journals? our conclusions and all responses suggest that our concerns
Brussard and Tull commend more direct expressions of are not unique to Borneo but reflect wider problems. We
advocacy for conservation biologists if such advocacy takes conclude that conservation research across most of the tropics
the form of: (1) advocacy for science (presenting a positive is failing to address conservation needs” (Meijaard and Sheil
view of science as a method and way of knowing about 2007). If Meijaard and Sheil are right, conservation requires
things, including conservation issues, to the general public); an approach to advocacy made of sterner stuff than the
(2) advocacy for ecosystem services (speaking as an advocate traditional “professional advocacy” described by Brussard
for the economic and material value of goods and services and Tull, even if one does hand out the right reprints at the
provided by ecosystems and ecosystem processes); and managers’ meeting. What would such advocacy look like?
(3) advocacy for the natural world (speaking as an advocate Over 20 years ago, environmental lawyer Daniel Rohlf
for preserving undisturbed nature and encouraging human memorably described conservation biologists as focused
experiences in it) (Brussard and Tull 2007). One can hardly advocates (Rohlf 1995a), which he defined as a person
find any argument with these forms of advocacy as appropri- reporting data concerning an area in which he or she has
ate for conservation biologists. One can hardly find any expertise as well as deeply held convictions, and who works
inspiration in them either. Endangered species and to ensure that the information presented is correctly interpreted
ecosystems will not be saved by extolling platitudes about and rightly applied. Many conservation biologists have long
the value of “the natural world” when the threats to individual believed that focused advocacy, including the development of
species and habitats are particular, direct, and obvious. If regulations and policies to conserve biodiversity, is an inherent
particular species, habitats, and ecosystems are to be pre- responsibility of being a conservation biologist (Noss 1989;
served, what kind of advocacy will get this done, and can a Thomas and Salwasser 1989; Dudley 1995; Rohlf 1995b).
conservation biologist engage in it? Focused advocacy provides specificity that avoids vacuous
Carol Brewer, former Education Editor of Conservation platitudes like supporting “advocacy for the natural world”
Biology, expressed the problem of traditional professional because it speaks to particular needs of particular species,
advocacy this way. “We do not have time,” wrote Brewer, places, and processes. Brussard and Tull, in our earlier exam-
“to wait for our discoveries to “trickle-down” to the public ple of their efforts to influence leaders of government agencies
through the filters of textbooks and other media. We must by passing out reprints of their study on the effects of climate
take more responsibility for translating the results and signif- change on pikas, were not, in that case, acting as advocates for
icance of our research in a way the public – our families, science, ecosystem services, or the natural world. They were
neighbors, and communities – can understand” (Brewer acting as advocates for pikas and they were trying to influence
2001: 1203). Brussard and Tull apparently did not think specific management actions and policies that would affect
that pikas in Nevada had time for the “trickle-down” method
pikas. This is nothing to be ashamed of. It is an example of they refer to as “integrative leadership” (Manolis et al. 2009:
focused advocacy at its best. 881). The role of advocacy in conservation leadership is
Just as there are legitimate forms of advocacy to pursue, critically important. But such advocacy, as an exercise of
such as focused advocacy, there are also forms of advocacy to leadership, must be intentional, never inadvertent.
avoid, expressions of advocacy that can create conflict of
interest or of the appearance of using scientific information
as propaganda. We now examine more carefully the different 13.6.5 Making Advocacy Intentional – Avoiding
forms that advocacy can take, which forms of advocacy can Inadvertent Advocacy
be legitimately pursued in specific circumstances, and what
forms of advocacy should be avoided. In 2019, the SCB stated that its mission was to advance “the
science and practice of conserving Earth’s biological diver-
sity” (SCB 2019b). Affirmation of this statement makes
13.6.4 Can Conservation Biologists Not Be every SCB member an advocate for biodiversity preserva-
Advocates for Conservation? tion. Problems can arise, and credibility can decline, how-
ever, when advocacy is unintended or “inadvertent” in the
As we have seen in the previous section, every conservation course of conservation debate. For example, George Wilhere
scientist arguably works as an advocate for biodiversity pres- of the Washington (USA) Department of Fish and Wildlife,
ervation. Things can become more controversial, however, has noted that “Scientists may engage in inadvertent advo-
when a conservation scientist advocates a specific conserva- cacy in one of two ways. First, they express opinions they
tion policy. As Scott et al. put it plainly, “. . .science alone believe are scientific judgments but . . .are actually ethical
does not dictate policy. . .Societal values are translated into judgments or personal policy preferences. Second, they
policy goals, a policy is selected, management strategies are express opinions they know are ethical judgments or personal
adopted, and actions are taken. . .Too often, discussions about policy preferences, but inadvertently fail to effectively com-
how scientists might engage in the policy process are framed municate the nature of their opinions to policy makers or the
as simplified dichotomies: scientists can either act as public” (Wilhere 2012: 39). For example, notes Wilhere,
advocates for particular issues or policies, or they can shun scientists of the IUCN are engaging in inadvertent advocacy
the policy realm and focus only on science” (Scott et al. 2008: when they assert that “. . .the extinction risk threshold for
866–867). As Scott et al. (2008) go on to explain, this view is categorizing a species as threatened. . .[is described as] ‘the
a gross oversimplification. As introduced in Chap. 12, there highest level of risk that is biologically acceptable.’ Wilhere
are four generalized roles that scientists can play regarding the points out that “There is no such thing as biologically accept-
development of conservation policy. The simplified view able. Appropriate modifiers for acceptable would have been
identifies two dichotomies (shun policy or advocate specific socially, politically, economically, ethically, or culturally
policies). But, between these extremes, they can also provide [acceptable]. . .” (Wilhere 2012: 43).
information on policy-relevant issues when the information is In his summary judgments, Wilhere is not kind to those
requested, or they can evaluate the full range of policy guilty of inadvertent advocacy. “. . .I believe inadvertent
options in light of scientific information and scientific uncer- advocacy is professional negligence. . .If scientists unknow-
tainty (Pielke 2007; Scott et al. 2008). In these second or third ingly express policy preferences, develop quasi-scientific
roles, if scientists begin to advocate certain policies, instead definitions of recovery, criticize a recovery plan because
of merely provide information to decision-makers, their they do not understand its policy context, or do not acknowl-
information may begin to be mistrusted (Scott et al. 2008). edge the values inherent to a definition of threatened, then
In these roles scientists would not only be providing policy they have stepped outside their role as scientists and engaged
relevant information, but in advocating specific policies, in inadvertent policy advocacy” (Wilhere 2012: 44, emphasis
providing policy prescriptive information (Revkin 2007). his). If the advocacy is subsequently identified and
All of these examples reveal that conservation scientists recognized as advocacy, political decision makers and the
will inevitably interact in some way with conservation policy. public will not know if it was inadvertent. They might there-
Therefore, some conservation scientists have argued that fore suspect that the advocacy was intentional and the scien-
students studying conservation should be specifically and tist tried to hide the advocacy and mislead her hearers. When
intentionally trained for such interactions. Manolis et al. this is the perception, the public perceives attempted manip-
(2009), who have already helped us understand the impor- ulation and the scientist’s credibility declines. As Wilhere
tance of leadership in conservation, stress that conservation explains, “Inadvertent advocates facilitate amoral policy
leaders inevitably and invariably interact with conservation making by expressing their values as policy preferences in
policy, especially as advocates of integrating conservation ways that are nearly indistinguishable from science. Success-
science into policy, management, or society at large, a role ful conservation of biological diversity depends on sound
science, but will ultimately be determined by society’s arrangement is perfectly ethical under the constraints
values. Inadvertent policy advocacy undermines the rational described. Neither the firm nor the energy company takes
political discourse necessary for the evolution of society’s the role of an advocate. But suppose that a group of local
values (Wilhere 2012: 44). citizens becomes concerned about the drilling activity, not
Advocacy, at some level, is inherent and intrinsic to the only because of potential detrimental effects on elk and other
makeup of conservation. Scientific credibility can coexist wildlife and their habitat, but because of perceived negative
with advocacy as long as an individual transparently effects on the aesthetics of the landscape, and the potential for
identifies what he advocates and the reasons, both scientific negative effects on local economies and culture by creating a
and ethical, for this advocacy. “gas boom” in an area that has traditionally been dominated
by an agricultural economy. They form an organization
whose mission is to stop natural gas exploration in the
13.6.6 Avoiding Conflicts of Interest local area.
in Advocacy As a biologist who lives in and enjoys the area, you are
sympathetic to the aims of the local citizens’ organization.
Conservation scientists can maintain credibility with most But should you join the organization and be an advocate for
people and groups if they are transparent in their advocacy its mission while this study is in progress? The answer is
and the basis for it, but a conservation biologist can be “no.” Taking a position against drilling for natural gas while
unethical in what she advocates, as would be the case if at the same time conducting a study to determine the effects
such advocacy was motivated by a desire for personal gain. of drilling on wildlife and habitat conditions represents a
When this happens, advocacy will be compromised by a conflict of interest, an unacceptable way of expressing advo-
conflict of interest, and credibility will be lost. An obvious cacy for a particular policy (ban drilling!). Your interest in
conflict of interest might be accepting bribes or exploiting a scientific objectivity (what really happens on a drilling site
relationship by advocacy of a particular position. But most after drilling is over) is in conflict with your interest in
conflicts of interest are much more subtle, and therefore, stopping the drilling altogether, which would be your stated
more professionally sinister and potentially damaging to aim as a member of the protective organization. In this case,
your career. Let’s inspect one example. the right thing to do is to NOT join the protective organiza-
Suppose you are a conservation biologist who works for a tion, but to continue your research and present the outcomes
private environmental consulting agency in the USA. As of it in a fair and professional manner. It is for this reason that
shortages of oil make natural gas a more attractive fuel option many private conservation organizations, consulting firms,
for some forms of energy use, an energy company is and government agencies prohibit, as part of their employee
motivated to search for prospective natural gas supplies on contract agreements, membership in other conservation
large private ranches in a remote area in a western state. The organizations whose specific missions might create a conflict
ranch lands are heavily used by wildlife, one of the most of interest with their mission. Thus, when you consider
visible and charismatic being elk (Cervus elaphus). Drilling employment with any kind of conservation organization or
to find and remove the natural gas destroys vegetation on the entity, consider the implications of that employment on your
drill site, but the energy company has various options for own role and activities as a conservation advocate, and
revegetation and reclamation. The company asks your firm to whether you can accept those implications and limitations
conduct a study of the various restoration methods and report that may come with the job and the agency you work for.
how elk respond to each of them. They want to know, if the Your own view of advocacy should be one of your most
elk used drilled areas before disturbance, do they return to the important criteria to consider as you explore different kinds
site after drilling is over or not? If so, are their use rates the of conservation vocations and employment. Some kinds of
same or different than they were before drilling? Does the work in conservation biology are almost entirely about advo-
operation cause a permanent loss or degradation of habitat, or cacy. Other types of conservation efforts require that work
a long-term change in a site’s biodiversity, or is it limited to and advocacy be kept separate from one another, and many
short term effects? Which reclamation techniques best pre- kinds of work in conservation biology fall somewhere in
serve natural biodiversity and natural levels of elk use of the between. Specifically, you should ask, “Am I pursuing a
site? Your employer takes on the project, with a contractual career in conservation biology in order that I might provide
understanding that the energy company will pay your firm for information about the biology of endangered species or in
the service of its investigation regardless of the results. With order that I might provide expertise in how to save
that commitment in the contract, your firm agrees to prepare endangered species? Do I wish to focus on the measurement
and present the results of its study in a final report to the of biodiversity, or to provide an informed analysis of how to
company at a specified time. preserve it?” Perhaps the most important question is, “Do I
This kind of scenario is not uncommon. One of us has wish to work with an institution or community that provides
participated in studies like it (Van Dyke et al. 2012a, b). The information about how to conserve species, or do I wish to be
a member of an organization that actively uses such informa-

tion for species preservation?” opportunity to investigate and discover the nature
An individual doing work in one category is not more or and causes of biodiversity loss, and the solutions to it,
less of a conservationist than one working in another cate- through direct investigative research. You can demon-
gory. And, you may find yourself doing each of these things strate and apply the results of such research as a con-
at different points in your career. However, if that person is servation manager or conservation practitioner who has
you, it is your responsibility to be happy, and free from actual influence and responsibility to care for systems
internal conflict, in the professional life you have chosen. and species. You can educate and interpret the work of
Therefore, the issue of advocacy is one you must address in conservation to people who want to know more, but
advance, before the choice of employment is made, so that don’t yet know enough, to make a difference them-
you will not have to make hurried and, perhaps, unethical selves. The education you provide can change that.
responses to issues of advocacy in the heat of crises or You can advocate and inspire other people to take
conflicts that will inevitably occur in the work, including decisive action on particular issues in conservation,
your work, of conservation. like raising money to acquire land for preserved
areas, working for or against significant political and
policy decisions that will shape conservation manage-
Synthesis
ment, or bringing people into contact with places and
This chapter has taken the risk of offering some pre-
species that will move them to take actions of their
scriptive advice because conservation is performed by
own. You can follow a path to legislate and influence
conservationists, not by words in a textbook that magi-
human behavior and actions through work in conserva-
cally assemble themselves into correct conservation
tion law, regulation, and policy, or through efforts in
actions. We encourage all students of conservation
the social sciences that change community perception
biology to consider what the future may hold for them-
of conservation goals, or, at the personal level, contrib-
selves and their discipline. We make the future every
ute to the formation of conservation ethics that will
day by every daily choice. Our choices reflect our
guide individual choices about how to interact with
commitments to what we truly value. There are three
the Earth’s non-human species, which form the sum
key principles to success in pursuing an effective edu-
of its biodiversity. You can administrate and lead
cation and vocation in conservation biology. The first is
conservation organizations, government agencies, aca-
the principle of personal responsibility. You, not your
demic clusters or professional societies or their work-
college, advisor, roommate, parents, income level,
ing groups to enable people to achieve more together
national origin, or upbringing are responsible for the
than they could have achieved alone.
outcomes of your education and your vocation. You
Few meaningful jobs in conservation are devoted
cannot always control the circumstances and stimuli
solely to one of these categories. The differences are
thrown at you in the great adventure of life, but you
more often matters of emphasis. But in any given
will always have the freedom to choose how you will
position, effective work in conservation requires a per-
respond to them. Value and use that freedom. The
son to integrate and apply their knowledge and
second principle is to define what you value. There is
abilities in all these roles, not only in work, but in
no satisfaction in climbing the ladder of success only to
life. It is precisely this integration and application that
find it was leaning against the wrong wall. Be sure of
makes conservation a true vocation, or calling, that
what you value and pursue that. The third principle is:
influences all aspects of how one lives their life, as
take action. If you accept personal responsibility for
well as a noble and respected profession in which
how you are going to respond to the circumstances and
they work, on their own and with others, to benefit
opportunities of your life, and your career in conserva-
other people and preserve the Earth’s biodiversity.
tion biology, and if you know what you value achiev-
In the end, the most basic and fundamental goals of
ing in this career, then take concrete action in a
conservation biology, the preservation of Earth’s bio-
pre-considered plan to pursue that goal.
diversity in its living systems, with their health and
There are many paths to a career in conservation. To
integrity, will not be achieved by improvements in
name these by job titles would be tedious and uninspir-
technology, economics, or information transfer. An
ing. Let us instead return to the basic categories of tasks
Earth in which the world’s biodiversity flourishes will
that enable the work of conservation to succeed
be one in which nature interacts with a different kind of
(Fig. 13.9). You can choose a path that provides
human being, and a different kind of human
(continued)
(continued)
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Glossary
A nity and “evenness” the relative abundance of each species

in the community.
acidification The process through which the pH of surface alpha rarity Rarity or reduction in numbers of individuals of
fresh waters, especially lakes, declines because of inputs each species on a site or in a community that occurs as more
of acidic precipitation in the form of rain, snow, or fog. species are added to the site or community (i.e., rarity in
action research An approach in which individuals who will numbers of individuals of a species in a site-specific popu-
use the research (usually the stakeholders) are themselves lation that occurs as the site increases in species diversity).
involved in doing and directing research in which they alteration of flow regimes Human-caused changes to
have a strong vested interest. stream flow, including the quantity, duration and seasonal
adaptive management The structuring of policy or man- pattern of flows. Examples include dams, reservoirs
agement actions as a set of testable hypotheses to promote (impoundments), hydroelectric facilities, locks, levees,
learning from policy implementation, and the subsequent and water withdrawal.
design and use of management actions as ongoing alternative stable states In lake systems, the potential for
experiments intended to provide reliable information different conditions to prevail at similar nutrient levels,
about the system and inform future management with rapid transitions occurring between states (e.g., from
decisions. abundant submerged macrophytes in clear water, to dense
age structure The proportion of individuals in a population phytoplankton in turbid water).
at each age, or in each age category. aquatic ecological system Stream networks representing a
agent-based dispersal models A particular type of dis- range of areas with distinct geomorphological patterns
persal model that assesses the “connectedness” between connected by similar environmental processes such as
the model’s habitats, or, habitat connectivity of the model, hydrologic, nutrient, and temperature regimes.
determined by the model’s neighborhood or movement aquatic zoogeographic unit (AZU) The highest level of
rules, which specify the distance across which sites are classification in a coarse-filter classification system for
accessible to organisms (the “agents” of the model) by streams, lakes, and rivers, which also serves as the overall
virtue of their dispersal or gap-crossing abilities. planning unit in initial conservation assessment. AZUs
alleles The variety of gene forms found at the same place on conform to major freshwater drainage boundaries, generally
a chromosome. The number of alleles present for a given 10,000–100,000 km2, and are distinguished by differences
gene is one measure of the genetic variation of a in the continental and regional distribution of animals.
population. assisted migration Intentional translocation by humans of
allelic diversity The average number of alleles per genetic organisms to areas more suitable for their persistence.
locus. A loss of unique alleles leads to a loss of allelic average heterozygosity The average proportion of indivi-
diversity and overall genetic diversity. duals in a population that are heterozygous (carrying two
alpha diversity The diversity of species within a commu- different alleles) for a particular trait. This metric reflects the
nity on a specific site, normally described as a measure of proportion of heterozygous individuals measured across
two attributes – species richness and species evenness, several loci. As a metric, average heterozygosity is used as
where “richness” is the number of species in the commu- a measure of genetic diversity in a population.

F. Van Dyke, R. L. Lamb, Conservation Biology, https://doi.org/10.1007/978-3-030-39534-6
572 Glossary
B bleaching See coral bleaching

bycatch The inadvertent capture of animals that have
basic rule of conservation genetics The assertion that nat- become hooked, trapped, or entangled in fishing gear
ural selection for performance and fertility can balance deployed with the intention of catching something else
inbreeding depression if the change in the inbreeding during commercial fishing activities.
coefficient (ΔF) is no more than 1% per generation. The
1% rule is referred to as the basic rule of conservation
genetics because it serves as the basis for calculating the C
irreducible minimum population size consistent with the
short-term preservation of fitness. carbon sequestration The process of capturing and storing
bequest value The value of knowing that something is atmospheric carbon dioxide. One natural method is refor-
preserved for future generations. estation or afforestation, where trees store carbon as a
benefit cost analysis See cost benefit analysis component of their biomass as they grow. Another
beta diversity A measure of the rate of change in species method is to inject CO2 produced via industrial processes
composition of communities across a landscape (some- deep into underground wells or beds of underground
times called beta richness). Beta diversity would be minerals so that it does not enter the atmosphere.
0 when sampled sites all have the same species composi- charismatic megafauna See phenomenologically signifi-
tion, indicating a highly homogeneous landscape. cant animals
beta rarity Rarity that occurs in species that are habitat chemical alteration A form of water pollution or water
specialists which are abundant in one (optimal) environ- contamination caused by organic or inorganic substances.
ment, but rare or absent from environments which mani- It can take many forms, but two of the most common are
fest even slight changes in one or more critical variables, eutrophication and acidification.
and therefore show a pattern of overall rarity across a chemical manipulation A form of human intervention
landscape. within aquatic systems involving the conversion of
bioclimate envelope model Models that characterize contaminants to other chemical states, or the use of
associations between aspects of climate and species’ chemicals to prevent contaminants from entering or
occurrences to estimate the conditions that are suitable to interacting with the system.
maintain viable populations, such as minimum and maxi- climate niche The climate component of an organism’s
mum temperature. fundamental niche, or the sum of a species’ tolerance to
biodiversity The entire array of earth’s biological variety, ranges of climate variables.
contained in genes, populations, communities, and climate velocity A method to map the pace of climate
ecosystems. change across space, characterized by the equation
biodiversity hotspot A relatively small area with a dispro- ( Cyr-1/ Ckm-1 ¼ kmyr-1).
portionately high level of biodiversity in multiple taxa. club goods A type of good in neoclassical economics,
biological species concept The idea that species are defined sometimes classified as a subtype of public goods, that
by reproductive isolation. By this definition, species are are excludable but non-rivalrous. An example might be a
organisms that breed together to produce viable offspring movie theater ticket, where the price of the ticket restricts
resembling the parents, but do not breed and reproduce access but broadly speaking, a high number of moviegoers
viable offspring with other species. does not take away from the value of the film.
bio-manipulation Direct alteration or manipulation of an coercive measures In the context of international law,
ecosystem with species additions or removals. Often a sanctions, penalties, or loss of membership in international
procedure based on enhancing the suppression of phyto- organizations or of privileges in international dealings
plankton by zooplankton, and control of fish populations which are used to motivate unwilling parties to comply
in a manner so as to achieve efficient phytoplankton with legal standards, such as international environmental
reduction by zooplankton filtration. agreements.
biotemperature In the Holdridge Life Zone classification co-management A conservation strategy that attempts to
system, the mean value of daily temperature above 0 C simultaneously address biological, cultural, economic,
divided by 365, which provides a measure of the heat and political concerns through collaboration and integra-
available during the growing season. tion of conservation efforts between local communities
blast fishing The harvesting of fish through the use of and government authorities.
explosives. Blast fishing, when employed near corals, common pool resources Resources or goods where one
leads to destruction of coral reefs. user’s consumption subtracts from the supply available
Glossary 573
for others, yet the physical nature of the resource makes it communicating such purpose includes understanding
difficult to exclude users. how human values, beliefs, attitudes and behaviors
community-based ecotourism (CBET) Travel to and rec- towards biodiversity work themselves out in organizations
reation in natural environments by nonresident individuals and socio-political structures.
(tourists) for the purpose of enjoying the biodiversity or conservation reliant species A species of conservation con-
ecosystem characteristics of such environments but cern that requires continuous management intervention,
practiced in such a way that it conserves the environment even after a self-sustainable population is achieved;
and sustains the wellbeing of local people. threats cannot be completely eliminated, only managed.
compliance In the context of international law, the extent to conservation triage A term used to describe the process of
which the behavior of a state, as a party to an international determining which species or subpopulations to abandon
treaty, actually conforms to the conditions of the treaty. (usually approaching unavoidable extinction), in order to
compliance information systems In the context of interna- focus limited resources on others with higher chances of
tional law, systems that are built into government survival. This approach may also be applied in allocating
structures and whose aim is to ensure compliance and conservation efforts between sites or among program
report non-compliance. types.
conceptual model A visual or narrative summary that contingent valuation (CV) The assignment of economic
describes or identifies important components of a system values to nonmarket goods through analytical methods
and the possible interactions among them. that determine an individual’s willingness to pay for
connectedness The presence of physical linkages between such goods or willingness to accept compensation for
landscape elements. their loss.
connectivity – See also habitat connectivity, hydrologic converters See parameters
connectivity A parameter of landscape function that coral bleaching A phenomenon that occurs in corals
measures the processes by which subpopulations of exposed to extended periods of elevated water
organisms are interconnected into a functional demo- temperatures resulting in the loss of the cells and pigments
graphic unit, achieved only if organisms actually move of symbiotic dinoflagellates (zooxanthellae) that live
between connected units. within the coral.
connectors – See also sink, source, and stock In a model, corridor A pathway that connects habitat patches across a
elements that explicate or display the path through which landscape often fragmented by human activity and
material or individuals are transferred from one stock to facilitates organism movement among them.
another or from sources to sinks. cost benefit analysis (CBA) An economic tool used to
conservation easement – See also zoning A special case of transparently compare measure the benefits of a decision
land-use zoning applied specifically to conservation and with the costs associated with taking that action; the tool
developed to make the value of conservation on private is often used to compare the costs and benefits across a
land more explicit and profitable to landowners. In an range of policy options. Often ecosystem goods and
easement, the landowner agrees to restrict some activities services are not included unless they are assigned a mone-
or forms of development on his or her land to achieve tary value.
specific conservation goals. Such restrictions lower the critical habitat As defined by the US Endangered Species
assessed value of the land, generating a reduction in prop- Act, the specific areas within a geographic area, occupied
erty taxes for the owner and a reduction in inheritance by the species at the time it was listed, that contain the
taxes for the owner’s heirs. physical or biological features that are essential to the
conservation management agreement (CMA) An agree- conservation of endangered and threatened species and
ment between two or more private or public actors defined that may need special management or protection.
by four elements: (1) a partnership agreement enabling critical threshold See percolation threshold
effective management actions, (2) a management plan crucial habitats Season-specific use areas, population
informed by critical elements of species recovery, (3) suf- sources areas, movement paths, or some portions of
ficient funding and (4) legal enforcement. annual home ranges for populations within protected areas
conservation management unit (MU) A population or
group of populations that show evidence of genetic relat-
edness, but also are arranged spatially in such a way that D
they can benefit from a common management strategy.
conservation mission As a discipline, conservation is damage costs avoided Method which uses either the value
mission-driven in that it proceeds with purpose; of property protected, or the cost of actions taken to avoid
574 Glossary
damages, as a measure of the benefits provided by an another, measured in terms of rate (proportion of
ecosystem. individuals that leave the natal area), distance (how far
data mining See mining of legacy data an organism travels from the natal area before it resumes a
deadweight social losses Distortions in economic efficiency settled existence) and direction (which way is the dispers-
and market function caused by the diversion of earned ing individual going).
income to the government through taxes. distance effect – See also decay value The rate of dissipa-
decay value – See also distance effect A variable some- tion of the edge effect at a boundary between two different
times incorporated into models that attempts to simulate habitats over distance from the edge into the interior of the
effects of edge influences. The larger the decay value, the habitat.
faster the variable changes per unit distance from the edge. dredging Direct removal of sediment from a lake, pond, or
Low values indicate little decline over a long distance. wetland, usually by physically scraping it from the bottom
High values indicate rapid decline in effect over a short with large, earth-moving machines.
distance.
decision analysis A management approach that attempts to
determine the probability of different population events, E
such as persistence or extinction, that would result from
specific management decisions, actions or strategies. ecocentrism An ethical position asserting that the value of
deme A subdivision of a population consisting of closely land and its component plants and animals is derived from
related individuals typically breeding mainly within the their contributions to the function and integrity of the
subdivision. ecosystem and its processes; therefore, the highest good
demersal species In aquatic environments, bottom-dwelling is to preserve the functional integrity of the ecosystem.
species or “groundfish” such as flounder and haddock. ecological drainage unit (EDU) Regional biodiversity
demographic monitoring Monitoring approach which distinctions within aquatic zoogeographic units (AZUs),
follows the fates of individuals in a population over time generally 1000–10,000 km2 in size, which are delineated
and makes repeated on-site visits to gain direct demo- and classified by identifying areas with similar biotic
graphic measurements of the population. patterns.
demographic stochasticity Random fluctuations in birth ecological economics A school of economic theory
and death rates, emigration and immigration, or sex ratio designed to achieve a union of economics and ecology
and age structure of a population. such that the economy is conceived as a subset of the
density dependence A negative feedback relationship global ecosystem sustained by the flow of energy and
between population growth rate and population size material from and back to that ecosystem.
resulting in the regulation of the population fluctuations ecological flows Movements of material, organisms, and
around a mean population size. energy that are waterborne or airborne, as well as
density independent A population whose growth is solely a movements of disturbances such as fire.
function of chance events (independent of density) which ecological succession A pattern of continuous, directional,
act on increasing or decreasing the size of the population. nonseasonal change of plant populations on a site
deterministic factors Factors that affect a population in a over time.
constant relation to the population’s size. economic institutions A diverse set of rules (formal or
diadromous fish species A species which moves from fresh informal) to govern economic behavior, developed and
water to saltwater in their normal life cycle, such as the enforced by a diverse set of actors.
Atlantic sturgeon and sea lamprey. ecosystem A biological community of interacting
differential fitness species concept Species defined as organisms and their physical environment.
groups having features that would have negative fitness ecosystem-based management (EBM) A management
effects in other groups and cannot be regularly exchanged approach that defines management strategies for entire
between groups upon contact. ecosystems, not simply individual components of
direct payment (DP) – See also payments for ecosystem ecosystems; as a consequence, EBM takes into account
services In conservation, payment to individuals or local interactions among ecosystem components and manage-
communities for protecting or restoring ecosystem ment sectors, as well as cumulative impacts of a wide-
services which benefit others. spectrum of ocean-use sectors.
dispersal The movement from an individual’s natal area to ecosystem management A pattern of prescribed, goal-
one of more permanent residence, usually as an adult, or oriented environmental manipulation that (1) treats a
the permanent movement of an individual from one area to specified ecological system as the fundamental unit to be
Glossary 575
managed; (2) has a desired outcome of assuring the per- endemism Restriction of a species to a particular area or
sistence of historical components, structure, function, region; in some areas, like islands, endemism is character-
products, and services of the system within biological istic of nearly all species.
and historical ranges and rates of change over long time enforcement In the context of international law, actions
periods; (3) uses naturally occurring, landscape-scale pro- used to force states to first implement and then comply
cesses as the primary means of achieving management with international laws and agreements.
objectives; and (4) determines management objectives entropy – See also ecological economics A measure of the
through cooperative and deliberative decision-making by amount of unusable energy in a system; as entropy
individuals and groups who reside in, administer, or have increases, the amount of energy available for work
vested interests in the state of the ecosystem. decreases; according to ecological economics, the econ-
ecosystem monitoring Periodic, regular measurement and omy uses low entropy energy and matter from its
tracking of indices of ecosystem system properties and surrounding natural environment (such as coal or oil), to
status in a given ecosystem; a critical tool for managing produce consumption goods, and discards high entropy
uncertainty in attempting to describe or predict the behav- wastes and dissipated heat back into the environment
ior of ecosystem processes. (such as carbon dioxide).
ecosystem properties Natural structures and processes of Environmental Kuznets Curves (EKC) A family of
ecosystems, which form the basis of any kind of ecosys- graphical representations of the relationship between envi-
tem services utilizable by humans. ronmental deterioration and per capita income. A typical
ecosystem services Subset of natural ecosystem process that Kuznets Curve shows increasing environmental deteriora-
generate benefits to people (e.g., water purification). tion with increasing income to an “inflection point” or
ecotourism See community-based ecotourism (CBET) “turning point income,” past which further increases in
edge effects A suite of processes and factors associated with income are associated with declining levels of environ-
edge environments which become more pronounced when mental deterioration.
habitat is fragmented and the relative amount of interior environmental stochasticity Fluctuations in the probability
(non-edge) habitat decreases. of birth and death in a population because of temporal
edge influences See edge effects variation in habitat parameters, climatic variation,
effective population size The size of an “ideal” (randomly competitors, parasites, predators, diseases or other envi-
mating) population that would undergo the same amount ronmental factors external to the population.
of genetic drift as a particular real population. ethics Systematic organizations of values that establish
effectiveness In the context of international law, the degree principles for conduct and behavior.
to which the fulfillment of the conditions of a given law, eutrophication The process in which the release of
treaty or convention actually achieves the objectives of it. nutrients, particularly phosphorus, into streams, lakes, or
Ehrlich Identity An expression of the relationship between estuaries triggers a chain of events resulting in oxygen
human population, human resource consumption, and depletion, turbidity, and radical alteration of the biological
human technology and environmental impact, formally community.
expressed as I ¼ P A T; where I is environmental evolutionary species concept The idea that a species is one
impact, P is population, A is affluence (a measure of lineage evolving separately from other lineages.
consumption), and T is technology (an index of efficiency excludable goods Goods or resources in which ownership
of resource use and pollution abatement). The Ehrlich permits the owner exclusive use of the goods or resources
Identity asserts that environmental impact is a function and provides the owner with the ability to exclude others
of the combined effects of human population density, per from such use.
capita consumption (A) and efficiency of resource use (T ). existence value The value derived from knowing that some-
elasticity analysis A type of sensitivity analysis that thing exists.
determines the effect of a variable on model outcomes; exotic species Also known as alien species, invasive spe-
the degree to which a change in the value of a model cies, non-indigenous species, and bioinvaders, these are
variable changes the value of D, the population’s rate of species of plants or animals present in a nonnative
growth, in relation to other model variables. environment.
emigration – See also dispersal The act of leaving one’s exponential growth A model of population growth in
current area or population to enter another, a form of which population size increases at an ever- increasing
dispersal. rate, and only the population’s size (N ) and intrinsic rate
576 Glossary
of increase (r) determine the change in numbers of fragmented habitats and landscapes A condition in which
individuals. areas of vegetation with similar characteristics are
externalities – See also market failure An economic cost separated from one another by matrix habitat which differs
or benefit that is incurred by a third party with no control from habitat characteristics of this vegetation.
over its creation; as such these costs and benefits are not fund-service resources – See also stock-flow resources
explicitly captured in a decision or action and left outside Resources that can only be used a given rate, and whose
of the market. A negative externality might be pollution production is measured as an output per unit time and
while many ecosystem service represent positive therefore cannot be stockpiled; the maintenance of water
externalities. quality by an intact forest is an example of a fund-service
extractive reserve An area of land, generally state-owned resource.
where access and use rights, including natural resource
extraction, are allocated to local groups or communities.
extinction Permanent loss of entire species because of envi- G
ronmental forces (habitat fragmentation, global cli-
mate change, natural disaster, overexploitation of species GAP analysis Originally an acronym for the Gap Analysis
for human use) or because of evolutionary changes in their Program (GAP), a type of analysis that determines,
members (genetic inbreeding, poor reproduction, decline through the use of computer overlay maps and other
in population numbers). forms of spatial analyses, whether populations of species
extrinsic factors Environmental or ecological events and targeted for conservation fall within the boundaries of
processes external to the population that influence popula- currently protected areas and which elements of landscape
tion growth over time. biodiversity are underrepresented in reserve systems. The
final outcome of a GAP analysis is to attempt to identify
the “gaps” in the conservation reserve network where
F significant biodiversity resources have been left unpro-
tected in the landscape.
FBO A faith-based organization engaged in conservation GIS An acronym for Geographic Information Systems, or
that uses and applies religious principles and teaching to sometimes Geographic Information Science, that
solve conservation problems, formulate management designates a computer-assisted system designed to cap-
decisions, or determine long-term conservation strategies. ture, store, manipulate, analyze, manage, and present spa-
factor resolution A population monitoring tool in which tial or geographic data.
experiments are conducted to determine which factors GONGO A Governmental and Non-Governmental Organi-
actively limit population growth. zation (GONGO), such as the World Conservation Union
fecundity The number of gametes produced per female per or International Union for the Conservation of Nature
unit time. (IUCN), whose members and contributors are nations
fixation index The proportion of genetic variance contained and organizations rather than individuals.
in the various subpopulations relative to the genetic vari- gamma diversity The total number of species recorded for
ance in the total population. the group of sites or communities that make up a land-
fixation of deleterious alleles A condition in which all scape; gamma diversity is the product of the alpha diver-
individuals in a population possess only the harmful allele sity of a landscape’s communities and the degree of beta
among multiple alleles at a particular locus, such that the differentiation among them.
trait or traits associated with the allele become permanent gamma rarity Species that may have large populations in
or “fixed” in the population. The risk of such fixation local communities and demonstrate broad environmental
increases with decreasing population size. tolerances, but are restricted to particular geographic areas
floodplain A low-lying area adjacent to a river with its and so become increasing rare with increasing distance
species and systemic productivity dependent on seasonal from their population centers.
pulses of flooding. gene diversity The probability that two alleles from the
flows See ecological flows same locus sampled at random from the population will
focused advocate A person or group reporting data not be identical by descent. Mathematically, Gene Diver-
concerning a subject in which he, she or they have exper- sity (GD) is equal to GD ¼ 1 MK, where MK is
tise as well as deeply held convictions, and who works to population mean kinship.
ensure that the information presented is correctly gene flow The effective movement of genes between
interpreted and rightly applied. populations or population subdivisions, which can be
Glossary 577
mathematically determined, in one way, from the expres- habitat connectivity – See also connectivity The degree to
sion N e m ¼ 1F
4F ST , where Ne is the effective population
ST which individuals in a population can move between
size, m is the rate of immigration and FST is the total spatially disjunct patches of the same kind of habitat in a
genetic diversity found among all populations. The result landscape, such that individuals in different patches func-
is expressed in immigrants per generation. tion as a single demographic unit (population).
generalized random tessellation stratified (GRTS) habitat conservation plan (HCP) Under the US
samples – See also spatially balanced sampling design Endangered Species Act, an agreement between a federal
(SBSD) Probability sample designed with reverse hierar- agency (usually the US Fish and Wildlife Service) and a
chical ordering such that for any sample size, n, the units private landowner or non-federal government land owner
in the sample will be spatially balanced (i.e., “spread that stipulates actions that will be taken by the landowner
out”), having coverage of survey effort over the target to enhance the population or habitat of an endangered
region. species in return for permitting some mortality of the
genetic drift Random fluctuations in gene frequencies that species on the property in the course of other activities
occur as a result of nonrepresentative combinations of carried out by the land owner (incidental take).
gametes during mating, especially in small populations. habitat degradation The gradual deterioration of habitat
genetic stochasticity Fluctuations in demographic quality, which can co-occur with habitat loss, fragmenta-
parameters, especially of small populations, through tion, or isolation, such that the habitat loses critical
increased rates of inbreeding, genetic drift, and accumula- resources or structures needed by individual species,
tion of unfavorable mutations. thereby affecting the amount of available habitat.
geographic-based approaches Approaches to biodiversity habitat fragmentation The process through which a single
conservation that focus on the qualities of habitat and block of contiguous habitat is broken into fragments of
landscape that sustain resident populations rather than on patches of similar habitat no longer directly connected or
the dynamics of individual populations. adjacent to one another. Habitat fragmentation is created
governmental subsidies Incentive payments that the gov- through habitat disturbance and alteration, and results in
ernment provides to individuals or firms to influence their increased amounts and proportions of habitat edge.
economic behavior; often encouraging pro-environmental habitat generalists Species that can exploit a variety of
and pro-conservation outcomes that would not otherwise habitats in a given geographic range and, thus, are rela-
result from market transactions. tively resistant to extinction through habitat loss or land-
gradient models Models of habitat distribution in which use changes.
different habitats are not clearly defined and environmen- habitat heterogeneity Differences in habitats, at a variety of
tal conditions change slowly and gradually at fine spatial spatial scales, which may be natural (due to a rich internal
scales. structure of differing habitat patches) or artificial (due to
greenhouse gases Atmospheric gases, notably water vapor fragmented habitats resulting from human activity).
(H2O), methane (CH4), ozone (O3), nitrous oxide (N2O) habitat isolation The separation of blocks of habitat from
and carbon dioxide (CO2), that, because of their capacities other blocks of similar habitat, a result of habitat
to absorb infrared radiation radiated from the surface of fragmentation.
the Earth, create a net influx of energy to the Earth and its habitat loss Physical destruction of habitat for a particular
atmosphere. species by human activities.
habitat specialists Species that are typically highly success-
ful in only one or a few types of habitat and, thus, are
H vulnerable to extinction through loss of their preferred
habitat.
habitat The physical surroundings of an organism, habitat structure – See also habitat complexity and habi-
characterized by both physical and biological features, in tat heterogeneity The amount, composition, and three-
which a species can persist, whether consisting of discrete dimensional arrangement of biotic and abiotic elements
patches or of gradients with indiscernible boundaries. where an animal lives as a specific time and location.
habitat complexity The relationship of scale to the amount habitat suitability model (HSM) Model developed to pre-
of physical components (for example, food resources) in dict species occurrence based on species-specific environ-
the habitat. mental variables; however, predictions of HSMs are not
habitat configuration The spatial arrangement of habitat the same as actual species distributions.
and the proximity and relatedness of different parcels of haplotypes Mitochondrial DNA (mtDNA) groups that can
habitat to one another in the same landscape. be used to determine rates of gene flow among populations.
578 Glossary
hard law In the context of international law, formal expressed as days per year); (2) desiccation frequency
conventions and treaties adopted by many nations, with (how often the site dries up); and (3) predictability or
explicit mechanisms for enforcement. regularity of alternating flooding and desiccation events.
harvest refugia – See also no-take zones Areas, usually in
marine environments, designed to protect a particular
commercial stock or group of stocks from over- I
exploitation by keeping areas closed to all types of fishing
and harvesting. implementation In the context of international law, specific
headwaters The source of a river or stream; the furthest actions taken to make international treaties operational in
place in that river or stream from where it empties or their own national legal system.
converges with another river. immigration In population demography, the addition of
heat capacity The number of heat units needed to raise the individuals to a population through the entrance of new
temperature of a system or body by one degree; water has individuals from another population or populations
the highest heat capacity of any liquid. inbreeding The mating of individuals with close relatives,
hedonic property model A model of property value that with whom they may share many genes.
treats such value as a function ( f ) of its structural inbreeding as nonrandom mating A quantitative measure,
characteristics (S), neighborhood (N ), and environmental FP, of genetic drift in a population, where FP represents
quality (Q), expressed in the identity Pi ¼ f(Si, Ni, Qi), the average probability of inbreeding by descent, a mea-
where the subscript i refers to each value for an individual sure of the effect of genetic drift on a population relative to
(ith) property. an “ideal” population experiencing completely random
hedonic travel cost method See travel cost method mating.
heterozygosity – See also average heterozygosity In the inbreeding by descent See inbreeding as nonrandom
context of genetics, the condition of or degree to which mating
individuals in a population carry two different alleles of a inbreeding by population subdivision – See also panmictic
gene for a particular trait, one from each parent, at the index Inbreeding quantified as the panmictic index, f,
corresponding loci of a pair of chromosomes. which measures inbreeding as a deviation from a reference
homozygosity In the context of genetics, the condition of or population which has a system of mating in which alleles
degree to which individuals in a population carry identical at a locus are paired in proportion to their frequencies in
alleles of a gene for a particular trait at the corresponding the overall population (by definition, random mating). The
loci of a pair of chromosomes. panmictic index evaluates deviations from heterozygosity
hotspot See biodiversity hotspot frequencies expected under random mating through the
hybrid Offspring of matings between individuals of differ- relationship f ¼ 1 HO/He, where He is expected hetero-
ent species, subspecies, or populations. zygosity under random mating and Ho is observed
hybridization The mating between individuals of different heterozygosity.
species, subspecies, or populations; often a waste of repro- inbreeding coefficient The probability that two alleles at the
ductive effort by potential parents, and especially threat- same locus in an individual are identical by descent.
ening to rare species due to reduced fitness of hybrids. inbreeding depression A sequence of events initiated by
hydric (water-logged) soils Soils formed under conditions matings between closely related individuals, especially in
of saturation, flooding or ponding long enough during the small populations of normally outbreeding species,
growing season to develop anaerobic conditions as well as whereby heterozygosity and fecundity are reduced and
distinct structural and chemical characteristics in the upper mortality is increased through expression of deleterious,
portions of the soil. recessive alleles.
hydrologic connectivity The water-mediated transport of inbreeding load The proportional decline in survival that
matter or organisms within or between elements of the can be attributed to inbreeding of a given magnitude.
hydrologic cycle. incidental take – See also taking Under the US
hydrology The science and study of the occurrence, move- Endangered Species Act, harm or harassment done to a
ment, and storage of water in the earth system. protected species that is incidental to, and not the purpose
hydroregime The total contribution of different hydrologi- of, carrying out an otherwise lawful activity.
cal variables to the stability of a temporary aquatic habitat indicator species – See also surrogate species A species
including, but not limited to, (1) hydroperiod (the length whose conservation status is assumed to reflect the status
of time there is standing water on the site, usually of other species with which it shares the community.
Glossary 579
institutions See economic institutions intrinsic factors In population demography, characteristics

instrumental value The value of something that is realized internal and specific to a population itself that influence or
through its usefulness, utility, or instrumentality to meet determine its demographic traits.
the needs or promote the good of another; a value that is intrinsic value The value of something in and of itself,
realized in an object or entity through its use for some without regard to its usefulness or utility to others.
other purpose. introductions of exotic species The acts or processes, via
intact habitats and landscapes Habitats and landscapes direct or indirect human action or natural events, through
characterized by contiguous natural vegetation covering which species from one area rapidly enter a new area
more than 90% of their land surface area. where they did not previously occur. The introduction of
integrated conservation and development projects exotic species is a leading cause of global species endan-
(ICDPs) Projects in which local people share benefits of germent and biodiversity loss.
using or harvesting plant or animal resources in their introgression The long-term acquisition and incorporation
environment at sustainable levels, take ownership of the of genetic material from one species into the genome of
conservation of such resources, and have an active role in another species, especially when individuals of a rare
decisions affecting the use and management of these species hybridize with those of a closely related, but
resources in ways that benefit them economically and more numerous species.
culturally. ICDPs generally attempt to achieve conserva-
tion as an outcome of sustainable community develop-
ment rather than through exclusive focus on J
management or protection of the resource to be conserved.
integrated ecosystem assessment (IEA) A framework used Judeo-Christian Stewardship Environmental Ethic An
for organizing information needed to guide ecosystem- organized system of values for environmental care based
based management (EBM). An IEA is created in a five- on principles taught in the Old and New Testaments of the
step process including (1) an abstraction of the ecosystem Bible, including especially a view of the intrinsic good-
into subsystems relevant to management issues; (2) an ness of created things, the responsibility of human beings
issue identification and scoping process that identifies to care for and protect the non-human world, and the
specific ecosystem management drivers and pressures; inclusion of non-human creation in the redemptive plans
(3) an identification of indicators, most sensitive to such and purposes of God.
drivers and pressures and thus provide the basis for assess-
ment of the status and trends of the ecosystem; 4) a risk
analysis that evaluates the probability of harm to the K
ecosystem indicators posed by human activities and natu-
ral processes; and (5) ongoing monitoring and assessment kinship coefficient A measure of the degree of relatedness
of ecosystem indicators. between two individuals, expressed as the probability that
integro-difference equation (IDE) models A category of alleles randomly selected from homologous loci in such
models that describe the predicted spread of an invasive individuals are identical by descent from a common
species by breaking dispersal and population growth into ancestor.
separate stages.
interior habitat Habitat sufficiently distant from an edge
(border of two or more different habitats) as to experience L
negligible effects of edge influences.
International Union for the Conservation of Nature landscape-level conservation An approach to conservation
(IUCN) An international conservation organization, that focuses on landscape integrity and connectivity,
also known as the World Conservation Union, whose including privately owned lands, urban areas, and agricul-
members consist of nations or organizations rather than tural areas, taking into account human dependence on
individuals. IUCN declares its vision to be that of creating resources.
“a just world that values and conserves nature”, and its landscape A large area that is comprised of more than one
mission to “influence, encourage and assist societies type of habitat distributed in numerous discrete and dis-
throughout the world to conserve the integrity and diver- tinct patches.
sity of nature and to ensure that any use of natural landscape genetics The study of the interaction between
resources is equitable and ecologically sustainable.” loss of genetic diversity and habitat fragmentation.
580 Glossary
lentic systems Freshwater aquatic environments character- market, price, and other economic variables to provide behav-
ized by non-moving or slow-moving water, such as lakes ioral incentives. Some examples of market-based economic
or ponds, rather than flowing water (lotic) environments policy solutions are tradable permits, taxes, and subsidies.
characteristic of streams. market equilibrium A market state where the supply in the
lethal genes Genes which, although recessive and unex- market for a good or service is equal to the demand in the
pressed in a heterozygous state, will, in a homozygous market. The equilibrium price is the price of a good or
condition, result in the death of the individual. service when the supply of it is equal to the demand for it
lethal load The proportion of lethal genes in a population, in the market.
which often rises when alleles are lost during a period of market failure – See also externalities A market system
population reduction. condition that occurs when the price mechanism fails to
limit indicators – See also performance indicators In account for all of the costs and benefits necessary to
ecosystem management, variables that detect or foretell a provide and consume a good or service. The market fails
point at which a critical resource begins to come under because it does not supply the socially optimal amount of
stress. When the level of the variable reaches a the good or service.
pre-determined threshold or critical value, the manager is market goods Things that can be traded in standard
facing an unacceptable risk of harm to the resource or the currencies of exchange, such a money, in normal arenas
system that produces it and should take immediate action. of exchange, such as markets.
limited access A mitigation strategy designed to reduce the marketplace See market goods
effects of human activity and movement to and from sites matrix habitat Habitat which separates non-contiguous
within wildlife habitat by restricting use of roads to the site “pieces” or “fragments” of a given type of natural or
as well as actual use of the site to essential activities reference habitat and differs, to varying degrees, from
performed by a relatively small number of designated characteristic of natural or reference habitat.
individuals. maximum-avoidance-of-inbreeding (MAI) strategy A
living modified organism (LMO) An organism whose strategy of captive breeding management which avoids
genomes have been engineered or “spliced” so as to incor- mating between relatives. In an MAI strategy, managers
porate genes from other, usually very different, kinds of examine kinship between potential mates, which is equiv-
organisms in order to preserve, enhance, or add traits alent to the inbreeding coefficient of potential offspring of
favorable to increased production, fertility, survivorship, the pair, and pair individuals with little or no kinship to
or adaptability to particular environmental conditions. one another (i.e., pairings that produce offspring with a
Also known as genetically modified organism (GMO). low or zero inbreeding coefficient).
logistic growth A model of population growth in which maximum sustained yield – See also sustained yield
population size increases at a decreasing rate as it forestry The largest amount or level of removal of a
approaches an upper asymptote, set by environmental particular resource from a system that can be taken indefi-
limits. nitely for successive time increments without depletion of
lotic systems Freshwater aquatic environments characterized the resource or loss of productivity of the system. If
by flowing water (lotic) such as is characteristic of streams. correctly determined, the maximum sustainable yield is
equal to the regenerative rate of the resource in that system
in each time increment.
M mean generation time The average age at which animals
produce offspring.
macroeconomics A branch of economics that studies the mean kinship The average kinship between a single indi-
behavior and performance of an economy as a whole. It vidual and all other individuals in the population.
focuses on the aggregate changes in the economy such as mean kinship (MK) strategy A strategy of captive breed-
unemployment, growth rate, gross domestic product and ing management in which individuals with similar mean
inflation. kinship values are paired for breeding, especially if such
macrohabitat In the habitat classification system used by values are low, leading to the production of offspring with
The Nature Conservancy, a system of finer scale classifi- an increased representation of rare alleles.
cation units that can be used to define an Aquatic Ecolog- mentor A trusted counselor, guide, tutor or coach, particu-
ical System which may be a target as a potential protected larly in one’s own profession, who takes active interest in
area. the welfare, development, and advancement of a younger
market-based approaches – See also economic institutions; or less experienced colleague or student.
marketplace A suite of economic institutions that use the
Glossary 581
metapopulation A population that exists as spatially dis- N

junct subunits at different densities in habitat patches of
varying carrying capacity. natural catastrophes Extreme forms of normal environ-
metapopulation theory A conceptual model to describe mental variation (e.g., flash floods or severe and prolonged
collections of subpopulations of a species in a given drought) or unusual and infrequent natural events (e.g.,
area, each occupying a suitable patch of habitat in a forest fire, volcanic eruption) that have the potential to
landscape of otherwise unsuitable habitat. eliminate all individuals in a small population.
micro-colony fusion A technique used in coral reef restora- natural resource rights A category of property rights
tion in which fragments of different colonies can be fused which establish rules ogverning activities that individuals
together to produce larger colonies with faster rates of or groups may exercise with natural resources including
growth. defining access to the resource, withdrawal (the right to
microeconomics A branch of economics that analyzes the remove the resource from the system for other use), man-
economic tendencies of individuals, households, or firms agement, exclusion (defining who may not have access to
in decision making and the allocation of resources. It the resource), and alienation (the right to sell or lease other
applies specifically to markets for goods and services. rights to other parties).
microsatellites A type of satellite DNA that consists of naturalistic fallacy The error of arguing for an imperative
short tandem repeats 2–4 nucleotides long, and whose conclusion or normative action based on a descriptive
variability is useful in determining pedigrees of premise; arguing from “what is” to “what ought to be.”
individuals. In science, attempting to prove an imperative or ethical
minimum viable population (MVP) The minimum num- conclusion based on a fact of nature.
ber of individuals required for a population to persist for a neighborhood rules – See movement rules; neutral land-
specified length of time at a specified level of probability. scape models A landscape model in which the distribu-
mining of legacy data The systematic search and interpre- tion of habitats in the landscape is a theoretical distribution
tation of past records to inform present conservation man- of habitat independent of (i.e., “neutral” toward) actual
agement decisions or policies. biophysical processes that shape landscapes.
minisatellites A type of satellite DNA that consists of neutral landscape models Landscape models in which
sequences up to 100 base pairs long, and whose variability arrangement of habitats is independent of (i.e. “neutral”
forms the basis for DNA fingerprinting. toward) biophysical processes that shape landscapes.
mitigation In conservation, the lessening of the effects of no surface occupancy A method of mitigation, often used
human disturbances on populations, habitats, and in mining and drilling operations, in which humans do
landscapes. not extract resources directly from beneath high quality
movement rules Rules used in landscape models which habitat, but remove the resources through directional
specify the distance across which sites are accessible to (side) drilling from a more remote site in a lower-quality
organisms by virtue of their dispersal or gap-crossing habitat.
abilities. no-take zones – See also harvest refugia In marine
moral agent An entity capable of discerning between right environments, areas designed to protect a particular com-
and wrong, and therefore considered morally responsible mercial stock or group of stocks from over-exploitation by
for its actions and their consequences. prohibiting any harvest (i.e., “take”) within the
moral subject An entity that can be treated rightly or designated zone.
wrongly in a moral sense, even though the entity itself non-excludable goods and services Goods and services for
might not be not capable of acting in a morally right or which it would be extremely difficult and costly to exclude
wrong manner. anyone from receiving the benefits (e.g., protection from
multiple use A management approach, initially developed ultraviolet radiation by the ozone layer).
and practiced by the US Forest Service, that manages non-rival goods and services Goods and services whose
lands and ecosystems for multiple objectives simulta- use by one person does not reduce or restrict use by others
neously, such as wildlife conservation, timber production, (e.g., breathing oxygen from the atmosphere).
and human recreational opportunity. nonspatial models Models in which knowledge of spatial
multiple-use module An approach to habitat conservation locations of entities of interest is not known, nor are
in which a fully protected core area is surrounded by processes and transfer rates affecting movement from
concentric zones of natural areas used in progressively one point to another. Nonspatial models are often used
more intense fashion for recreation and commodity to predict the spread of invading organisms by using
production. projections derived from population demography, such
582 Glossary
as exponential or logistic growth equations, or to predict P

changes in numbers of the invasive species through time.
null model Models, particularly in population demography, panmictic index A measure of inbreeding as a deviation
that assume that changes in the variable of interest reflect from the heterozygosity frequency expected under random
random events. mating. Expressed mathematically, the panmictic index, f,
is 1 HO/He where He is the expected heterozygosity
under random mating and H0 is the observed heterozygos-
O ity in the population.
parameters Values of variables that determine rates of flow
occupancy estimation and modeling Occupancy estima- or movement of resources or individuals from one state to
tion is the estimation of the probability of detecting the another or from one area to another.
occupancy of sites or areas by individual organisms or partnerships In ecosystem management and conservation,
groups of organisms that are imperfectly detected. Occu- dynamic relationships among actors with vested interests
pancy modeling is a method of estimating the density or in a system or resource based on mutually agreed
abundance of individuals in an area based on making objectives and pursued through an understanding of divi-
multiple visits to some or all of the sample sites sion of labor based on the respective comparative advan-
representing that area to determine occupancy and detec- tage of each member.
tion probability, which are then used to adjust for patch The fundamental unit of a landscape, containing only
estimates of distribution, density and abundance of the one type of habitat.
sampled species or group. patch contrast The degree of difference between the two
occupancy theory A branch of conservation science adjoining habitats. The greater the level of contrast
concerned with estimation of the probability of a species’ between the habitats, the greater the effect of edge
or organisms’s occupancy of sites or areas and the proba- influences on both habitats.
bility of detection of individuals on the site or area in order patchiness A quality of habitat arrangement, manifested as
to compensate for underestimation of occupancy and contrasting, discrete states of physical or biotic
detection that result from imperfect detection methods. phenomena.
oligonucleotide A short piece of DNA used in the polymer- patch models Models of habitat distribution or habitat use
ase chain reaction. by organisms which arrange habitats in patches, i.e., small
opportunity cost The best alternative that is given up or lost areas within landscapes which contain only one type of
when a choice is made. For example, costs or losses habitat.
associated with the inability to use a resource to produce payments for ecosystem services (PES) A conservation
goods A, B and C if the resource is used to produce good strategy in which an individual or group receives direct
D. payment from a conservation organization or government
optimal niche gestalt An approach to habitat management, agency for providing a specified, contracted ecosystem
based on the idea that identifiable structural features of an service, such as carbon sequestration, soil stability, or
environment allow a species to thrive, rather than merely enhancement of site-specific plant or animal biodiversity,
persist, and that such features can be recognized and usually on land owned or controlled by the individual or
identified by definitive criteria. group receiving the payment. Often such services are
option value The value of a resource’s expected future use, quantifiable (i.e., price per ton of carbon dioxide
or what a person would be willing to pay to guarantee that sequestered).
the resource would be available for future use. pedigree analysis An analysis used in captive breeding and
outbreeding depression A decline in fitness that occurs population management in which managers determine
when individuals from normally inbreeding populations the relatedness of individuals in the population to one
breed with individuals from other populations of the same another by determining each individual’s parental ances-
species, breaking up uniquely coadapted genetic try and subsequent offspring, and then use such analysis to
combinations and resulting in subsequently reduced fit- inform choices of pairings for reproduction of new
ness and fecundity. offspring.
overexploitation A level of harvest of a population that pedigree inbreeding Inbreeding by descent, or the measure
cannot be replaced by the population’s own recruitment of an individual’s ancestry shared in its maternal and
and is therefore not sustainable over time. paternal lines.
Glossary 583
pelagic species In marine environments, free-swimming physical habitat alteration The physical conversion of
species that may disperse and move widely throughout habitat to unusable non-habitat (habitat loss), breaking
ocean waters. large, contiguous blocks of habitat into smaller patches
percolation cluster In habitat and landscape ecology, a (habitat fragmentation), increasing separation of blocks of
single group of habitat patches in sufficient proximity to habitat from one another (habitat isolation), or changes in
one another such that organisms can move throughout the habitat that affect composition, structure, or function (hab-
entire system (i.e., “percolate” from patch to patch). itat degradation).
percolation theory Originally a theory of physics to policy A set of principles and intentions used to guide deci-
describe and explain the physical properties of gels, sion making; in environmental conservation, a set of
polymers, and glassy materials, particularly as a means principles and intentions used to guide decision making
to understand the flow of liquids through material about human management of natural capital and environ-
aggregates. In conservation biology, a theory of landscape mental services.
ecology that provides a quantitative analysis of habitat polluter pays principle Economic strategies based on the
connectivity in spatially structured landscape systems to principle that polluters, rather than society, should pay for
describe and explain rates and types of movements of the pollution they create, thus preventing polluters from
organisms between habitat patches or other elements. externalizing pollution costs. Mechanisms associated with
percolation threshold - The level of disturbance in a land- such strategies can include pollution-specific fees, taxes or
scape (e.g., the proportion of sites modified, degraded or fines assessed on a polluter for exceeding prescribed pol-
destroyed relative to their reference state) at which the lution limits.
transition from a connected to a disconnected landscape polyculture cropping The practice of growing many crops
system occurs. Above the threshold value, the landscape is together on the same site, an agricultural technique that
considered to be connected (i.e., to consist of a single can contribute to enhanced biodiversity in agricultural
cluster of habitat spanning the entire system); below the landscapes.
threshold, the landscape is considered to be disconnected polymerase chain reaction (PCR) A genetic technique that
and to consist of numerous small clusters of similar but uses DNA polymerase to repeatedly copy (amplify) a
isolated habitat. short region of a DNA molecule for various types of
performance indicators In ecosystem management, variables analysis, such as direct sequence of the PCR products to
used to monitor the state or performance of an ecosystem, determine genotypes of individual animals.
such that when the indicator variable reaches a critical state polymorphism A genetic locus that has two or more forms
or value, it signals the need to initiate a predetermined (alleles). In a population or population subunit, polymor-
management action. phism is expressed as the probability of encountering a
performance payments – See also payments for ecosystem polymorphic loci among all loci in the population.
services (PES) In conservation, an arrangement in which pools In stream ecosystems, areas of deeper, slower water
money is given directly to individuals or communities for associated with conditions that are less photosynthetically
meeting specified conservation objectives. productive and where decomposition is the dominant eco-
persistence likelihood An estimate of the probability of logical process.
persistence or extinction of a population, often determined population A group of individuals of the same species that
as an outcome of a population viability analysis (PVA). is spatially, genetically, or demographically discontinuous
phenomenologically significant animals Animals that with other groups.
evoke strong emotional or empathetic responses in population bottleneck A drastic, temporary reduction in pop-
humans; often large animals with symbolic appeal such ulation size through catastrophe or dispersal of individuals to
as the elephant, panda and tiger. a new area, resulting in loss of genetic variation.
photic zone In an aquatic system, the depth to which light population demography The branch of science concerned
penetrates with sufficient intensity to permit photosynthe- with the study of a population’s size, composition and
sis. Also referred to as the euphotic zone. distribution across space and through time, especially
phylogenetic species concept The idea that species should through measurement and prediction of the quantity,
be defined by measuring genetic similarities, differences, rates, and probabilities of birth, death, immigration, emi-
and distances among populations or groups of gration and other events in a population which affect its
populations. A species, according to a phylogenetic spe- growth, composition, structure, and distribution, and the
cies concept, represents a group of organisms with an processes which affect these variables to induce change in
assumed or determined common ancestral lineage whose the population.
genetic similarities, differences, and distances are distin- population fitness The reproductive efficiency or success of
guishable from other such groups. a population or any relative or absolute measure thereof,
584 Glossary
often integrated and expressed by the symbol λ (lambda), professional advocacy The act of informing policy makers,
the population’s growth rate. managers and the public about issues that arise in one’s
population genetic diversity The degree, range, distribu- area of expertise through publication in scientific and
tion and array of genetic difference among individuals in a professional books and journals, and avoiding other,
population, often measured and expressed as the level of more direct or personal methods of information transfer.
polymorphism (P) (P ¼ number of polymorphic loci/total protected areas In conservation, areas with definable
number of loci), average heterozygosity (H ) (the average boundaries set aside for biodiversity conservation that are
proportion of individuals in a population that are hetero- recognised, dedicated and managed through legal or other
zygous for a particular trait), or allelic diversity (A) (the effective means to achieve long term conservation of nature
average number of alleles per locus among individuals in with associated ecosystem services and cultural values,
the population). usually through the exclusion or mitigation of human
population mean kinship The arithmetic mean of all indi- activities or presence detrimental to native biodiversity,
vidual mean kinships in a population. natural environments, ecosystem services or cultural values.
population viability analysis (PVA) The use of analytical protected area-centered ecosystems (PACEs) Ecosystems
or simulation models to make precise estimates of the which contain or are dominated by a protected area or
likelihood of species persistence within a defined time group of proximately located protected areas in which
period at a given level of probability, and to identify and threats to the PACE are identified using comprehensive
rank or weight threats to such persistence from specific scientific methods to map and analyze land-use changes in
causes. and around the protected area.
pre-mating isolating mechanism Any differences in distri- public goods Goods that are not easily transacted between
bution, appearance, anatomy, behavior, physiology or individual buyers and sellers, and that can be used and
genetics that prevent or discourage encounter, attraction, enjoyed by all in such a way that no individual can easily
copulation or fertilization associated with potential exclude others from the use, benefit or enjoyment of
matings between different species, subspecies, or the good.
populations.
Preservationism The philosophy and practice that conser-
vation should be advanced through the designation of Q
protected areas which exclude most human activities,
including and especially extraction of natural resources, quasi-option value The value of preserving options for
and most permanent human residence. future use, given an expectation of growth in knowledge
primary environmental ethic An organized system of or applied technology that might lead to a future, but
values that treats environmental entities, such as as-yet undiscovered or unrealized use for the resource.
non-human creatures or natural objects, as moral subjects
that can be treated or used rightly or wrongly by humans,
and thus the primary benefactors or victims of ethical R
decisions affecting their welfare.
private goods Goods that can be bought, sold and enjoyed radiative forcing A change in the energy balance of the
individually by private buyers and sellers, and which can earth-atmosphere system in response to a change in factors
be used and enjoyed by an individual in such a way that such as greenhouse gases, land-use change, or solar radia-
the individual can exclude others from the use or enjoy- tion. Positive radiative forcing refers to an increase in the
ment of the good. temperature of a system, such as the atmosphere, that
probability design In survey design for detection of occu- occurs as a result of absorption of energy, while negative
pancy and presence of organisms, survey techniques in radiative forcing refers to changes in the system that create
which each unit of the target population in the survey has a cooling.
known, non-zero probability of being included in the reaction-diffusion model A type of species invasion model
sample, and there is some random component to the in which populations travel as a wave of a given velocity
selection of sampled sites. (V ) determined by the population’s intrinsic rate of
production function In economics, an analytical method increase (r) and rate of movement or distance traveled
that relies on the fact that ecosystems may be inputs into (D), expressed as travel to a site for a particular activity,
the production of other goods or services that are them- to determine the value associated with a user’s preferences
pffiffiffiffiffiffi
selves marketed, such as fisheries. for such activity V ¼ 2 rD.
Glossary 585
recessive lethal alleles An allele which, although unex- Romantic Transcendentalism A view, popularized by
pressed in a heterozygous state, will, in a homozygous Ralph Waldo Emerson, Henry David Thoreau and other
condition, result in the death of the individual, which US essayists and philosophers that the highest and best
usually occurs in the zygote. use of nature was as a place of spiritual renewal, moral
reclamation The preparation and enhancement of degraded formation and encounter with the divine.
land to fulfill its former use or a new use.
recruitment The entry of young organisms into a population.
regional habitat representation A “coarse-filter” approach S
used by The Nature Conservancy for representing biodiver-
sity in regional conservation planning by attempting to safe harbor agreement – See also habitat conservation
include or represent examples of all regional habitats within plan (HCP) A type of habitat conservation plan (HCP)
preserves and protected areas in a given region. under which a landowner agrees to actively maintain
relictual habitat A habitat that covers a much smaller geo- suitable habitat (a “safe harbor”) for a predetermined
graphic area than it did in the past, often because of number of individuals of a species equal to the number
environmental change - a remaining relict of a once com- present on the site when the agreement is formulated. In
mon and widespread type of habitat. return, the landowner receives an incidental take permit
replacement cost The amount an entity would have to pay that authorizes future land-use changes or management on
to replace an asset at the present time, according to its other parts of the site that may be occupied by additional
current worth. When used in the evaluation of ecosytem individuals of the endangered species, and removes liabil-
services, it is not a measure of the value of the ecosystem ity for harm to those individuals of the species incurred
service itself, but the cost of replacing that service by some from the authorized activity of the landowner.
alternative means. satellite DNA Short, highly repetitive segments of DNA in
Resource Conservation Ethic A view, popularized by an organism’s genome with base sequences differing from
Gifford Pinchot, that the highest and best use of nature those of other forms of DNA.
was the management of its resources in the most efficient secondary environmental ethic An organized system of
way to achieve the greatest human good for the greatest values that treats environmental entities, such as
number of people in present and future generations. non-human creatures or natural objects, as morally neutral
revealed preference methods – See also stated preference agents or means to advance human welfare, such that the
methods A family of behavior-based, analytical ethical consideration of the effect of any decision on the
techniques that use metrics associated with specific condition or welfare of natural objects or nonhuman spe-
preference-driven behaviors, such as costs associated cies is secondary to or an expression of ethical
with travel to a site for a particular activity, to determine considerations of the decision’s effect on humans.
the value associated with a user’s preference for such sectoral management A type of ecosystem-based manage-
activity. ment (EBM) in which definably different systems or
riffles In stream ecosystems, areas of fast moving, turbulent, “sectors” within the ecosystem, having definable ecologi-
shallow water such that all areas are within the photo zone cal characteristics or spatial boundaries, are managed
such that photosynthetic organisms like diatoms, green under different management plans specific for each sys-
and blue green algae, and aquatic moss are able to persist tem. Sectoral management is also used in areas subject to
in the water column or on the substrate, making riffle areas multi-agency or international management, in which each
important zones of primary production. agency or nation state manages the “sector” corresponding
risk analysis (pathway analysis) A management approach to its administrative or legislative authority or within the
in which managers develop different responses unique to spatial boundaries of its jurisdiction.
particular decision pathways and their associated levels of sex ratio The ratio of males to females in a population.
risk. simulation model A model that provides mathematical esti-
risk assessment A systematic process of evaluating poten- mation of the state of a system through time and, in
tial risks involved in a proposed activity or undertaking. In landscape and ecosystem management contexts, across
conservation biology, methods and techniques of risk space.
assessment are frequently employed in population viabil- sink In an ecological model, an area into which a stock or
ity analysis (PVA) and other kinds of population resource is absorbed. In habitat assessment, an area of
modeling, particularly for endangered and threatened low-quality habitat in which a population cannot replace
species. itself without immigration.
rival goods Goods whose consumption or use by one person soft law In the context of international law, nonbinding
reduces the amount available for everyone else. agreements that, although having no official means of
586 Glossary
enforcement, eventually come to define norms and species list A list of species present in a community.
standards for international behavior among nations with species richness The number of species present in a com-
regards to the topic or objectives addressed by the munity standardized to reflect the number of species
agreement. recorded per sampling area or observation effort.
sociocultural values – See also value; ecosystem Society for Conservation Biology (SCB) The premier
services Those values that measure the importance of international nonprofit professional organization of
ecosystems for human wellbeing; specifically, the mate- professionals, students, and non-profit employees engaged
rial, moral, spiritual, aesthetic motivations people have in the science, study and practice of conservation biology
regarding the environment and its conservation. and dedicated to advancing the science and practice of
source In an ecological model, an area from which a stock conserving biodiversity.
or resource originates. In habitat assessment, an area of soft law In the context of international law, non-binding
high-quality habitat in which a population produces sur- agreements that, although having no official means of
plus individuals which provide potential colonists for new enforcement, eventually come to define the norms and
areas or habitat patches. standards for international behavior.
spatially balanced sampling design (SBSD) – See also stakeholders Individuals or groups possessing vested
generalized random tessellation stratified (GRTS) interests in the persistence, health, products, state or
samples A sampling design in which sampling units are services of a system to whom managers have legitimate
more or less evenly dispersed over the area or resource and defined functional, ethical, legal or contractual
being sampled, ideally with equal numbers of sampling obligations.
units in each part of the area. SBPDs are intended to more stasis rate – See also transition matrix The diagonal
efficiently cover the sampling area and be more represen- elements in a matrix representation of present conditions
tative of the resource’s distribution than simple random (rows) and future conditions (columns) representing the
sampling. proportion of individuals of life stage (for populations) or
spatially explicit model A metapopulation or landscape habitat type (for landscapes) that do not change from the
model that incorporates differing degrees of connected- present time (t) to a specified future time (e.g., t +1).
ness between population or landscape subunits and stated preference methods – See also revealed preference
features localized interactions between subunits or methods A suite of tools within microeconomics to
individuals whose rules are defined by the spatial measure what individuals are willing to pay or willing to
relationships between interacting agents. accept for the gain or loss of a good or service in a
spatially implicit model A metapopulation model (e.g., hypothetical market.
Levins’ model) or landscape model in which local stochastic factors Factors whose effects on a population
populations and habitat patches are discrete, but equally vary randomly, but usually within a limited range of
connected with one another. values.
spatial planning – See also sectoral management In stochastic variation In population demography, random or
ecosystem-based management (EBM), management chance events that determine population growth and struc-
planning based on and specific to spatially-defined areas ture without relationship to the population’s density.
within the ecosystem. stock A standing crop or source from which a resource
spatially realistic model A metapopulation or landscape originates or is produced by ecological processes.
model that incorporates the specific geometry (e.g., size, stock-flow resources – See also fund-service resources
shape, and arrangement) of particular habitat Resources, usually produced from a standing crop or
patches within the landscape. “stock,” that arise through a transformation process, usu-
spatial scale A measure of habitat patchiness that relates ally self-renewing, that occurs within the ecosystem itself.
interpatch distance to a species’ dispersal ability. Such resources can be used at any rate desired. Forage for
species In the now most currently used phylogenetic species herbivores produced by a grassland is an example of a
concept, a group of organisms with a unique set of defin- stock-flow resource.
able and measurable genetic similarities. Among sexually stratified diffusion model A type of species invasion model
reproducing organisms, the older biological species con- that incorporates long- distance dispersal and density
cept may be used to define a species as a group of dependent rates of spread.
interbreeding populations reproductively isolated from stream order A measure of the magnitude of a stream based
other such groups. on the magnitude of tributary streams that have formed
species evenness The equitability of relative abundance or it. At its origin, a stream is referred to as a first order
density of different species in the area or community stream. When that stream joins another first order stream,
Glossary 587
the new stream becomes a second order stream. When two target indicators – See also performance indicators. In
second order streams join, they form a third order stream. production-oriented ecosystem management, variables
Whenever two streams of the same order join, they form a that measure outputs of what the system is intended to
stream of the next highest order. Increasing stream order produce, such as a specified level of stock or biomass of a
reflects an increasing area of associated watershed, the particular resource.
area of landscape drained by the stream. target population size In captive breeding management, the
subsidies See governmental subsidies population size at which managers attempt to maintain a
succession model Models of organism distribution that captive population.
assume that such distributions are determined by changes temporal scale In the context of habitat conservation, the
in vegetation or other environmental qualities which duration of a habitat’s persistence relative to a species’
change over time. generation time. More generally, the units and anticipated
sunshine approach An approach to international treaty span of time appropriate to an object of investigation.
enforcement which focuses on mechanisms to bring the timing limitations A method of mitigation limiting human
behavior of key parties (usually nation states) into the activities in habitats which are seasonally or periodically
open for public scrutiny, including mechanisms such as occupied by wildlife to those seasons or times when
regular reporting, peer scrutiny, on site monitoring and wildlife are absent.
media access and coverage. toll goods See club goods
surrogate species – See also indicator species A species tradable permits A market-based mechanism where
whose status is assumed to reflect the status of other regulators identify a maximum acceptable level of pollu-
species with which it shares the community. tion or depletion that is ecologically sustainable and dis-
sustainable development – See also ecological tribute permits under this level across polluting industries;
economics A paradigm originating in the 1970s focused however, if a polluter does not “use up” all of his pollution
on ‘development without economic growth’ emphasizing rights (because of increased efficiency and cleaner pro-
instead qualitative improvements. The United Nations has duction), he can sell his permit to another polluter, offer-
since emphasized that the practice of sustainable developing a cost-effective way to achieve overall reductions.
ment should harmonize three core elements: economic transition matrix In population viability analysis or habitat
growth, social inclusion and environmental protection. successional analysis, an arrangement of rows and
sustained yield – See also maximum sustainable columns that display the probability of an individual at
yield The amount of an exploited population that can be one life stage or habitat type in the present (column
harvested in a present time increment leaving the headings) changing or transitioning into a new life stage
population’s productivity undiminished in future time or habitat type in a subsequent time period (row headings).
increments. travel cost method (TCM) – See also revealed preference
sustained yield forestry A practice of forest management methods A method of determining the values of environ-
that permits the same amount of timber to be removed mental amenities, goods and services based on the
from the forest annually without loss or interruption of assumption that the more valuable an environmental ame-
forest productivity. nity, good or service, the farther people are willing to
systematic conservation planning An approach to conser- travel to get it, the more they will spend per trip, and the
vation planning that employs algorithms for identifying more trips they will make.
and ranking conservation priorities in reserve design by trend analysis A population monitoring tool for calculating
integrating the relative threat of loss of an area or sys- one or more specific demographic variables in a popula-
tem (vulerability) with its irreplaceability (ecological tion and, based on the value of the variable, then deter-
uniqueness). mining whether the population is growing, stable or
declining.
trophogenic zone In aquatic environments such as lakes, a
T region of high light intensity (shallower water) dominated
by the process of photosynthesis.
taking In the context of traditional property law, hunting, tropholytic zone In aquatic environments such as lakes, a
fishing, collecting, or trapping a creature to kill it or bring region below the trophogenic zone of lower light intensity
it into personal possession. Under the US Endangered in which the dominant process is decomposition.
Species Act, the traditional concept is expanded to include turtle excluder device (TED) A grid trapdoor installed
any intentional or unintentional act that harms or harasses inside a shrimp trawling net that keeps shrimp in the net
a protected species in any way. but directs other, larger animals, such as sea turtles, out.
588 Glossary
typological species concept The idea that species are dis- whole catchment management (WCM) A strategy of man-
tinguished by morphological characteristics that can be agement that perceives freshwater aquatic ecosystems,
determined by gross observation. particularly rivers and streams, as products of terrestrial
environmental constraints or “filters” which control inputs
of abiotic and biotic inputs to the stream ecosystem. WCM
U thus emphasizes management of land use practices within
the entire watershed (catchment) as the foundational basis
umbrella species A species, or group of species (e.g., large for management of freshwater systems within the
mammals), of particular conservation or public interest, watershed.
and whose protection is assumed to provide benefit or a wilderness ideal The view that large landscape areas rela-
protective “umbrella” to many other species in other taxo- tively unaffected by human activity or residence represent
nomic categories. ideal ecosystems that should be given priority in nature
use value The value derived from the actual use of a conservation and preservation in order to better study their
resource. ecological processes and conserve their species
utility In neoclassical economics, the want-satisfying components.
“power” of any commodity or the capacity of a commod- Willingness to Accept Compensation (WTA) A form of
ity to give satisfaction or happiness. contingent valuation (CV) analysis in which the market
value of a non-market good, such as a non-human species,
is estimated by what a person would be willing to accept
V as compensation for its loss.
Willingness to Pay (WTP) A form of contingent valuation
value The basis for an estimation of worth or importance. (CV) analysis in which the market value of a non-market
variegated habitats and landscapes Environments where good, such as a non-human species, is estimated by what a
human alteration has reduced the amount of natural habitat person would be willing to pay to prevent its loss or harm.
and introduced a greater degree of landscape heterogene-
ity and variety of vegetation.
visual minimization A method of environmental mitigation Y
that involves reducing the distance at which animals can
see objects associated with human disturbance. Yellowstone Model A pattern of establishing national parks
based on the characteristics of Yellowstone National Park,
USA, emphasizing the conservation of regional-scale
W landscape areas characterized by relatively little human
impact and residence, or the removal of resident humans
western privilege Privilege is a special, unearned advantage and their activities and impacts from such areas if neces-
or entitlement, used to one’s own benefit or to the detriment sary to fulfill conservation objectives.
of others. In the context of moving certain conservation
models from western Europe and North America to other
parts of the globe, this privilege was often exercised as Z
a disregard or deemphasisis of local, place-specific history
and culture as well as place-specific social, economic, and zoning – See also conservation easement An arrangement
political traditions and customs, and subsequent replace- in which some of the property rights normally associated
ment of these with western norms and practices. with the individual owner of a property are transferred to
wetland An area or ecosystem in which the water table is at or held by the community in which the property is located,
or near the surface or the land is covered by shallow water effectively restricting what individual property owners
during at least part of the year, vegetation is dominated by can do on their property because of the context in which
hydrophytic (literally, “water-loving”) plants, or the sub- the property is located. Conservation easements are a
strate is characterized by hydric (wet or waterlogged) soil special case of zoning applied toward conservation
types. objectives.
Index
A Akhirah (accountability), and Islam, 431

Aammiq Wetland (Lebanon), 437 Ǻland Islands, 187
Abandoned habitats, 104–106, 111 Alaska (US), 4, 88, 147, 148, 204, 349, 495
A-B cleavage, 27, 28 Aleutian Islands, 204
Absolute family richness, 69 Allee effects, 103
Abundance, and species richness, 324 Allee, W.C., 84
Academic credentials, and conservation education, 539 Allele
Acidification, of freshwater habitat aspen, 311 recessive lethal, 176–177
Action research, and conservation education, 545 Allele frequencies, 172
Adaptation to captivity Alliance for Zero Extinction (AZE), 106
minimization of, 196, 201 Alliance of Religions and Conservation (ARC), 440–442
Adaptive capacity, to climate change, 129, 149–151 Allozyme electrophoresis, 180
Adaptive management Alpha diversity, 48–52, 74
role in ecosystem management, 370 Alpha rarity, 51
small and declining populations, 256, 257 (see also Management) Alteration of flow regimes, 312
Adaptive science, conservation biology as, 29 Alternative stable states
Adirondack Mountains (USA), 311 lakes, 318, 319
Adventitious root systems in wetland plants, 323 novel ecosystems, 94
Advisory groups and National Environmental Policy Act, 499 Altitude and species diversity, 57, 139
Advocacy Altman, I., 365–367
alternative views of, 563–566 Aluminum and acidification of aquatic ecosystems, 311
conflicts of interest in, 567–569 Amazon Basin, 23
by conservation biologists, 490, 563, 565 Amazonia, 23, 25, 63
focused advocates, 565 Amenity values, of biodiversity, 458
inadvertent, 566–567 American alligator (Alligator mississippiensis), 323
intentional, 566–567 American Association for the Advancement of Science (AAAS), 40, 41,
professional, 164, 563–565 71, 72, 84, 186, 429
role of, in conservation leadership, 566 American chestnut (Castanea dentata), 270
Aerial photographs, 280, 339, 340, 386 American Museum of Natural History
Aesthetics and aesthetic value, 367, 413 Center for Biodiversity, 560
Africa American Ornithologists Union, 9, 61
coral reefs and ecosystem management, 371 American robin (Turdus migratorius), 136, 227
grazing mammals, 38, 395 Amino acid sequences, 181
habitat suitability model of reserves, 291–292 Amphibians, and biodiversity, 325
nonindigenous species in Lake Victoria, 375 Amphibian Research and Modeling Initiative (ARMI), 229
wildlife harvests and local communities, 480 (see also South Africa) Amsinckia grandiflora, 234–236
African elephant ivory, and CITES, 515 Analysis of the likelihood of extinction (ALEX), 187
African Muslim Environmental Network (AMEN), 440 Analytical-deterministic population viability analysis, 220
Agapow, P.M., 47, 48 Analytical disciplinary skills, and conservation education, 552
Agenda 21 (United Nations, Rio Summit), 479, 516, 517, 519 Andasibe-Mantadia National Park (Madagascar), 461
Age structure, of population, 103, 214, 215, 220, 242 Andrewartha, H.G., 217, 218, 220
Agostinho, A.A., 313, 315, 316 Angermeier, P.L., 412
Agriculture Animal rights, 423, 425, 433
eutrophication, and aquatic, 311, 317 Anomalies, and climate change, 158, 360
freshwater habitats, 317, 328 Antarctic Peninsular Plume (APP), 346
grazers and grazing, 394 See also Southwest Atlantic Section (of Southern Ocean)
pastoral, 4, 38, 295, 461 Anthropocene Epoch, 81–83, 95
pesticides, 396, 465 Anthropocene Working Group (AWG), 81, 82
polyculture cropping, 38 Anthropocentric values, 421
slash, and-burn, 197 Anthropogenic biomes, 83–86, 88–90, 93–95, 106
systems, 37, 38, 42, 117 (see also Fertilizer) Anthropogenic habitat, 104–106

F. Van Dyke, R. L. Lamb, Conservation Biology, https://doi.org/10.1007/978-3-030-39534-6
590 Index
Antwerp (Belgium) Zoo, 198 Beever, E.A., 29, 32

Appalachian Mountains, USA, 270 Beiji (Lipotes vexillifer), 348
Appendices of, CITES, 514 See also Yangtze dolphin
Apps, M.J., 130 Bennett, N.J., 549–551
Aquaculture, 26, 316, 336, 339 Bennetts, R.E., 26, 382, 384, 385
Aquatic ecosystem management Benthic communities
biomanipulation, 317, 318 numerical complexity score, 332
chemical manipulation, 317, 318 Bequest value, 419
dredging, 317 Beta diversity, 49–52, 327
input regulation, 317 Beta rarity, 51, 253
landscape filter model, 319 Bet hedging and fisheries management, 353
Aquatic macrophytes and role in lake transition states, 319 Bevers, M., 284, 285
Aquatic systems Bhuma devi and Hinduism, 431
introduced species, 310, 330 Bhumi Project and Hinduism, 440
Aquatic zoogeographic units (AZUs), 75, 320, 321 Big bluestem (Andropogon gerardii), 391
Arcadia, 7 Big Cypress National Preserve, 377
Arcadian ideal/Arcadian vision, 7, 12 Bighorn sheep (Ovis canadensis canadensis), 142–144, 204
Archived data, and ecosystem management, 382, 386 Bioclimate envelope, 151–154, 158
Arctic bearded seal (Erignathus barbatus), 502 Biodiversity
Ardoin, N.M., 549 aquatic habitats, 325, 326
Aristotle, 45 climate-wise strategy, 73, 74
Arizona (USA), 96, 107, 239, 289, 290 definition and concepts of, 40–42, 44, 76
A Rocha International (ARI) eco-functionally wise (EcoWise) strategy, 73, 74
A Rocha Kenya, 437 ecosystem-based approach to conservation of, 68, 75, 161, 385, 456
A Rocha UK, 437 effects on
Arrhenius, S., 129 below ground biomass, 37
A Sand County Almanac, 16, 17, 427 infectious disease, 38
Ashe juniper (Juniperus asheii), 190 plant communities, 36, 37
Asian carp (Hypophthalmichthys nobilis), 182 system stability, 36
Asian clam (Corbula amurensis), 100 as focus of conservation biology, 22, 24, 26, 29, 30, 32, 38–40,
Asian open-billed stork (Anastomus oscitans), 433 61–66, 68–73, 106–117, 125–166, 171, 206, 262, 292, 326, 327,
Aslan, C.E., 349, 542, 543 353, 412, 437, 438, 455, 465, 469, 471, 482, 484, 533, 543, 560,
Aspen 563
Populus spp., 88 geographic-based approaches to conservation of, 50–52, 57, 76
quaking (Populus tremuloides), 263, 394 hotspots of endangered species and, 57, 58, 62, 63, 66, 69, 71, 72,
Assisted migration, 161 106, 110, 153, 158, 162, 351
Association for the Protection of Sea Birds (APSB), 8, 9 indices, 48, 69, 74
Association of Zoos and Aquariums, 197, 202 and invasive species, 95, 97, 98, 104, 105, 125, 141
Atlantic right whale (Eubalaena glacialis), 348 inverse relationship to latitude, 57
Audubon Society, 254, 471 measurement, 48–51
Australia National Forum on, 44
aquatic systems and invasive species, 317 rare species, contributions to, 36, 68, 189, 326, 424
corridors and small mammals, 298 socioeconomically wise (SocioWise) strategy, 73, 74
endemism of mammals, 54 thought styles, 45
Great Barrier Reef Marine Park (GBRMP), 350–352, 402, 561 in urban areas, 62, 64, 106–117, 121
Auto industry, 463 Biodiversity Benefits Index, 32, 420
Avoidance of inbreeding strategies, 201–202 Biodiversity coldspots, 62
See also Maximum avoidance-of-inbreeding (MAI) strategies Biodiversity in Development Project, 30
Biogeochemical cycles, 86, 130, 131
Biogeographic approach, to minimum viable population experiments,
B 136, 327
Babbitt, B., 506 Biological control, of non-indigenous species, 221
Back to the land movement, 8 Biological productivity and ecosystem management, 467
Baker, C.S., 205, 206, 251 Biological species concept (BSC), 46
Bald eagle (Haliaeetus leucocephalus), 138, 420, 500 Biomanipulation, of freshwater ecosystems, 318
Ballast-water discharges, 98, 182, 313 Biome boundaries, 134
See also Invasive species Biomes, 44, 69, 71, 83–86, 90, 93–95, 106, 155, 275, 280
Baltimore, Maryland (USA), 107, 108 Biophilia Hypothesis, 422
Barcelona Convention for the Protection of the Mediterranean Sea Biosafety Protocol, 18, 240, 518
Against Pollution, 513 Biosphere reserves
Barcode of Life Data (BOLD) System, 182 Spain, 59
Barry, D., 412, 490 Biotemperature, 145, 146
Basic rule of conservation genetics, 172 Biotic assemblages, 97
Bean, M.J., 506 Biotic community of Möbius, 84
Beaver (Castor canadensis), 97, 323, 324, 393 Biotic resources and instrumental values, 291, 353, 418–420
Index 591
Birch, L.C., 217, 218, 220 Bycatch regulation

BirdLife International, 62, 361 behavior effect, 349, 542
Birds and biodiversity, 40, 50, 55, 57, 61, 62, 65, 66, 71, 98, 104, 107, deployment effect, 349
136, 325, 471
Biscayne National park, 377
Bison (Bison bison), 61, 173–174, 178, 218, 219, 378 C
Black-capped chickadee (Parus atricapillus), 225, 227 Caldwell, L.K., 362, 363, 493, 495
Black-footed ferret (Mustela nigripes) California
captive breeding program, 36, 236, 424, 542 grassland restoration, 44, 142, 234
restoration, 235 Joshua Tree National Monument, 27
Black-footed Ferret Recovery Team (BFRT), 237 Mineral King Valley, 425, 426
Black heron (Egretta ardesiaca), 434 nonindigenous invasive species, 100
Black-legged kittiwake (Rissa tridactyla), 374, 375 San Bruno Mountain controversy, spotted owl and ecosystem
Black rhino (Diceros bicornis), 61, 514 management, 507
Black stilt (Himantopus novaezelandiae), 313, 392 Santa Clara red ribbons (Clarkia concinna concinna), 119 (see also
Blickley, J., 551, 552, 555 Los Angeles, and plastic trees biodiversity and indicator species)
Blue jays (Cyanocitta cristata), 227, 228, 277 California condor (Gymnogyps californianus), 424
Blue whale (Balaenoptera musculus), 346, 347 Callicot, J.B., 12, 15, 418, 421, 427–429, 431
Bohlen, E.U. Curtis, 500 Calvert Cliffs vs. Atomic Energy Commission, 495
Boitani, L., 292, 293 Canada
Bonelli’s eagle (Aquila fasciata), 247, 249, 250 British Columbia (Canada), 183, 187, 204, 298, 299, 549
Bornean orangutan (Pongo pygmaeus), 442–444 climate change and national parks, 125, 126
Borneo Orangutan Survival (BOS) Foundation, 443 federal policy on wetland conservation, 465, 563
Bottlenecks and small populations, 172–176, 178, 179 Prince Albert National Park, 126
Boulder, Colorado (USA), 93 Canadian Environmental Assessment Act, 499
Bowdish, S., 535 Candidate species, and Endangered Species Act, 505
Boyce, M.S., 213, 313 Canopy management, 275
Boyle, T., 206 Cape seahorse (Hippocampus capensis), 335
Bramhaputra-Salween Landscape, 64 Captive breeding
Brandis, D., 15, 21 genetic management, 195, 200, 201, 237
Brazil, 6, 22–24, 31, 58, 59, 63, 66, 313–315, 483, 515, 524 mean generation time, 201
Atlantic Forest Region, 66 (see also Amazonia) okapi (Okapia johnstoni), as example of, 197–199 (see also Pedigree
Brewer, C., 552, 565 analysis)
Brewer’s sparrow (Spizella breweri), 415, 416 strategies of, 197
Bridger, J., 10 target population size, 201
British Royal Society for the Protection of Birds (BRSPB), 9 Carbon
Brown bear (Ursus arctos), 141 sequestration, 114, 133, 165, 457, 483
Brown, C., 400–402, 542 sink, 131–133, 164, 165
Brown-headed cowbird (Molothrus ater), 228, 253, 424 source, 131
Brown, M., 351, 353 Carbon 12 (12C), 82
Brown tree snake (Boiga irregularis), 98, 100, 101 Carbon 14 (14C), 82
Browsers and browsing, 264, 375, 376, 382, 391, 393–395 Carbon removal capacities, 132
Brundtland Report, 479 Careers, in conservation biology, 531, 548, 550, 557
Brussard, P.F., 1, 563–565 Caribbean basin
Buddhism, 432–434, 440–441 species-area relationships, 324
Buddhist Peace Fellowship, 441 Carlsen, T., 234, 236
Buddhist Perception of Nature Project, 440 Carp (Cyprinus carpio), 182
Buddishm, Law of Dependent Origination, 432 Carpenter, S.R., 318, 319
Buffalo (Bubalus bubalis), 62 Carruthers, D., 478, 479
Buffering and acidification, 311 Carrying capacity (K) 3
Buffers assumptions, 218, 219
corridors, reserve design, and landscape planning, 160, 161, 285 Cartagena Protocol, 518
riparian vegetation, 398 Catch variation (in fisheries management), 354
strips and wetland restoration, 396, 563 Categorical exclusion (CE), and environmental impact
wetlands as, 311 assessment, 496, 497
Bunn, S.E., 313, 328–331 Cattle and ecosystem management, 173
Bureaucratic organizational model, 361 Caughley, G., 175, 176, 185, 563, 564
Bureau of Land Management (BLM), 239, 362, 404, 496, 534 Center for biological diversity (CBD), 66, 503, 504, 511, 518, 525
Büscher, B., 119, 120 Centre for orangutan protection (COP), 443
Butler, J., 396, 398, 399 Centers of plant diversity (CPD), 71, 72
Butler, K., 359 Cereal leaf beetle (Oulema melanopus), 103
Bycatch Céréghino, R., 327
cryptic, 347 Chapin, F.S., 88, 90
definition, 347, 348 Charismatic megafauna, 420, 509
ghost, 348 Charlesworth, B., 217, 219
mitigation of, 349 Charter of the United Nations, 510
mortality, 347–349 Cheatgrass (Bromus tectorum), 96, 142
592 Index
Chen, J., 274, 275 Commitment

Chestnut blight fungus (Cryphonectria parasitica), 270 effective, 114, 115
Chicago, Illinois (USA), 112 normative, 29, 114, 115
See also DuPage County, Illinois (USA) overall, 114, 115
China, 5, 25, 26, 40, 64, 190, 265, 297, 310, 348 Commodity production and ecosystem management, 27, 408
China Valley, British Columbia (Canada), 298 Common European adder (Vipera berus), 190
Chinook salmon (Oncorhynchus tshawytscha), 4 Commission on Sustainable Development (CSD), 516, 518
Chipko movement, and Hinduism, 440 Commission on Sustainable Development, and Agenda 21, 516
Choice modeling/choice experiments, 461 Common pool resource, 453, 467–469
Chum salmon (Oncorhynchus keta), 138, 139 Commons Preservation Society, 8
Cicero Swamp Wildlife Management Area, New York (USA), 271 Community
CITES, see Convention on the International Trade in Endangered alpha diversity, 48, 49, 51, 52
Species (CITES) beta diversity, 49–52
Citizen jury, 499 gamma diversity, 51, 52
Citizen of a professional culture, 534 Community-based ecotourism (CBET), 481, 482
Cladistic species concept, 47 Community-based environmental protection, 522
Clark’s nutcracker (Nucifraga columbiana), 141 Community-based natural resource management (CNRMs), 62, 479
Clark, T.W., 10, 412, 541 See also Integrated conservation and development projects (ICDPs)
Classical metapopulation, 222, 223 Comoros Islands, 351
Clean Air Act, 463 Compliance information systems, and global cooperation with
Clean Power Plan, 456 conservation laws and treaties, 519
Clean Water Act, 75, 463 Conata Basin, South Dakota (USA), 237, 239
Clements, F., 84 Conceptual model, 89, 91, 107, 272, 297, 332, 333, 382, 384, 389, 407,
Climate change and extinction of local populations, 29, 117, 144 519
Climate change, forms of Conflict of interest and conservation advocacy, 566, 567
discontinuities, 127 Congressional record, 363
increasing variability, 127 Connectedness and connectivity
jumps, 127 defined, 282
trends, 127 distinguished, 93, 320
Climate change, global fingerprint of, 131, 134–149 habitat fragmentation, 273
Climate change, human-induced, 131 metapopulation theory, 220, 223
Climate change-integrated percolation cluster and threshold, 283
conservation strategies, 73, 126, 127, 149, 151, 159–164 Connectivity algorithm, 290
recommendations for, conservation actions, 159 Conservation Alliance, 406
vulnerability assessments, 150, 151, 164 Conservation basic income (CBI), 120, 121
Climate change metrics, 155, 157 Conservation Biology (journal), 1, 2, 490, 537, 563
Climate niches, 126, 152, 161, 163 Conservation Biology Institute, 2
Climate refugia, 153, 160, 163 Conservation easements, 255, 469–471, 563
Climate variations Conservation education
regular periodic variation, 127 creative thinking, 542, 543
Climate velocity, 155, 158, 161 foundational elements, 532–533
Clinton administration, 507, 514, 523 international relations, 542
Closed-ended questions, and surveys, 419 political science, 542, 549
Cloud forests, Central American, 36, 145 social and cognitive psychology, 542
Club goods/toll goods, 453 sociology, 542, 549
CO2 Conservation entrepreneurs
atmospheric concentrations, 130, 133 existing opportunities, 560, 561, 563
correlation with global temperatures, 130, 131 potential opportunities, 560, 561
Coalition on the Environment and Jewish Life (COEJL), 437 traction opportunities, 560–562
Coarse woody debris (CWD) Conservation International, 113, 161, 550
forests, 298 Conservation law, 19, 42, 60, 61, 75, 76, 206, 302, 411, 416, 431, 444,
Coarse-filter approach, 160–163 492–521, 525, 526, 561, 568
Coarse-filter, climate informed conservation strategies, 159, 161, 162 common characteristics of effective, 493
Cody, M.L., 51, 52 compliance and effectiveness factors, 519, 520
Coercive measures, and international treaties on conservation, 521 creation and enforcement, 492, 518–521
Cognitivist view, of conservation ethics, 412 ecosystem management, 93, 499, 574
Coho salmon (Oncorhynchus kisutch), 138 genetics, role of, 190, 206
Collared lizard (Crotaphytus collaris), 278 global cooperation, 17, 164
Colobus monkey, Black-and-white (Colobus vellerosus), 434 interdependence and global, 521–527
Colorado chipmunk (Tamias quadrivattus), 186 marine resources, 351, 353
Colter, J., 10 origins of, 3–11
Colter’s Hell, 10 public trust, doctrine of, 412
See also Yellowstone National Park relationship to conservation biology, 2, 28–30 (see also Federal
Columbia, 64, 65, 69 government; Litigation; Policy, environmental)
Columbia University, 560 Conservation leadership, and conservation vocation, 566
Co-management, 120, 351, 353, 369 Conservation Letters (journal), 196, 537
Index 593
Conservation management agreement (CMA), 254–257 Critical habitat and Endangered Species Act, 36, 240, 289, 302, 313,
Conservation mission, 533 325, 345, 375, 378, 484, 500, 505–506, 509
Conservation positivism, 564 Currents, global marine, 133, 328, 345
Conservation reliant species, 42, 252–257 Cyprinid fishes (Family Cyprinidae, carps and minnows), 311
Conservation reserves, 10, 22, 24, 70, 287, 291, 292, 321, 350, 560 Cytochrome oxidase I site gene, 203
Conservation science, xiii, 26, 28, 30–32, 117, 119, 229, 273, 411–413, Czech Republic, 536
489–491, 526, 533, 537, 540, 546, 550, 554, 556, 558–560
Conservation Science and Practice (journal), 537
Conservation social sciences, 548–551 D
Conservation triage, 449 Daehler, C., 104–106
Constructed wetlands, 62 Daily, G.C., 450
Consumer surplus and environmental economics, 459 Dalai Lama of Tibet, 440
Consumer surplus and microeconomics, 459 Daly, H.E., 32, 412, 453, 465, 474, 475
Contingent valuation analysis (CVA), 460–461, 477 Darwin, C., 45, 51
Convention between the United States of America and Great Britain for Data Basin, 2
the Protection of Migratory Birds, 18 See also USGS
Convention Concerning the Protection of the World Cultural and Data Basin Climate Center, 2
Natural Heritage, 521 Data mining (mining of legacy data), 387
Convention on Biological Diversity (CBD), 38, 66, 206, 504, 511, 518, Davis, B., 47
520, 524, 525 Deadweight social losses, 463
Convention on Climate Change, 164, 165, 510, 517 Debinski, D.M., 280
Convention on the International Trade in Endangered Species, 206 Debt bondage system, 23
Convivial conservation, 119–121 Decision analysis and management of small populations
Coral bleaching, 145, 340 adaptive management, 29, 94, 106, 115, 233, 252, 256, 370, 371
Coral reefs classification problems, 104
biodiversity of, 145 selection problems, 178, 188, 264
biological restoration, 343, 350 Declaration on the Human Environment, 512
blast fishing, 340, 341, 344 Declaration on the Rights of Indigenous Peoples, 524
bleaching, 145, 340, 342 Declining population paradigm, 28, 360
conservation status, decline and endangerment, 330, 335 Deep ecology, 425–427, 432
micro-colony fusion, 341 Deer mouse (Peromyscus maniculatus), 277
structural restoration, 341–343 Defenders of Wildlife, 461, 503
symbiotic relationship with, 145 Demersal species, 343, 344
transplants, 340, 341, 343 See also Groundfish
Core habitat, 109, 160, 273, 279, 470 Demes, 172, 183
Corncrake (Crex crex), 266, 267 Democratic Republic of Congo (DRC), 197, 198
Correlative approach, to modeling, 151–154 Demographic monitoring, 233, 234
Corridors See also Factor resolution; Trend analysis
connectivity and metapopulations, 75, 153, 160, 161, 290 Demographic stochasticity, 214, 215, 240, 246, 256, 345
design of nature reserves, 289 Demonstration-oriented research and conservation education, 114, 424,
gene flow, 75 438, 440, 532, 545, 546, 549, 558
movement enhancement between habitat fragments, 280 Density dependence, population regulation by, 103, 144, 213, 217–220
studies of, 280, 340 Dependent Co-Arising, Law of, 432
Costa Rica Design, see Nature reserves
cloud forests, 36, 41, 145, 146 Designated critical habitat, 500
coffee farm and ecosystem services, 457 Designed habitat, 104, 105
Forestry Law and ecosystem services, 483 DeSoto National Wildlife Refuge (Iowa), 27, 49, 535
National Forestry Financial Fund, 483 Detection
travel cost method (TCM), 458 computer programs for calculation of, 182, 227
Costa Rican National Park Service, 459 methods of, 225
Costanza, R., 335, 450, 473 probability, 212, 227, 229–231
Cost-benefit analysis (CBA)/benefit-cost analysis, 413, 420–421, 456, problem of, 224–229
463 threshold distance, 225, 228
Cottonwood (Populus spp.), 88, 317 variability, 230
Courter, J., 543, 546, 547 Deterministic factors, 219, 241, 246, 270
Covey, S.M.R., 405, 532 Deterministic models, 241, 243, 246, 270
Coyote (Canis latrans), 193–196, 237, 239 Dhammanaat Foundation, 441
Crane Diadromous fish species, 316
blue (Anthropoides paradiseus), 62 Dichlorodiphenyltrichloroethane (DDT), 335
red-crowned (Grus japonensis), 25, 26 Diet selection, 219
Siberian (Grus leucogeranus), 62 Dietz, J.M., 540, 547, 548, 557
whooping (Grus americana), 216, 217 Differential fitness species concept (DFSC), 46
Crisis Ecoregion (CE) strategy, 71, 72, 162 Diffusion coefficient, 102, 103
Crisis-oriented discipline, conservation biology as, 29 Diffusion theory, 103
Critias, 4 DiMento, J.F., 512, 513
594 Index
Dinesen, I., 424 Ecological stressor, 125, 126

Dispersal and dispersal behavior Ecological succession, 50, 270, 271
aquatic invertebrates, 311 Ecological uniqueness, as criteria for protected area selection, 70, 289
climate change, 54, 142, 151, 154, 155, 158 Ecological value
corridors, role in, 290 Ehrlich Identity, 478
gene flow, 172, 189, 203, 212, 350 instrumental value, 418–421
invasive species, 104–106, 111, 142 Kuznets Curves, 478
landscape spatial structure, 269 market-based solutions to conservation problems, 465–466
limitations of, 154 property values and stream restoration, 457
marine reserves, 349–355 steady-state, 474
population demography, 212–214, 241, 288 sustainable development, 466, 477
spotted owl, 288–290, 363 valuation methods, 418, 419, 453, 456, 475–477
Dispersal models, 282 Ecologistic values, 417
Distance effect, 274 Ecology, as foundational discipline for conservation biology, 11–16
Distance sampling, 225 The Ecology of Invasions by Plants and Animals, 96
Di Stefano, J., 375, 376 Economic, 443
Distinct population segments (DPS), 504 Economic behavior, 449, 450, 484, 506
The Distribution and Abundance of Animals, 217, 220 See also Economic institutions
Disturbance Economic growth
ecosystem, 329, 382, 384, 388, 477 assumptions, 476
role of, in modeling, 154 capital and resources, 449, 473, 474
Diversity and inclusiveness in conservation biology, 1, 28, 35, 47, challenges to, 449, 479
51–57, 109, 412, 557, 559, 563 redefinition of, 474 (see also Ecological economics)
DNA Economic institutions, 107, 461–463
barcodes and barcoding, 181, 203 Economics, 30, 365, 418, 419, 421, 485, 516, 519, 549, 551, 552, 568
fingerprinting, 176, 180, 181 See also Cost-benefit analysis
Doane, G.C., 10, 16 Ecosystem
Dobson, A.P., 45 definition of, 89, 90, 359, 360, 377, 379, 450, 477
Dolphins and Marine Mammal Protection Act (MMPA), 522 effective size of, 378
Dominionistic value, 417 Ecosystem engineers, 97, 393–396
Douglas fir (Pseudotsuga menziesii), 269 Ecosystem management
Dourado (Salminus brasiliensis), 315 governance systems, 396–408
Draft EIS (DEIS), 497, 498 limit indicators, 373, 382
Dragonflies (Order Odonata), 53 marine reserves, 371
Dredging and freshwater ecosystems, 317, 318 multiple interests, 396, 398, 399
Drew, J.A., 560 National Environmental Policy Act (NEPA), 360, 493–500, 522,
Drivers-Pressures-State Change-Impact-Response (DPSIR) approach to 526
ecosystem-based management, 374, 389 performance-based management, 371
Drosophila, 172, 177, 184, 185 performance indicators, 371, 373, 375–377
Duchelle, A.E., 543–545 scientific basis of, 408
Dugong (Dugong dugon), 335, 348, 402 sectoral management, 379
DuPage County, Illinois (USA), 112 spatial management, 381
See also Chicago, Illinois (USA) target indicators, 373 (see also Management definition and goals of)
Dusky gopher frog (Lithobates sevosus), 505 Ecosystem modeling
Duvall, B., 426 connectors, 390
Dwyer, J.W., 363 flows, 389
Dynamic global vegetation models (DGVM), 154, 155 parameters/converters, 390
Process model, 389
sinks, 389
E sources, 389
Earth Charter Consultative Committee, 440 Sphere model, 389
Earth First, 427 stocks, 389
Earth Island Institute, 523 Ecosystem monitoring
Earth Summit, 516 Geographic Information Systems (GIS), use of, 75, 378, 379, 385
Eastern hellbender (Cryptobranchus alleganiensis alleganiensis), 182 lake and bog sediments, use of, 387
Eastern massasauga rattlesnake (Sistrurus catenatus catenatus), 271 long-term, 382
East Maui Watershed Partnership, 407 Parker transect, 382
Ebell, M., 506 probability design, 382
Ecocentrism, 17, 427–428 regularly collected data (continuously collected data), 382–384
Ecological drainage units (EDU), 75, 320–322 spatially balanced sampling design, 382
Ecological economics, characteristics of, 473–475 survey design, 382
Ecological knowledge, in indigenous societies, 434 Ecosystem processes
Ecological match and invasive species, 100, 101 fire, 391, 392
Ecological processes, and variations in biodiversity, 44, 62, 97 flooding, 392
Ecological Society of America (ESA), 36, 37, 203, 249, 253, 345, 362, herbivory, 393–395
439, 493, 526, 538 Ecosystem restoration, 92, 163, 165, 404, 454
Index 595
Ecosystem services Elk, C.B., 435

benefits of, 364, 450, 451 Ellis, E.C., 83–86, 88, 89, 95, 154
carbon sequestration, as a type of, 457, 483 Elton, C.S., 96, 98, 224, 428
cultural, 37, 364, 450, 477 Emerson, R.W., 12, 432
provisioning, 31, 37, 364, 450, 476, 483 Emigration, 186, 212, 213, 215, 287
regulating, 37, 364, 397, 399, 450, 477 See also Populations
soil conservation, as a type of, 454 Emlen, J.T., 225, 227
supporting, 37, 364 Emotivist view, of conservation ethics, 416
valuation of, 456 Empirical models and species response to climate change, 149
value of, 335, 450, 451, 475–477, 480 Endangered Species Act (ESA)
Ecosystem structure, 89, 96, 147, 384, 388, 392, 394, 454 application of, 203, 416
Ecotourism, 119, 481–482 assessment of value, 203
Ecotrust, 550 candidate species, 503
Edge habitat destruction and degradation, 500–503
expansion, 275, 276 history of, 500–501
sealing, 275, 276 hybrid policy, 193, 493
softening, 275, 276 property rights conflicts, 506
Edge effects, 273–275, 297, 378 recovery plans and species recovery planning, 27, 236, 249, 250,
See also Edge influences (EI) 252, 503, 509, 510
Edge environments, 273, 274 species listing and delisting, 501–503
Edge influences (EI) strengths of, 501, 509–510
compositional responses, 274, 276 taxonomy, 203
decay value, 274, 275 weaknesses of, 509–510
distance of edge influence (DEI), 276 wolf restoration, 195, 196
expansion, 275, 276 Endangered Species Committee, 501
primary, 275 Endangered Species Conservation Act, 500
process response, 275, 276 Endangered Species List, 20, 47, 253, 255
sealing, 275, 276 Endangered Species Preservation Act, 500
secondary, 275 Endemic Bird Areas (EBA), 71, 162
softening, 275, 276 Endemism
structural responses, 275 and conservation value, 59
Edge species, 111, 273, 274, 279, 280 Energy efficiency, 475
Education Energy–renewable/alternate sources, 164, 473
careers in conservation biology, 555 Enquist, C.A., 145
environmental, 114, 353, 433, 437, 441, 442, 544 (see also Entrance fees and travel cost method, 459
Universities) Entropy and ecological economics, 472, 473
Effective population size Environmental assessment (EA) and environmental impact
adaptation to captivity, 196, 197, 201 assessment, 496
ecological and evolutionary processes, 75 Environmental DNA (eDNA), 181, 182
GD, 188, 201 Environmental dumping, 478
gene flow, 75, 186, 189, 190, 204, 205 Environmental economics, 478
genetic diversity and bottlenecks, 175–177 Environmental impact statement (EIS), environmental impact
inbreeding effective size, 177 assessment, 420, 493–499
maintenance of, 75 Environmental justice, 492, 551
minimum, 173, 214, 233, 240, 292 Environmental law, based in
mtDNA, 181, 182, 204 equitable distribution of risks, 491, 492
population recovery, 175 ethical rights, 491
subdivision of populations, 189 utilitarian interests, 491, 492
variance effective size, 177, 201 Environmental policy
Ehrenfeld, D., 2, 3, 428 governmental regulation, 463
Ehrlich, A.E., 478 governmental subsidies, 463
Ehrlich identity, 478 governmental taxation, 463
Ehrlich, P., 478 tradeable permits, 465–466
EIS implementation plan (IP), 497 voluntary programs, 463
Electricity Company of São Paulo State, 316 Environmental property rights, 484, 492
Electrophoresis, 180 See also Tradable permits
Elephant Environmental Protection Act (New Zealand), 499
African (Loxodonta africana), 61, 394 Environmental Protection Agency (EPA), 131, 455, 456, 463
Asian (Elephas maximus), 204 Environmental stochasticity, 214, 215, 220, 240, 246, 256, 289
Elevational life zones, 145 Environmental Water Allocations (EWAs), 313
See also Holdridge Life Zones Epistasis, 172
Elevational shifts, 139 Epps, C.W., 142–144, 204
Elk (Cervus elaphus) Espoo Convention and environmental impact assessment, 500
effective population size, example of, 177, 178, 218 Ethical value, 411, 564
harem mating, example of, 177, 178 European Endangered Species Program (EEP), 197
Line Creek population, 298–302 European frog-bit (Hydrocharis morsus-ranae), 311
596 Index
European Studbook (ESB), 197 Fire ant (Solenopsis invicta), 102

European Union (EU) Fire return interval (FRI), 96, 104
Common Fishery Policy (CFP), 349 Fire, role in
Habitats Directive, 349 ecosystem management, 377, 378, 388, 391, 392
Member States, 349 Hinduism and conservation ethics, 431–432
Eutrophication and aquatic ecosystems, 311, 317, 318 history of conservation, 391, 392
Evangelical Environmental Network (EEN), 437 red-cockaded woodpecker habitat, 506
Evangelicals for Social Action (ESA), 437 sagebrush (Artemisia) habitat, 262, 388, 404, 415, 416
Everglades National Park, 11, 377 First International Conference on Conservation Biology, 1
Evolutionarily significant unit (ESU), 48 Fischbach, A.S., 147, 148
Evolutionary species concept, 46 Fish assemblages, shifting under climate change, 139, 140
Evolutionary time and conservation biology, 29 Fisher, B., 457, 485
Exclusive economic zones (EEZs), 524 Fishing industry
Existence value, 419, 460 fisheries management, 25, 26, 28, 343, 349, 353, 373, 401, 354, 371,
Experiential value, 421 400
Exponential population growth, 102, 213 marine bycatch, 349, 542
Exposure, climate change, 129 marine protected areas (MPAs), 353–355
External costs and externalities MMPA, 522
negative externalities, 455 whaling, 346
positive externalities, 456 (see also Cost-benefit analysis; Market Fixation index, 190
failure) Fixation of deleterious alleles, 171, 175
Extinction Flather, C.H., 284, 285, 289, 509
coefficient of, 185, 187 Fletcher, R., 119–121
crisis of, 35, 36, 117, 188 Floodplains, 50, 51, 316, 392, 397, 398
inbreeding depression, relationship to, 184–185 Floods and ecosystem management, 392, 393
patterns of, in relation to climate change, 36 Florida (USA)
Extinction risk, 42, 143, 149, 161, 187, 205, 216, 240, 251, 566 coral restoration, 343
Extractive reserve, 22–24, 30, 31 gene flow in endemic perennial shrubs, 204
Lake Okeechobee, 377
management of Florida panther, 54, 192, 214
F marine reserves, 343
Factor resolution, 233–236, 256 Pelican Island and US wildlife refuge system, 14, 61
Faculty advisor and graduate school, 542 scrub jay and metapopulation models, 222
Fahrig, L., 267, 268, 270, 273, 277, 284–287, 289, 295 University of, Tropical Conservation and Development Program,
Faith-based organizations (FBOs), 436, 437, 439–442 543 (see also Everglades National Park)
Faith for Earth Initiative and United Nations Environment, 441 Florida Audubon Society (FAS), 61
Family richness Florida panther (Puma concolor coryi), 54, 192, 214
absolute, 69 Florida scrub jay (Aphelocoma coerulescens), 222
proportional, 69 Flow regimes and river restoration, 307, 312, 313, 316, 392
proportional weighted for species richness, 69 Flushing distances, 225
Fancy, S.G., 382, 384, 385 Focused advocates, 29, 565
Farley, J., 453 Food and Agriculture Organization (FAO), 518, 524
Feasibility, as criteria for protected area selection, 290 Foote, E.N., 129
Fecundity Ford of Firth, 375
population demography, 212–214 Forest Department of India, 15
population viability analysis (PVA), 245 Forest of Harissa and Holy Forest of Our Lady of Lebanon (Lebanon),
Federal government, 12–14 441
ESA, 501, 506 Forest Preservation Act, 13, 14
history of conservation, 11–16 Forest reserves, 12, 14, 23
NEPA, 494, 498 Forests
payment for ecosystem services, 482–484 commercial, 296, 297
user fees, 459, 460 (see also Conservation law; Policy, fragmentation, 204, 273
environmental) mitigation from logging, 298
Federal lands and NEPA, 495 plantation, 53, 104, 296–298, 398, 442
Federal Register, 497, 498, 502, 503 slash, 197, 298 (see also National Forest Management Act; US
Fengshui (China), 5 Forest Service)
Ferraro, P.J., 482, 483 Forman, R., 282, 283
Fertilizer Forsman, E.D., 362
air pollution sludge, 464 Foster, M., 560
eutrophication, 311, 317 Founders (genetic), 177
logged forests, 270 Fox, H., 341, 343
Finding of No Significant Impact (FONSI) and environmental impact Framework-protocol approach and international conservation law, 510
assessment, 497 France, 17, 20, 190, 323, 430, 521
Fine-filter approach, 162 Frankel, O.H., 172, 173, 175, 179, 185
Fine-filter, climate informed conservation strategies, 160, 161 Frankham, R., 29, 45, 46, 171, 172, 177, 184–186, 188, 196, 203, 204
Finland, 187 Franklin, I.R., 173, 218, 233, 240
Index 597
Franklin, J.F., 283, 361 Genetic drift

Free trade and conservation law, 525 bottlenecks, 175–177
Freshwater fauna captive breeding strategy, 178
extinction rates, 310 concept and definition of, 172, 175–178
Freshwater habitats, 317–319, 328–332 effective population size, 177–178
Freudenberger, L., 73 Genetic restoration, 190, 192
Friesen, V.L., 204 Genetics
Fronhoffer, E.A., 222, 223 metapopulations, 212, 221–223
Frontier forests, 71, 162 small populations, 171, 172, 175, 178, 179 (see also Genetic
Fruit flies (Drosophila), 98, 172, 184, 196 diversity; Genetic drift; Molecular genetics, and assessments of)
Functional conservation areas (FCA), 75–76 Genetic stochasticity, 214
Functional ecosystems, 36, 93, 350, 363 Genetic variation
Fundamental niche, 152, 154 impact on inbreeding, 178, 179, 186–187, 233, 242
Fund-service resources, 450–452, 454 small populations, 171, 179, 188, 204
Genkai-Kato, M., 318, 319
Genotyping, of endangered species, 193
G Genuine Progress Indicator, 475
Gamma diversity, 51, 52, 326, 327 Geographic-based approach to conservation, 57, 262
Gamma rarity, 51 Geographic distribution, of species, 52, 53, 291, 292
Gandhi, Mahatma, 548 Geographic information systems (GIS), 76, 263, 269, 384–386, 563
Gap analysis program (GAP), 76, 291, 294, 386 ArcGIS, 385
GAPPS, 206 QGIS, 385
Gardner, R., 283 Geographic scales and biodiversity, 51
Geller, G.N., 66 The Geologic Time Scale 2012, 82
Gene diversity (GD), 188, 200–202, 237–239 Georgescu-Roegen, N., 472, 473
Gene flow Georgia-Pacific Corporation, 507
connectedness in population subunits, 75, 187, 220 Giant jaú (Zungaro jahu), 315
ecological and evolutionary processes, 75 Giant panda (Ailuropoda melanoleuca), 264, 265, 267, 292, 387, 420
estimation, 203, 204 Gibbons, A., 1, 2, 10
inbreeding effects, 186 Glanville fritillary butterfly (Melitaea cinxia), 187
invasive species, 163 Global Amphibian Assessment (IUCN), 292
loss of genetic variation, 186, 204 Global biodiversity strategy (World Resources Institute et al.), 70
marine reserves, 349, 350 Global carbon cycle, 130
models of, 350 Global carbon tax, 464, 465
reproductive ecology, 203 Global climate change, diagnostic fingerprint of, 135
General Agreement on Tariffs and Free Trade (GATT), 523, 524 Global climate model (GCM), 147, 149, 155, 156, 158
Generalist predators, 280, 281 Global Earth Observing System of Systems (GEOSS), 66
Generalized random tessellation stratified (GRTS) samples, 383 Global ecoregions, 40, 71, 162
Genetic adaptation (GA) Global Environmental Facility (GEF), 351, 519
to captivity, 196 Global Mammal Assessment (IUCN), 292
Genetic applications in conservation Global marker, 81, 83, 134
clarification of phylogeny, 203 Global Stratotype Section and Point (GSSP), 81–83
clarification of relatedness, 203 Global Vegetation Models (GVM), 158
clarification of taxonomy, 203 Goal-rational approaches, in conservation, 436
determination of patterns of reproductive ecology, 203 Goats, 21, 407
determination of population management units, 203, 206 God Squad, and Endangered Species Act, 501
determination of sources of wildlife products (conservation Golden-cheeked Warbler (Setophaga chrysoparia), 189
forensics), 203, 206 Golden eagle (Aquila chrysaetos), 251, 253
determination of time since past bottlenecks, 172–174 Golden toad (Incilius periglenes), 146, 147
estimation of gene flow and dispersal, 203 Goodnight, C., 173
Genetic constraints, 256 Goshawk (Accipiter gentilis), 214
Genetic diversity Governmental and Non-governmental Organization (GONGO), 20
allelic diversity, 171, 172, 174, 176, 200 Gradient, and species diversity, 50, 75, 109, 139
applications in conservation, 203–207 Gradient model, 266
average heterozygosity, 174, 175, 177 Graduate professor, 540
concept of in conservation biology, 171–172 Graduate Program in Sustainable Development and Conservation
effect on population size, 75, 172–177, 188, 189, 201 Biology (CONS), and the University of Maryland, College Park,
fixation of deleterious alleles, 171 xi, 547, 548
heritable variation, 171 Graduate record exam (GRE), 538–539
hybrids, 174, 196 Graduate school, and graduate program
inbreeding resulting in loss of, 172–174 advisor, 254, 540, 542, 568
measures of polymorphism average heterozygosity allelic , courses, 531, 532
diversity,174 curriculum, 539
Nei’s measure of, 207 financial support, 539
polymorphism, 174 record of employment or placement, 539
restoration of, 190, 192 record of publication and grantsmanship, 539
small populations, 172, 173, 178, 179, 188, 205 student awards, 546
598 Index
Granek, E., 351, 353 patches, 65, 66, 102, 143, 188, 190, 220, 222, 223, 268, 269, 272,
Gran Pantanal, 392 277, 280, 283, 287, 289
Grasslands, 37, 44, 49, 65, 86, 90, 94, 96, 142, 155, 234, 262, 277, 278, patchiness, 266, 267
295, 384, 391 predators and changes in preference, 277
Gray wolf (Canis lupus) proportion, 105, 263, 284, 286
controversial delisting, Endangered Species Act, 503 relictual, 271
hybridization with red wolf, 193, 195, 196 selection, 263–265 (see also Ivlev’s Selection Index (SI))
reintroduction to Yellowstone National Park, 461 structure, 96, 261, 264, 296, 392 (see also Habitat fragmentation:
restoration in Great Lakes states, 504 Habitat isolation: Habitat loss)
Grazers and grazing, 394 Habitat amount hypothesis, 285, 287, 295
See also Ecosystem management See also Fahrig, L.
Great Acceleration, 84 Habitat clustering, 287
Great Barrier Reef (GBR), 145, 340, 350, 351, 396–399, 402, 403, 561, Habitat connectivity, 158, 161, 278, 282, 284, 289
562 Habitat conservation areas (HCAs), 363
Great Barrier Reef Marine Park (Australia), 350–352, 402, 403 Habitat conservation plans (HCPs), 506, 507, 509
Great Barrier Reef Marine Park Authority (GBRMPA) Habitat fragmentation
ecosystem management, example of, 403 alpha diversity, 48–52, 74, 326
Representative Areas Program (RAP), 396 climate change, 126, 150
Great Basin, 404, 405, 407 corridors, 280
Great Lakes, 96, 182, 212, 312, 324, 355, 435, 504 dispersal distance and home range size, effects on, 268
Great tit (Parus major), 217, 218, 220 distinction from habitat degradation, isolation and loss, 290
Greece, 4, 5, 7 edge, relationship to, 273
Greedy Richness Algorithm, 290 field and experimental studies of, 284–289
Greenhouse gases fragment specialist species, 278
carbon dioxide, 129 invasive species, 275, 388
methane, 129 marbled murrelet (Brachyramphus marmoratus), 204
nitrous oxide, 129 metapopulation biology, gene flow and gene diversity, 189, 190, 204
ozone, 129 penetration depth and security of interior habitat, 277
water vapor, 129 percolation theory, 283–289
Green Mosques Project, 439 process and effects of, 272
Green sea turtle (Chelonia mydas), 335, 351, 353 species interactions, effects on, 287
See also Sea turtles threats from, 188, 189
Green woodpecker (Picus viridus), 266 wolf spiders, effects on, 279–281
Grenyer, R., 56, 57, 59, 68 Habitat generalists, 52, 53
Grid sampling, 382 Habitat islands, 279
Grinnell, J., 125, 126, 152 Habitat isolation, 262, 272, 273, 280
Grizzly bear (Ursus arctos horribilis), 141 Habitat loss
Gross National Happiness Index, 475 climate change, 150, 316
Groundfish, 344 distinction from habitat degradation, fragmentation and isolation,
See also Demersal species 126
Group on Earth Observations (GEO), 66 eastern massasauga (Sistrurus catenatus catenatus), 271
Group on Earth Observations Biodiversity Observation Network edge, relationship to, 279
(GEOBON), 66 fragmentation, relationship to, 278–289
Growling grass frog (Litoria raniformis), 212, 223–226 habitat generalists, effects on, 53
Guam, 98, 101 hotspots, 66
Guha, R., 8, 15–17, 21 wolf spiders, effects on, 279
Gulf of Maine, 332, 365, 368, 369 Habitat modifications, 104, 108, 278, 506
Gypsy moth (Lymantria dispar), 100, 103 Habitat patches
corridors and connectivity of, 75
habitat and landscape models, 281, 297
H habitat selection and movement by animals, 219, 279
Habitat invasive species, 272
amount, 273, 277, 284, 285, 287, 288, 295 metapopulation dynamics and movement, 143
availability, 262, 284, 288, 289 percolation theory, 283–287
classification framework, 104 Habitat specialists, 41, 51–53, 103, 190, 378
complexity, 261, 287, 313, 333 Habitat split, 325
configuration, 273, 281, 287–289, 291 Habitat suitability models (HSMs), 152, 265, 266, 289, 291, 292, 387
definition and importance of, 261 Habitat transitions – Markov model, 268–270
design of nature reserves, 264 Habitat types, 41, 45, 51, 66, 104, 105, 113, 266, 294, 326, 333, 386,
destruction, 66, 125, 126, 158, 214, 256, 272, 279, 284, 332, 477
335–337, 339, 340, 444 Haddad, N.M., 273, 280, 282
dynamics, 268 Haida tribes, 4
heterogeneity, 221, 262, 266, 267, 294, 391, 392 Halbert, N., 173, 174
loss, isolation and fragmentation, 280 Haleakala silversword (Argyroxiphium sandwicense), 54
matrix, 278 Half-Earth: Our Planet’s Fight for Life, 117
non-reserve lands and management, 302 Hannah, L., 125, 127, 134, 161, 165
Index 599
Hansen, L., 134, 377–380 Hoffman, M., 40, 42

Haplotypes, 172, 204 Hogna helluo, 279
Happy Planet Index, 475 See also Edge effects; Wolf spider
Harbor porpoise (Phocoena phocoena), 348 Holdridge life zones, 145, 146
Hardin, G., 425, 468 Holling, C. S., 408
Harmonic mean, 102, 178 Holocene Epoch, 82
Harris, G., 66 Holt, R.D., 280
Harvest refugia, 350 Homozygosity and lethal genes, 177, 201
See also No-take zones Honest broker, scientist role in policy, 490
Hawaii (USA) Honnay, O., 38, 188, 189
CO monitoring, 129, 131 Hotspots, of biodiversity and endangered species, 57, 69, 110, 111
ecosystem management, 93 House Finch (Haemorhous mexicanus), 103
endemic species, 54, 348 House mouse (Mus musculus), 111
invasive species, 93 House wren (Troglodytes aedon), 227, 277
restricted distribution of plants, 54–57 Hudson, M.E., 182
Year of the Hawaiian Forest, 407 Huffaker, C. B., 220, 221
Hawai’i Island (Hawaii, USA), 93 Human benefits and cost-benefit analysis, 421
He, P.M., 261, 262, 287, 288 See also Human relationships and interests, and ecosystem
Headwaters, 59, 309, 317 management
Heard, G., 223–226 Human Ecosystem Framework (HES), 107, 108
Heat capacity, 107, 308 Human Genome Project, 181
Heath hen (Tympanuchus cupido cupido), 214 Human impact matrix, 365
Hedges, C., 10 Humpback whale (Megaptera novaeangliae), 348
Hedonic property model, 457 Hungerford’s crawling water beetle (Brychius hungerfordi), 460
Heimlich, J.E., 549 Hunter, M.L., 2, 161
Herbicides, 234 Hurtt, G.C., 95, 154, 155, 165
managing succession, use in, 270 Hutchinson, G.E., 151, 161
restoration of endangered species, use in, 235 Hybridization
water pollution, 334 (see also Pesticides) in captive populations, 196–202
Herbivores, and ecosystem management, 393–395 fitness effects of, 193
Hernández-Matias, A., 247, 249, 250 genetic determination of, 174
Heterogeneity Genetically Modified Organisms (GMOs), risks from subunits
environmental, 220, 266, 267 rare species, threats to, 193
fire, effect on, 391, 392 red wolf (Canis rufus) with coyotes (C. latrans), 193, 195, 196
flooding, effect on, 392 Hybrids
genetic, in metapopulations, 221, 256 coyote-wolf, 195
habitat, 221, 262, 266, 267, 294, 391, 392 definition, 193
invasive species, 102, 275 endangered species act and protection of, 193, 424
metapopulations, 220, 221 Florida panther (Puma concolor coryi), 192
spatial, 102, 219, 266 genetic restoration, role in, 190, 192
temporal, 267 red wolf (Canis rufus), 193, 195, 196
wetland, 324 wild and domestic turkey, 196
Heterozygosity Hydric (water-logged) soils, 323
breeding system, relationship to, 172, 176, 183–185, 187, 192, 201 Hydrologic connectivity, 316
captive breeding, 184, 200, 201 Hydrologic cycle
effective population size, 175, 177–178, 201 rapid and energetic, due to climate change, 133
electrophoresis, measurement by, 180 Hydrologic regime
forest trees, reduction due to logging, 206 invasive fish, 100
gene flow, 186, 187 wetland subunits, 320
loss due to genetic drift, 172, 175–178 Hydrology, 76, 322, 324
loss due to inbreeding, 184, 185, 187, 256 Hydroperiods, 324
management decisions, 391 Hypericum cumulicola, 204
measure of genetic diversity, 174
measure of inbreeding, 183, 185, 187
panmictic index, mini- and microsatellites, 183 I
small populations, 172, 175 Iberian lynx (Lynx pardinus), 297, 298
Hey, J., 47 Ice anomaly, 135
Hidden hurdle, becoming a colleague, 533–534 Ice extent, 135
Higgins, J., 75, 322 Ice thickness, 147
High Biodiversity Wilderness Area (HBWA), 71, 162 Idaho (USA), 291, 294, 504
Hinduism, 431–434 Illinois, 112, 186, 192, 229, 231
Hispid cotton rat (Sigmodon hispidus), 277 Immanence and Islam, 431
Historical “experiments,” and ecosystem management, 386 Immigration, 143, 197, 212, 213, 215, 231, 287, 482
Hobbs, R.J., 89, 90, 92, 95, 109, 110, 116, 272 See also Populations
Hoegh-Guldberg, O., 133, 144, 145, 161 Impacts and alternatives analysis, and environmental impact assessment,
Hof, J., 243, 246, 284, 285 498
600 Index
Inbreeding, 186–187 International Court of Justice (ICJ), 510

definition, 182 International Dolphin Conservation Act, 524
maximum avoidance of, 184, 201, 202 International Dolphin Conservation Program, 524
measurement of in small populations, 183, 184 International Geological Congress, 82
as nonrandom mating, 183 International habitat reserve programs (IHRPs), 483
by population subdivision, 183 International Smart Gear Competition, 349, 542
Inbreeding coefficient, 172, 173, 183, 185–188, 200, 202 International treaties, 38, 510, 519–521
Inbreeding depression, 171–173, 184–188, 190, 197, 201, 202, 239 International Union for the Conservation of Nature (IUCN)
Incidental take, of endangered species, 506, 507 Protected Area Management Categories, 39
Inclusion and diversity, in conservation biology, 557–560 Red List Categories and Criteria, 39, 42, 336, 513
Independent Research Study, 533 Red List Index (RLI), 39, 40, 42
See also Conservation education Red List of Ecosystem Categories, 39, 68
Index of Sustainable Economic Welfare (ISEW), 475 Species Survival Commission, 20, 39, 40
India, 15, 21, 62–64, 204, 317, 379, 381, 435, 440 International Union for the Protection of Nature (IUPN), 20
Indianapolis Power and Light Company, 464 International Whaling Commission (IWC), 205, 348
Indian grass (Sorghastrum nutans), 391 Interpersonal skills and careers in conservation biology, 533, 548–551
Indian Ocean, 176, 339, 341, 351 Interviews, and data on attitudes and values, 420
Indicator species, 69 Intrinsic values
Indicator taxa, 55 approaches to, 426
Indigenous and Community Conservation Area (ICCA), 120 ecocentrism, 427–428
Indigenous species, 125, 234 ethical basis of conservation and conservation biology, 421, 429,
Indonesia, 40, 48, 53, 341, 342, 344, 442–444, 565 444
Industrial revolution, 8, 83, 117, 450 instrumental values and, 418–421, 444
Inouye, D., 136 religious traditions, 428–436
Input-oriented problems and freshwater habitats, 332 Romantic transcendentalism, 12
Insectivorous plants Introgression, 102, 193, 195, 196
sundew (Drosera spp.), 324 Invasion biology, theory of, 98, 102–103
Venus fly trap (Dionaea muscipula), 324 Invasive species, 102, 104–106
Institute for Theological Encounter with Science and Technology barrier zones, 103
(ITEST), 437 characteristics, 98, 99, 142
Instrumental values, 418–421, 444 competitive advantage of, 141, 142
Intact habitat and landscape, 271 dispersal, 98, 103, 142
Integrated conservation and development project (ICDP), 479–484 ecological match of, 100, 101
See also Community-based Natural Resource Management establishment, 96, 98, 99, 104
(CNRMs) interactions between and climate change, 142
Integrated ecosystem assessment (IEA), 368, 370, 381 patterns of spread, 98, 99, 102
Integrated Graduate Education and Research Traineeship (IGERT), 543 pre-adaptation to environment, 100
Integrative nature, of conservation biology, 28, 29 recommendations for control of, 106
Integro-difference equation (IDE) models and invasive species, 103 shifts in plant community composition due to, 142
Interagency cooperation and ecosystem management, 362, 501 spatially distributed populations, 98
Interagency Scientific Committee (ISC), 363 spread of, 95, 141
Intergenerational equity, 492, 510, 521 stages of establishment, 94, 96
and international conservation, 521, 522 Iowa (USA), 16, 27, 49, 535
Intergovernmental Panel on Climate Change (IPCC) Irrawaddy dolphin (Orcaella brevirostris), 348
Representative Concentration Pathways (RCP), 154 Islam, 431, 434, 439, 440
Intergovernmental Science-Policy Platform on Biodiversity and Islamic Declaration on Climate Change, 440
Ecosystem Services (IPBES), 165, 476, 477 Islamic Foundation for Ecology and Environmental Services (IFEES),
Interior species, 273, 274, 277, 279, 280 439, 440
Intermediate disturbance hypothesis, 295 Islamic Principles for the Conservation of the Natural Environment, 431
International Animal Rescue (IAR), 443 Island biogeography, 28, 222, 295
International Commission on Stratigraphy (ICS), 81, 82 Island fox (Urocyon littoralis), 251–253
International conservation law Island species and endemism, 54, 55, 59, 351
agreements, 492, 510, 512, 513, 519 Isle Royale National Park, 382, 504
charters, 510 Isolation by distance model, 273, 281
compliance Isolation models, 271
coercive measures, 521 Israel, 5, 430
positive incentives, 520, 521 Issue advocates, scientist role in policy, 490, 491
declarations, 512 Italy, 82, 266, 267
effectiveness, 519 Ives, C.D., 111
hard law, 518, 519 Ivlev’s Selection Index (SI), 263
implementation of, 519, 520 Iwamura, T., 162
protocols, 510, 513, 519, 520
soft law, 518, 519
Stockholm Conference, 511–513 J
treaties, 492, 510, 512, 513, 518–520 Jack pine (Pinus banksiana), 52, 253, 254, 257
International Convention for the Preservation of Useful Birds, 18 Jackson, W.H., 10
Index 601
Jacquemyn, H., 188, 189 L

James, S., 433 Lacey Act, US, 60, 494
Japan, 205 Lack, D., 217, 220
Japan, Sea of, 205 Lacy, R.C., 201, 202, 215, 240
Jenkins, C., 66 La Jolla (California) agreement of, 524
Jenkyn, T.W., 83 Lake Erie, Great Lakes (US), 380
Jewish Global Environmental Network (JGEN), 437 Lake Okeechobee, Florida, 377
Jewish Theological Seminary, 437 Lakes
Job skills as signals to employers, 552, 554, 555 hypolimnion, 319
Joshua tree, 27 lacustrine zone, 314
Journal of Wildlife Management (JWM), 26 riverine zone, 314
Judaism, 5, 430, 431 transition zone, 26, 314, 318
Judeo-Christian Stewardship Environmental Ethic, 429 Lake sturgeon (Acipenser fulvescens), 435, 436
Judeo-Christian tradition and conservation values, 428 Lakota people, of US Great Plains, and indigenous belief systems, 435
Juniper (Juniperus spp.), 190, 404 Lambda, 220, 243, 250
Justinian, 307 Lamoreux, J., 55–57, 59, 68
Juvenile survival and inbreeding depression, 185 Land-based pollutants, of marine habitats, 334
See also Infant mortality, in cheetahs Land Degradation Neutrality and Convention to Combat Desertification
(UNCCD), 518
Land ethic, 16, 28, 427
K Lande, R., 172, 362
Kabilsingh, C., 440 LANDIS-II, 270
Kainer, K.A., 474, 541, 543–545 Land Sabbath, and Judeo Christian Stewardship Ethic, 430
Kalimantan, 443, 565 Landscape
Kareiva, P., 30, 31, 62, 119, 271 concept of in conservation biology, 22, 62, 63, 261–302
Karma, 432, 433 definition of, 262
Karpf Stock, 9 ecosystem management, 377, 378, 382
Keller, L.F., 183, 187, 188 genetics, 51, 188–190, 203
Kellert, S.R., 113, 417, 419, 420, 422, 424 Landscape ecology, 283
Kemp’s ridley turtle (Lepidochelys kempii), 203 Landscape-level conservation, 62–64
See also Sea turtles Landscape pattern, 279, 282, 283, 391
Kennebec River, Maine (USA) Landscape planning, 285
Edwards Dam, 316 Large-flowered fiddleneck (Amsinckia grandiflora), 234–236
Fort Halifax Dam, 316 Laser leveling, 392
Kenya Last of the Wild (LW), 71, 72, 162
Amboseli Community Wildlife Tourism Project, 481 Latitude
A Rocha Kenya, 437 biodiversity, 57, 69
marine reserves, 349–355, 371, 372 Laudato Si’, Encyclical Letter, 437–439
poaching, 514, 515 Lavery, T.J., 346, 347
Keoladeo Ghana National Park, 62 Law of Dependent Co-Arising, 432
Key Biodiversity Areas, 40, 59–60 Law on Biodiversity (Brazil), 524
Keystone species, 141, 292, 336 Lawsuits, see Litigation
Khilafa (trusteeship) and Islam, 431 Leadbeater’s possum (Gymnobelideus leadbeateri), 68
Killer whales (Orcinus orca), 203, 205 Leal, D.R., 4, 466, 468, 471, 472
Kilpatrick, J., 70 Learning and action platform, 543, 544
King Castle, Niepołomice (Poland), 82 Least tern (Sternula antillarum), 313
Kinship Legally empowered science, conservation biology as, 489
coefficient (kij), 198, 199 Lentic systems, 309
mean (mk), 199, 200, 202, 237, 238 Leopold, Aldo, 16–17, 26–29, 32, 116, 196, 224, 271, 302, 427, 428
population mean, 199, 200, 202 See also Ecocentrism
probability, 198 Lesser sand eel (Ammodytes spp. and Gymnammodytes spp.), 374
Kirtland’s warbler (Setophaga kirtlandii), 27, 52, 253–257 Lethal load, 176
Kirtland’s Warbler Recovery Team (KWRT), 254 Levins model, 222
Kiss, A., 481 Levins, R., 221, 222
Komodo dragon (Varanus komodoensis), 48 Lewis and Clark Expedition, 10
Komodo National Park (KNP), 341, 344 Lewis, S.L., 81–84
Konza, Prairie (Kansas, USA), 93, 535 Lewison, R.L., 343, 348, 359
Koontz, T., 359 Liangshan Mountains, China, 265
Koteen, L., 141 Life history traits, in invasive species, 99, 142
Kowarik, I., 96, 97, 108, 110–112, 115 Life stages, in invasive species, 240, 242–244
Krieger, M., 413 Life stages, in models of population viability analysis, 245
Krill, 346, 347 Light Detection and Ranging (LiDAR) technology, 165, 266–267
Kueffer, C., 104–106 Limited access, and mitigation, 300
Kuznets Curve, 472 Limpets, and ecosystem modeling, 388–391
Kyoto Protocol Lincoln-Petersen Index (Lincoln-Petersen Estimator), 229
emission reduction units, 164 Lindenmayer, D.B., 68, 262, 271, 272, 296, 297, 302
602 Index
Linear habitat, 333 Mandarte Island, 183, 187

Lindsay, D.L., 190–191 Manel, S., 189
Linnaeus, C., 45 Manley, P., 384, 389
Lion (Panthera leo), 54, 192, 345, 479 Manolis, J., 547, 566
Liroff, R.A., 495, 498 Manuel Antonio Park, 459
Listing process, and Endangered Species Act, 500–505 Marbled murrelet (Brachyramphus marmoratus), 204
Litigation, 363, 425, 493, 499, 505, 507 Marble-in-a-cup model, 318
See also Conservation law; National Environmental Policy Act Marine environments, 57, 307, 335, 351, 355, 366, 367, 374
(NEPA) See also Dolphins; Fishing industry; Marine Mammal Protection Act
Little River Band of Ottawa Indians (LRBOI), 436 Marine Mammal Protection Act (MMPA), 522, 523, 526
Livestock grazing, 90, 236, 243, 398, 404, 405 Marine reserves and protected areas, 349–355
Living Modified Organisms (LMOs), 518 Market-based incentives and economic institutions
Loarie, S.R., 155, 156, 161 governmental subsidies, 465
Local adaptation, 104, 188 governmental taxation, 463, 464
Logistic population growth, 213, 214, 217–219 tradeable permits, 465–466
Long-term ecological research (LTER), 107 Market equilibrium, 454
Long-term natural repositories, and data on ecosystems, 382, 387–388 Market failure, 455, 457, 461, 463
Loomis, J.B., 457–459 Market goods and biotic resources, 419
Los Angeles, and plastic trees, 412 Market incentives and government coordination, 463, 467
See also California Market price and marketplace, 450, 454, 457, 458, 465
Lotic systems, 309 Marley Wood, England (UK), 217, 218
Louisiana (USA) Marshall, S.D., 278–280
Weyerhaeuser Company v. United States Fish and Wildlife Service, Marsh, G.P., 83
505 Martha’s Vineyard, Massachusetts (USA), 214
Lovejoy, T., 125, 127, 165, 282 Martinez-Alier, J., 472
Lyons, J.R., 37, 363, 547, 548 Martinich, J.A., 547, 548
Marvier, M., 30, 31, 62, 119
Mascaro, J., 93, 105
M Mascia, M., 549
Macroeconomics, 450 Maslin, M., 81–84
Macrohabitat, in aquatic ecosystems, 75, 320, 321 Massachusetts (USA), 13, 214, 520
Madagascar, 63, 461 Mast production, 141
Madsen, T., 190, 192 Mather, S., 428
Maine (USA) Mattson, W.J., 98, 141
dam removal, 316, 392, 393 Maui
diadromous fish species, 316 East Maui Watershed Partnership, 407
vernal pools, 325, 326 Maui dolphin (Cephalorhynchus hectori maui), 348
Mainland-island metapopulation model, 222 Mauna Loa Observatory, 129, 131
Major federal action and NEPA, 493, 495 Mauritius, 176
Major histocompatibility complex (MHC), 192 Mauritius kestrel, 176, 177
Malaysia, 40, 63, 69 Maximum sustainable yield (MSY), 343
Malcolm, J.R., 57, 158 Mayr, E., 46
Malthus, T., 472 McCormack, J., 6
Mammals and biodiversity, 55 McFarlane, A., 8
Managed breeding, 176, 197, 198 McGee, W.J., 15
See also Captive breeding; Managed matings McKinley, W., 13
Managed population size, 201 McKinney, M., 106–109, 111, 113
Management, 371 McNeill, J.R., 4
captive breeding, 197, 198, 200, 201 Mean generation time, 201
decision analysis and small populations, 216 Mean kinship (MK), 200, 202
ecosystem management, 93–95, 359–408, 454, 499, 533 MK strategy, in captive breeding, 202, 237, 238
freshwater habitats, 317–320, 322 Medicine, tropical plants and pharmaceutical agents, 223, 237, 418, 435,
genetic subunits, 203, 204, 216 514
habitat on non-reserve lands, 302 Mediterranean fruit fly (Ceratitis capitata), 98
hybridization and introgression, mitigation of, 193, 195, 196 Mediterranean Sea and invasive species, 513
marine reserves, 349, 350, 355 Megadiversity and megadiversity countries, 69, 71
multiple threats to small and declining populations, 144–145 Meijaard, E., 565
nonindigenous invasive species, 100 Mekong River, 348
populations of spatially disjunct subunits, 220 Memoranda of Agreement (MOA) and Endangered Species Act, 255,
resource management biodiversity at landscape levels, 62 507, 509
succession, 270–271, 391 (see also Adaptive management) Memorandum of Understanding (MOU), 254, 510
Management training models, 545 Mendes, C., 23, 24
Management units (MU), in conservation, 48, 203 Menke, J.W., 236
Man and Nature, 83 Mentors and education in conservation biology, 540
Man and Biosphere Program, 24 Mentor-student relationships, 537
Index 603
Metapopulations and metapopulation theory Molecular genetics and assessment of variation, 46

conditions to meet criteria of, 222 Mollusc (Tympanotonos fuscatus), 434
corridors and connections of subunits, 220 Mona monkey (Cercopithecus mona), 434
definition and concept of, 221–223 Mongoose (Herpestes spp.), 100
desert bighorn sheep (Ovis canadensis nelsoni), 142, 144 Monogamous populations and sex ratio, 178
Florida scrub jay (Aphelocoma coerulescens) as case history of Montana (USA), 2, 10, 96, 141, 236, 239, 262–264, 298, 299, 416,
genetics, 222 466–468, 504
Glanville fritillary butterfly (Melitaea cinxia), 187 Montego Bay Convention, 513
models of population viability analysis (PVA) of, 233, 240 Moon, K., 29, 560
origins of, 220, 221 Moose (Alces alces), 262, 382
theoretical development, 221–223 Moral agents, 421
Mexico, 40–42, 50, 69, 238–240, 348, 439, 465, 511, 522 Moral extensionism, 422–428
Michigan (USA) Moral obligation and economic analysis, 421, 431
amphibian biodiversity, 325 Moral philosophy and economics, 419
Audubon Society, 254 Moral subjects, 421, 430, 492
Department of Natural Resources, 254, 435 Moral value, 15, 421–429, 433, 492
Isle Royale National Park, 382 Moran, T., 10, 11
Kirtland’s warbler, 27, 52, 253–257 Moreton Bay (Australia), 328, 331
Manistee River, 435, 436 Morris, F.O., 8, 9
timber industry, 13 Morris K. Udall Scholarships in Environmental Policy, 536
wild turkey reintroduction, 196 Morris, W., 8
wolf restoration, 195, 196 Moses, and Biblical Sabbath, 5
Microclimates, 139, 154, 160, 297 Mount Desert Island National Laboratory, 535
Microeconomics, 450, 453–456 Mount Taylor Ranger District, New Mexico (USA), 289, 290
Microsatellites, 174, 180, 181, 187, 204 Mountain gorilla (Gorilla beringei beringei), 160, 418
Midland populations, 185, 222 Mountain lion (Puma concolor), 54, 192, 225
Mitigation of bycatch, 344, 345 Mountain pine beetle (Dendroctonus ponderosae), 298
Mitigation of climate change, 134, 139, 141, 152, 158, 160 Mowing and grassland restoration, 107, 404
Mitigation of environmental impacts, 492 Muir, M.J., 11–15, 22, 32, 428, 540, 551, 552, 554, 555
Mitigation of habitat loss, range displacement, 298–302 Multidisciplinary nature, of conservation biology, 29
Mitigation of wetland loss, 461 Multiple use and management of non-reserve lands, 302
Migratory Bird Treaty Act (MBTA), 18, 494, 500 Multiple-use module (MUM), 320, 350
Migratory species, international protection of, 18–19 Multi-stakeholder processes and conservation education, 544, 546
Milankovitch cycles, climate change as a result of, 127 Murray Rivers, 396
Millennium Development Goals (MDGs), 517, 519 Muskrat (Ondatra zibethicus), 102, 323, 324, 471
See also Sustainable Development Goals Mussels, 96, 310
Millennium Ecosystem Assessment (MEA), 85, 87, 262, 364, 365, 396, See also Zebra mussel
450, 451, 477 Myers, N., 62, 63
Miniat, C.F., 393 Myrica faya, 101, 102
Minimum critical size, of habitat, 392
Minimum population size, 173, 233, 240
Minimum viable population (MVP) N
conservation biology and concept of, 233 Naess, A., 29, 425, 426, 432
persistence likelihood, 240 Nagoya Protocol, and Convention on Biological Diversity, 518, 520,
PVA, 240 524, 525
Minisatellites, 181 Nasr, S.H., 431
Minke whale (Balaenoptera acutorostrata), 205, 206 National Academy of Sciences, 132, 137, 343, 403, 458, 491
Minnesota (USA) National Aeronautics and Space Administration (NASA)
Department of Natural Resources, 547 Carbon Monitoring System, 165, 457
Minority students, in conservation biology, 558 Earth Exchange (NEX), 154
Mission-driven/mission-oriented character of conservation biology, 411 Global Ecosystem Dynamic Investigation (GEDI), 165, 266
Mississippi River, 242, 466 National Association of Environmental Professionals, 498
Missouri (USA), 196, 439 National Audubon Society (NAS), 9, 471
Missouri River, 64, 65 National Council of Churches, 437
Mites National Environmental Policy Act (NEPA)
six-spotted (Eotetranychus sexmaculatus), 221 application of, 76, 420, 493
Typhlodromus occidentalis, 220, 221 components, 76, 526
Mitigation, of habitat disturbance, 160, 298, 300, 302 development of conservation law, 75, 493
Mitochondrial DNA (mtDNA), 181, 182, 204, 205 ecosystem management, 361, 499
Möbius, K., 84 environmental impact assessment (EIA), 495–499
Moheli Marine Park environmental impact statement (EIS), 420, 493
awards, 353 federal lands, 494, 495
ecoguards, 353 implementation in other countries, 497
Journée de la Tortue (Day of the Turtle), 353 preparation time, 498
Mwana wa Nyamba (The Baby Turtle), 353 shortcomings of, 495
604 Index
National Forest Management Act (NFMA), 362 Nonexclusive goods (nonexcludable goods), 450, 452–453, 467
National Marine Fisheries Service (NMFS), 345, 494, 500–503, 505, Nongovernmental organizations (NGOs), 39, 113, 116, 120, 254, 255,
524 436, 441, 442, 468, 469, 471, 481, 483, 493, 516, 519, 522, 523,
See also NOAA Fisheries 525, 539, 544, 547, 563
National Marine Sanctuary Program (NMSP), 343 Non-native species, 95–106, 110, 111, 234, 307, 312, 313, 377
National Oceanic and Atmospheric Administration (NOAA) See also Invasive species
NOAA Fisheries, 500 Non-response bias and surveys, 420
National parks, 9–11, 13, 14, 16, 17, 21, 22, 24, 51, 54, 61–63, 116, 120, Nonrival goods, 450, 452–453
125, 126, 128–129, 138, 141, 159, 160, 166, 173, 174, 178, 218, Non-spatial models and invasive species, 85, 90, 94–106, 111, 125, 141,
219, 239, 294, 341, 344, 362, 377–380, 382, 384, 385, 388, 413, 142, 163, 182, 275, 311, 388, 546
414, 426, 461, 480, 496, 504, 534, 550 Non-traditional education, 540–541
See also Bonaire Marine Park (Venezuela); Costa Rica; Everglades Non-use values, 419, 460
National Park; Marine reserves and protected areas; Nature Nordhaus, T., 436
reserves; Yellowstone National Park Normative postulates of conservation biology, 28, 30, 411
National Park Service (NPS), 10, 11, 14, 54, 128, 138, 178, 239, 362, Norse, E., 44
382, 426, 459, 494, 496, 534, 550 North American Free Trade Agreement (NAFTA), 478
National Religious Partnership for the Environment (NRPE), 437 North American Wetlands Conservation Act (NAWCA), 465
National Research Council, 324 Northern Arizona University (USA), 543
National Resource Conservation Service, 535 Northern leopard frog (Lithobates pipiens), 229, 231
National Science Foundation Long-term Ecological Research (LTER) Northern spotted owl v. Hodel, 288, 289, 361–364
program, 107 North Sea, 139, 140, 367, 374, 375, 400
National Science Foundation (NSF), 107, 534, 543, 547 Noss, R., 147, 262, 412, 490, 506, 552, 565
National Wildlife Refuge System (US), 14 No surface occupancy, 300
Natural catastrophes, 214–217, 220 No-surprises policy, 501, 507
See also Fire No-take zones, 350
Natural History and Antiquities of Selborne, 7 See also Harvest refugia
Naturalistic fallacy, 416 Notice of availability (NOA) and environmental impact statement, 498
Naturalization Society (New Zealand), 98 Notice of intent (NOI), and environmental impact statement, 497
The Natural Regulation of Animal Numbers, 217 Notice of review (NOR), 502
National Resource Conservation Service (NRCS), 535 See also Candidate species
Natural resource law, 492–494 Novel ecosystems
Natural resource rights alternative stable states, 94
access, 467 characteristics of, 88, 90, 95
alienation, 468 definition of, 90
exclusion, 468 first description of, 31
management, 468 formation of, 31
withdrawl, 468 stability of, 94
Natural rights, 119 Numerical complexity score and marine habitats, 332
Natural selection, 45, 172, 188 Nyaru Menteng Orangutan Rehabilitation Center, 443
Nature conservancy, The, 75, 109, 115, 117, 119, 289, 302, 320, 321,
341, 361, 385, 534, 550, 565
Nature reserve design and size design, 291 O
Nature reserves, 9, 76, 264, 289, 326 O’Brien, S.J., 171–206
See also Marine reserves and protected areas Occupancy probability, 563
Negativistic attitude, 417 Occupancy theory and modeling, 229–230
Neighborhood movement rules, 282 Oelschlaeger, M., 412, 490
Neoclassical economics, 449, 450, 453, 456, 472–473 Ohio (USA), 278, 280, 317
Neoclassical economics challenges to, 472–473 Oil drilling, 92, 301, 351
Neoprotectionism, 117–119, 121 Oil leasing on private property, 457–458, 469–471, 492
Nepal, 64 See also Audubon Society
NEPAssist geospatial IT tool, 499 Okapi Conservation Project, 198
Neutral Landscape Models, 281, 282 Okapi (Okapia johnstoni), 197–200, 202
New Forest (England), 6 Okavango River, 392
New Mexico (USA), 16, 17, 239, 289, 290, 317 Okeechobee, Lake, 377
Newsome, C., 558, 559 See also Everglades National Park
Newt, Great Crested (Triturus cristatus), 182, 231 Oligonucleotides, 179
New Zealand, 41, 96, 98, 182, 313, 499 Olive ridley turtle (Lepidochelys olivacea), 203
New Zealand mud snail (Potamopyrgus antipodarum), 182 See also Sea turtles
Nicaragua, 69 One Earth Sangha, and Green Sangha, 441
Niche, fundamental, 152, 154 O’Neil, R., 283, 376
Niche, realized, 152, 154 Ontario (Canada), 287, 325, 380
Niemelä, P., 98 Open-ended questions, and surveys, 419
Nirvana and Buddhism, 432–434, 440–441 Operation Uhai, 61
Nixon, R., 493 Opportunity cost and microeconomics, 257, 419, 450, 453–457, 466,
NOAA Fisheries, 500 467, 483, 484
Noble, J.W., 13 Optimal niche gestalt, 257
Index 605
Optimization models, and ecosystem management, 395 Percolation cluster, 283, 284
Option values, 419, 467 Percolation theory, 282–290
Orangutan Conservancy (OC), 443 Pérez, H., 540, 551, 552
Orangutan Foundation International (OFI), 443 Performance payments, 482, 483
Orangutan Outreach, 443 See also Payments for Ecosystem Services
Orangutan (Pongo pygmaeus), 442–444 Persistence likelihood of a population, 240
See also Sumatran orangutan (Pongo abelii) Persistence threshold, 285, 289
Oregon, 125, 188, 362, 363, 404, 415, 495 Persistence time and population size, 133, 215, 220
Oregon Endangered Species Task Force (OESTF), 362 Personal mission statement, 532, 538
Oregon jay (Perisoreus obscurus), 125, 126 Peru, 58, 59
Oreskes, N., 491 Pesticides, 82, 176, 334, 396, 465, 474
Orr, D., 436 See also Agriculture; Herbicides
Ostrom, E., 467 Petulla, J., 10, 11, 13, 14, 509
Outbreeding depression, 188 pH
Outer continental shelf, 333 acidification of aquatic ecosystems, 311, 317
Ovenbird (Seiurus aurocapillus), 225, 227 climate change effects in oceans, 133, 335
Oxford Centre for Hindu Studies, 431, 440 Phenomenologically significant animal, 420
Philippines, 63, 350, 354, 521
Phillips Petroleum, 299
P Phoenix, Arizona (USA), 107
Pacific Rivers Council, 406 Phosphorus and eutrophication, 67, 82, 93, 311, 317–319, 324, 325
Pacu (Piaractus mesopotamicus), 315 Photic zone, 308
Paddlefish (Polyodon spathula), 466, 467 Phylogenetic Diversity Index, 28, 47, 59, 60, 181, 420
Palawan Biosphere Reserve (Philippines), 350 Phylogenetic species concept (PSC), 46, 47
Palmer, M., 442 Pielke, R.A. Jr., 490, 491, 566
Panama, 387, 389, 524 Pielou’s index of evenness, 49
Panama Declaration, 524 Pimm, S., 44, 54, 66
Panama lake, 387 Pinchot, G., 12–16, 18, 428
Panmictic index, 183, 184 Pine (Pinus spp.)
Paraguay River, 322, 392 corridors, 106
Parasites, 96, 136, 253, 256, 257, 277, 371, 424 invasive species, 99
Paris Agreement Jack (Pinus banksiana) and Kirtland’s warbler (Setophaga
Nationally Determined Contributions (NDCs), 164, 165 (see also kirtlandi), 52, 253
UNFCCC) lodgepole (Pinus contorta), 141, 298
Parker, S., 109, 112, 115–117 longleaf (Pinus palustris), 392, 506
Parmesan, C., 134–136 ponderosa (Pinus ponderosa), 298
Partnerships scots (Pinus sylvestris), 82
defined, 329, 396 white (Pinus strobus), 141
ecosystem management, role in, 66, 329, 331, 396, 406–408, 441, whitebark (Pinus albicaulis), 141
517 Pintado (Pseudoplatystoma corruscans), 315
Paspalum grass (Paspalum spp.), 62 Piping plover (Charadrius melodus), 313
Patch contrast, 274, 275 Plant functional types, for ecosystem modeling, 154
Patch dynamics, 266–271, 294 Plants
Patch models, 266 Endangered Species Act, 250, 302, 500
Patchiness, of habitat arrangement, 266, 267 ESA, 114
Paterson, B., 433 hybridization, 102
Pathogens, spread of under climate change, 100, 139–141 inbreeding depression, 185, 188
Pavlik, B.M., 233–235 nonindigenous species, 100, 311
Payments for Ecosystem Services (PES) direct payments strategy, 480, outbreeding depression, 188
482–484 trend analysis and endangered species, 233, 234 (see also Tropical
Pearson correlations, 55, 56 plants; Vegetation)
Pedigree analysis Plato, 4, 5, 45
kinship coefficient, 198, 200 Polar bear (Ursus maritimus), 147–149
mean kinship, 197–200 (see also Captive Breeding gene diversity) Policy, environmental
Pedigree inbreeding, 183 definition and history of, 493–495
Pe’er, G., 240, 241, 249, 251 ecosystem management and policy evaluation models, 75, 361, 420,
Peer review and published work, 88, 134, 251, 370, 387, 491, 534, 540, 456, 484, 490, 493, 494, 499, 526
565 governmental regulation, 456, 463, 526
Pelagic species, 314, 344 governmental subsidies, 465
Pelican Island Wildlife Sanctuary (Florida, USA), 14, 61 governmental taxation, 463, 464
Peloponnesus, 7 sustainable development, 30, 166, 511, 517, 547
Penetration depths and edge habitats, 277 tradable permits, 465
People’s Climate March, 437, 440 voluntary programs, 463 (see also Conservation law; Federal
People’s Climate Shabbat (Sabbath), 437 government)
606 Index
Pollen Population genetics, 171, 174, 175, 188, 190, 200, 219
index to gene flow, 190 Population Management Plan (PMP), 197
sediment cores, 387, 389 Population mean kinship (MK), 199, 200, 202
Pollinators See also Mean Kinship (MK)
deforestation, 176 Population regulation
effect on Index of Sustainable Economic Welfare (ISEW), 475 contrasted with population control, 217
Ehrlich Identity, 478 density dependent mechanisms, 213, 218
freshwater habitats, 317 density independent mechanisms, 214–218
marine habitats, 137 extrinsic factors, 149, 211, 214
pollution, 8, 15, 85, 310, 311, 333–335, 396, 429 (see also intrinsic factors, 149, 211, 214
Acidification; Eutrophication, and aquatic ecosystems) Population sinks, 222, 223, 247
synchronization of, with phenological changes in plants, 137 Population subdivision, 183, 220–223
Polluter pays principle, 463, 465 Population viability analysis (PVA)
Poloczanska, E., 134 analysis of risk and management of small populations, 173, 216,
Polycentric management and natural resource rights, 467 233, 240
Polygamous mating systems, 178 analytical-deterministic models, 243, 246, 270
Polymerase chain reaction (PCR), 179, 180 conceptual foundation and definition, 240–242
Pond biodiversity construction of, 216, 241, 242, 268
alpha, 48–52, 74, 326 definition, 240–242
gamma, 51, 52, 161, 326, 327 desired characteristics of, 233
Pools, 44, 50, 177, 184, 202, 224, 229, 287, 309, 322–328, 453, elasticity analysis, 247, 250
467–469, 560 evaluation of relative risk, 216
Pope Francis, 437–439 factor resolution, 233–236
Population northern spotted owl (Strix occidentalis caurina), 285, 358
age-structure, 103, 214, 215, 220, 242 persistence time of population, 220
age-structured growth model of, 103 spatially explicit population model, 102, 222
ceiling, 246 spatially realistic model, 220
conceptual framework for management decisions, 287, 288 stage-based model, 242, 243
critical age class, 219, 220 stochastic model, 241, 270
definition of, 211 surrogate population studies, 57, 68, 69
disjunctions associated with, 256 trend analysis, 233–236, 240
exponential growth, 213 western prairie fringed orchid (Platanthera praeclara), 242–246,
fitness, 46, 149, 171–173, 175, 177, 185–188, 203, 220 248 (see also Minimum viable population; Populations)
forest insects as examples of delayed density dependence, 218 Population viability management (PVM), 251–253, 361
fundamental equation of, 213 Porter, W., 414
gene flow, 75, 161, 186, 187, 189–191, 203–206, 212 Positive incentives and global cooperation with conservation laws and
genetic analyses of, 179, 182, 203–205 treaties, 520, 521
genetic diversity, 45, 75, 149, 161, 171–182, 184, 188–198, Possingham, H.P., 162, 400, 401, 515
200–202, 205, 206, 284, 419, 526 Pounds, A., 146
genetic management of, 195, 200, 201, 204, 206 Prairie
logistic growth, 102, 213, 214, 217–220 cool season grasses, 391
mainland-island, 222, 223 ecosystem management of, 94, 391
managed size, 75 effects of fire suppression, 278, 391
management of hybridization and introgression in, 102, 193, 195, potholes, 325
196 restoration of, 93, 94, 192, 239, 270
management of invasive, 102 tallgrass, 48, 49, 93, 94, 242, 391, 460, 535, 536
marginal, 172, 204 warm season grasses, 94
metapopulations and subdivisions, 142–144, 150, 212, 220–223, Prairie chicken (Tympanuchus cupido pinnatus), 192, 214
244, 271, 278, 312, 510 Prairie dog (Cynomys spp.), 236–238
monitoring, 182, 251, 253–255, 507 Prairie vole (Microtus ochrogaster), 277
mortality, compensatory, 214 Prakrti (five elements of), and Hinduism, 431
mortality, competing, 213, 215, 234, 235, 243, 412, 458 Prance, G., 58, 59
patchy, 222, 268, 294 Pre-adaptation, 100
processes in small, 39, 51–54, 62, 103, 171–173, 175, 178, 179, Predators and predation
182–184, 188, 189, 192, 193, 204, 205, 214–216, 220, 233, 240, biological control, 221
241, 256, 257, 292, 509 community biodiversity, 266
recovery strategies and threat identification for, 234, 236, 237, 247 compensation for losses to, 460
restoration of, 212, 233, 234, 255 edge and interior species, 277, 279
sex ratio, 177–179, 181, 201, 207, 214, 215, 242 edge, effect on, 277
source-sink, 222, 223, 247 habitat preferences of prey species, effects on, 344
spatially-structured, 102, 223 invasive species, 100, 101
threshold criteria for IUCN endangerment categories, 47, 171, 190, wolves, effects on livestock, 461
249, 335, 336, 374, 442, 515 (see also Minimum viable population viability analysis, 251–252
population; Population viability analysis; Small populations predator-prey interactions, 220, 334
captive breeding) predator-prey interactions, shifts under climate ,
Population demography, 35, 102, 173, 187, 211–214, 241, 256, 288 change,136, 138
Index 607
Prediction, algorithms and models Q

biodiversity indicators, 49 Qi, D., 265
extinction likelihood, as result of inbreeding, 187 Qing Dynasty, 6
fish community composition, 139, 330 Quantitative and measurable standards, 46, 50, 52, 174
fish population persistence, 311, 354 Quasi-extinction
gene flow, 187, 189, 190 contour, 241, 242
habitat change, 268–271 curve, 241, 242
lake transformation, due to eutrophication, 319 probability, 241
marine habitat complexity, 332, 333 surface, 241
minke whale (Balaenoptera acutorostrata), 205, 206 threshold, 241
models, in ecosystem management, 363 value, 247
park visitor days, 353 Quasi-option value, 419
percolation theory, 283–289 Quaternary Period, 82, 129
population persistence, 143, 222, 251, 284, 288, 289 Queensland (Australia), 328–331
population viability analysis, 173, 220, 233, 240–249, 251–252, 362
red wolf (Canis rufus) population persistence, 193, 195, 196
species range shifts associated with climate change, 139, 147, 565 R
stream invertebrate composition, 330 Raccoon (Procyon lotor), 277
Pre-mating isolating mechanism, 46 Radiative forcing, 117, 129, 131
Prescribed fire, 75, 278, 391, 404, 415, 416 Rainey Wildlife Sanctuary (Louisiana, USA), 471
PRESENCE occupancy and detection program, 230 Rajiv Gandhi National Park, 63
See also Detection, Occupancy probability, Occupancy theory and Ramahlo, C., 109
modeling RAMAS, 206
Preservationism, 12, 13 Ramsar Convention on Wetlands of International Importance, 326
Pricing and value of nature, 450 Ramsar Designated Wetlands
Primary environmental ethic, 421 Ngiri-Tumba-Maindombe (Democratic Republic of Congo), 326
See also Secondary environmental ethic Queen Maud Gulf (Canada), 326
Prioritization and conservation strategies, 70, 159 Randall, A., 418, 419
Private goods, and biotic resources, 419 Random mating and effective population size, 177, 178
Private property Rare alleles and bottlenecks in small populations, 173, 178, 179
conservation easements and zoning, 469–471 Rarity of species and biodiversity, 51–57, 68, 413
conservation strategy, 454 Ratnam, W., 206
property values and stream restoration, 457 (see also Property Ray, J., 45
rights) Reaction-diffusion models, 103
Production function, and ecosystem valuation, 373, 457 Realized niche, 152, 154
Professional networks, in conservation biology, 536–537 Reasonable and prudent alternative (RPA), 345
Professional relationships, in conservation biology, 536, 556–557 Reclamation and habitat disturbance, 300, 567
Professional societies, 3, 13, 412, 537–538, 563, 568 See also Mitigation of habitat disturbance
Project organization model, 361 Recolonization, 220, 222, 223, 226, 269, 316
Proper Baltic Sea, 348 Record of Decision (ROD), environment, 498
Property rights Recovery plans, and Endangered Species Act, 250, 252, 503
biodiversity, 467 Recruitment
Endangered Species Act, 506 adder (Vipera berus) restoration, 190
exclusive economic zones (EEZs), 524 corals, 343, 344
pollution and environmental, 468 desert bighorn (Ovis canadensis nelsoni), 144
role in conservation, 13 fire and grazing model in Ponderosa pine, 384
tradable, 468 limit indicator in ecosystem management, 373
wildlife refuges, 14, 61 (see also Private property) mean generation time and captive breeding, 201
Property right systems, best practices for common pool resources, 467, population demography, 212
468 Red-backed shrike (Lanius collurio), 266
Property values, and stream restoration, 457 Red Book (US Fish and Wildlife Service), 362
Proportional family richness, 69 Red-cockaded Woodpecker (Leuconotopicus borealis), 187, 506–508
Protected area-centered ecosystems (PACEs) Red Data Books Survival Service (IUCN), endangered species, 20
crucial habitats, 378 Red fox (Vulpes vulpes), 270, 277
Protected areas, 2, 22, 23, 39, 42, 59, 61–63, 70, 121, 141, 155, 160, Red List Criteria, 39, 53
161, 164, 289–296, 316, 319, 320, 349–355, 371, 377–379, 388, Red List (IUCN), 39, 40, 47, 53, 60, 68, 150, 190, 307, 336
398, 442, 444, 471–472, 479, 520 Reduced emissions from deforestation and forest degradation (REDD),
Provisional Act No. 2186-16 (Brazil), 524 480
Public comment, on environmental impact statement, 498 Red wolf (Canis rufus), 193
Public goods, and biotic resources, 419 Reed canary grass (Phalaris arundinacea), 311
Public-issue identification environmental impact assessment, 497 Reeves, R., 347, 348
Public participation, and conservation law Reference habitat and habitat classification framework, 104
Endangered Species Act, 75, 360 Reforming the Forest Service, 518
National Environmental Policy Act, 75, 360, 493 Regional biodiversity management, 159
Public policy, definition of, 463, 492 Regional habitat representation, 320, 322
Purple loosestrife (Lythrum salicaria), 311 coarse-filter model, 159, 161, 162
Pyrrocoma racemosa var. racemosa, 188 fine-filter model, 159, 161
608 Index
Regularly collected data, 382–384 Rissmen, A., 470

Regulatory science, conservation biology as a, 411–412, 489, 563 River herring (Alosa spp.), 316
Rehabilitation River Meuse, 50
coral reefs, 340–343 RNA, 180, 181
environmental insurance bonding, 496 Roberts, J., 519
orangutans, 443, 444 Robertson, D., 359, 363
water conservation practices and Islamic conservation , Robinson, J.B., 564
principles,440 Rockström, J., 117, 118
Relatedness, inbreeding depression and degree of, 184, 186 Rodgers, W.H. Jr., 493
Relational indicators of wellbeing, in ecosystem valuation, 477 Rohlf, D., 29, 500, 509, 510, 565
Relational skills, and careers in conservation biology, 547–548 Rojas, M., 47
Relational values, 477 Rolston, H. III., 416, 418, 421, 422, 425, 430
Religious traditions, and understanding of intrinsic value Romantic Transcendentalism, 12
in conservation biology, 428–436 Rondinini, C., 292, 293, 386
Remote sensing Roosevelt, T., 11–12, 14, 18, 32, 61, 174, 495
employed in species distribution modeling (SDM), 66 Rosy finch (Leucosticte tephrocotis), 125
Global Ecosystem Dynamic Investigation (GEDI) LiDAR, 165 Roszak, T., 426
Landsat Thematic Mapper (LSM), 67 Royal Societies in the Commonwealth of Nations, 491
Removal experiments Rubber, 23, 24, 220, 221
large-flowered fiddleneck (Amsinckia grandiflora), 234 See also Chico Mendez
snow and two-lobe larkspur ((Delphinium nuttallianum), 152 Rule-based models, 4, 218
swamp wallaby (Wallabia bicolor), and ecosystem management, Rural-to-urban gradient (RUG), 108, 109, 111
374–376 Ruskin, J., 8
Replacement cost, in microeconomics, 460 Rypstra, A., 278
Representative Areas Program (RAP), 402, 403
Reproductive ecology, 203, 288
Reproductive isolation, species criteria, 46 S
Research Experiences for Undergraduates (REU), 534 Sabbath rest, in Judeo Christian Stewardship Environmental Ethic, 430
Research-Implementation Gap, 543, 546 Saccheri, I., 187
Reserve selection algorithms and criteria, 289–291 Safe harbor agreements and Endangered Species Act, 507
Reserve size, 292–295 Safe operating space for humanity, 117, 118
Reserve size, determination of, 292–295 Sagebrush (Artemisia spp.), 383, 388, 404–407, 414–416
Resource Conservation Ethic, 15–17 Sage grouse (Centrocercus urophasianus), 415
Resource management Salmon, 4, 17, 18, 138, 139, 160, 406
Aldo Leopold, 26 See also Atlantic salmon; Chum salmon; Chinook salmon; Coho
applied science, 26 salmon; Sockeye salmon Sampling units, and ecosystem
contrast to conservation biology, 25–28 management
contrast to ecosystem management, 353 Salt sacaton (Sporobulus airoides), 94
maximum sustainable yield, 17, 21 Salzman, J., 491, 492, 522–526
new expressions of, 21–22 San Bruno Mountain controversy (California, USA), 507
primitive cultures, 3 San Francisco Bay, 457
research coordination, 373–374 See also California
Resource polymorphisms, 203 Sandberg bluegrass (Poa secunda), 234
Restoration Sardelis, S., 560
adder, Common European (Vipera berusii), 190 Sariska Tiger Reserve (India), 435
gravel pit, 93 Sarkar, S., 22, 61, 62, 412, 413, 421
prairie, 93, 192 Satellite images, 269, 291, 386
prairie chickens, 192 Satellite markers and DNA fingerprinting, 180, 181, 204
Restricted distribution of species, 54–57 Savannah River Ecology Laboratory, 535
Revealed preference methods, and microeconomics, 457 Save the Orangutan (STO), 443
Reverse Transcription PCR (RT-PCR), 180 Schlager, E., 467
Reynolds, R., 362 Schlich, W., 21
Rhino horn, trade of, and CITES, 514 Science (journal), 2
Rhinoceros Science, and values, 431
black (Diceros bicornis), 61, 514 Scientific baselines and ecosystem management, 387, 388
white (Ceratotherium simum), 514, 515 Scientific Panel on Late-Successional Forest Ecosystems
Richter, B.D., 75 (“The Panel”), 363
Ricker models and fisheries management, 343 Scoping meetings, and National Environmental Policy Act, 497
Riffles, 309 Scott, D., 125, 126, 155
Rights of natural objects, 427 Scott, J.M., 75, 253, 291, 342, 541, 552, 566
Ringed seal (Pusa hispida), 147 Seagrass
Rio Declaration, 510, 516, 517, 519, 521 Chesapeake Bay, 336
Rio Summit, see Earth summit conservation, 335–337
Riparian habitat and zones, 313 distribution, 337
Riplox method, 318 ecosystem services, value of, 335
Risk analysis, 216, 240, 368, 370 eelgrass (Zostera marina), 336
wildebeest (Connochaetes spp.) model, 480 food source for marine animals, 335
Rissler, L., 59, 60 restoration, 336, 341–343
Index 609
Sea ice, loss of, 147, 148 genetic drift and loss of alleles, 175–177
Seattle Audubon Society, 362, 363 genetic restoration of, 188, 190
Sea Turtle Act, 36, 523, 526 hybridization and introgression, threats from, 193, 195, 196
Sea turtles inbreeding and inbreeding depression, effects on, 197
biodiversity, 351, 522, 526 invasive species, effects on, 95
coral reefs, 144, 145 IUCN criteria for endangerment, 47
international conservation issues, 522 management of, 224–231
molecular genetic studies, 203 population processes, effects of, 144, 154, 201, 211–257
overexploitation of, 347 population viability analysis of, 173, 216, 220, 233, 240–252, 268,
reproductive ecology, 203 292, 362, 564
Seaweed and ecosystem modeling, 388–391 risk analysis of, 216
Secondary environmental ethic, 421 trend analysis, 233–236 (see also Populations)
See also Primary environmental ethic Smith, J.A., 508, 509
Secretary of the Interior, and Endangered Species Act, 501 Snail darter (Percina tanasi), 501
Sediment cores, from lakes and bogs, 82 Snow goose (Chen caerulescens), 27
Sedimentation, and aquatic habitats, 317 Social dimension of conservation assessments, 546, 547
Selection index (SI), 263 Social process, conservation as, 537–538
Selection strategy and captive breeding, 197, 201 Society for Conservation Biology (SCB)
Self-pollination (selfing), 188 Equity, Inclusion, and Diversity Committee, 560
Sensitivity to, climate change, 145, 149 Social Science Working Group, 549
Sentient creatures, 423 Society for Range Management (SRM), 535
Serengeti, 391, 395, 480 Sociological surveys, and values, 419–420
See also Serengeti-Mara Sockeye salmon (Oncorhynchus nerka), 4
Sessions, G., 426, 427, 432 Socrates, 4
Seto, K.C., 106 Soil conservation, and watershed-based management, 381
Severns, P.M., 188 Solarz, S.L., 547
Sex ratio Song sparrow (Melospiza melodia), 183, 187, 228
demographic stochasticity, 215 Soulé, M.E., 1–3, 28, 29, 31, 32, 35, 171–173, 175, 179, 185, 242, 292,
effective population size, 177–178 411, 537
logistic population growth, 214 South Africa
Shaffer, M.L., 214, 240 Cape Floristic Region, 53, 158
Shannon Index, 49, 174, 202, 206 Fynbos region, 54 (see also Africa)
Sheep South Dakota, 54, 236, 237, 239, 435
domestic, interaction with desert bighorn, 144 South East Queensland Healthy Waterways Partnership (SEQHWP)
Shepherd’s Club, 468 Ecosystem Health Monitoring Programme (EHMP), 329
Sheil, D., 565 Environmental Management Support System (EMSS), 329
Shellenberger, M., 436 Percentage of Native Species Expected (PONSE), 330
Sheppard, J.K., 32 Plecoptera, Ephemeroptera and Trichoptera (PET) family richness,
Sheyenne National Grassland, North Dakota (USA), 246 146, 330
Short tandem repeats (STRs), 181 Stream Invertebrate Grade Number Average Level (SIGNAL) index,
Shrimp fishing, and protection of sea turtles, 523 330
Sieg, C.H., 242, 246 South East Queensland Regional Water Quality Management Strategy
Sierra Club, 12, 13, 425, 426 (SEQRWQMS), 329
Sierra Club v. Morton, 425, 426 Southern Ocean, 337, 346, 347, 349
Sierra Nevada Mountains and John Muir, 11, 14 Southern red-backed vole (Myodes gapperi), 298
Significant environmental effects, and environmental impact Southern white rhinos (Ceratotherium simum), 514
assessment, 12 Southwest Atlantic Section (of Southern Ocean), 346
Silva, M., 23, 24 See also Antarctic Peninsula Plume (APP)
Simpson Index, 174 Sovereign rights over resources, and international conservation, 521
Singer, P., 433 Spain, 59, 247, 312, 472
Single species management, 160 Spatial heterogeneity, 102, 219, 266
Six-spotted mite (Eotetranychus sexmaculatus), 220, 221 Spatially explicit, spatially implicit, and spatially realistic models of
Skellam, J.G., 102 metapopulation biology, 222
Slash-and-burn agriculture, 197 Spatial Planning for Protected Areas in Response to Climate Change
Sleepy lizard (Tiliqua rugosa), 287 (SPARC), 161
Small population paradigm Spatial scales, 71, 74, 75, 86, 160, 262, 264–268, 270, 272, 288, 300,
conceptual framework for management, 288 368, 369, 380, 383, 399, 482
effective population size, 75, 175 Species
species management, 363 (see also Small populations) biodiversity definition and concepts of, 20, 21, 26, 28–30, 35–42,
Small populations 44–57, 59–62, 64–66, 68, 69, 71, 74–76, 83, 84, 93, 95, 420
Allee effect, 103 distribution, 4, 47, 55, 60, 66, 125, 126, 138, 149, 152–154, 291,
basic rule of conservation genetics, 172 310, 386, 509
bottlenecks and rare alleles, 178, 179 Endangered Species Act and definition of, 27, 36, 75, 113, 193, 203,
characteristics of rarity associated with, 253 206, 236, 250, 252, 255, 302, 345, 360, 364, 416, 463, 489, 493,
Endangered Species Act, 252 494, 500–510, 521, 522, 524
factor resolution, 233–236 extinctions, 40, 83, 106, 126, 158, 429, 490
610 Index
Species (cont.) Streams

international conservation of commercially valuable, 18, 492, 509, abiotic conditions and invasive species, 101
513–515, 522, 526 access and property rights, 450, 468
loss, due to climate change, 27, 32, 41, 42, 55, 57, 61, 66, 85, 88, 90, aquatic ecological systems (AES), 320, 322
94, 95, 97, 111, 117, 125, 126, 134, 136–139, 141, 142, 145, as derived ecosystems, 309
146, 149–166, 220, 266, 270, 310, 316, 335, 360, 367, 393, 402, ecosystem management, 59, 97, 309, 313, 316, 319, 320, 322, 329,
437, 565 330, 361, 392, 430
range shifts, 134, 136, 139, 142, 154 Iberian Peninsula, conservation value of species in, 247, 312
types/evolutionary units, 47 invertebrate communities and biodiversity of, 330
worldwide estimates of total number, 98, 158 (see also Species network, 320, 322
richness conservation of) order, 309
Species-area relationship (SAR), 324, 327 prioritization for conservation, 545
Species evenness, 37, 48, 49 reaches, 11, 50, 51, 129, 215, 283, 284, 310, 373, 466, 473
Species list, 49, 250, 308, 336, 501–503, 505, 509 restoration of in hedonic property model, 457
Species management, 363 riparian vegetation, 330
Species response to climate change riparian zones, 320
interactions across trophic levels, 136–139 threats to conservation of, 316
observed range shifts, 139 urbanization effects, 325
phenological changes, 136–139 Streiner, C.F., 457
trends in local abundance, 139–142 Stuart, S., 39, 41, 430
Species richness Student Conservation Association, 534
alpha diversity, as measure of, 48–49 Subcommission on Quaternary Stratigraphy, 81, 82
coral reefs, 371 Subpopulations and metapopulation biology, 149, 222
correlations among taxonomic groups, 68 Succession
correlation to habitat heterogeneity and patchiness, 266 ecological, 50, 270, 271
endemism, relationship to, 55, 56, 308 management of, 271
hotspots, as centers of, 58 Sulawesi Tonkean macaque (Macaca tonkeana), 53
hybrid zones, 193 Sumatran orangutan (Pongo abelii), 442
indicators of, 69, 292 Sumatran Orangutan Conservation Programme (SOCP), 443
latitude, relationship to, 57 Sunshine approach, to global cooperation with conservation laws and
patch size, relationship to, 268 treaties, 519
wetland, 484 Supply and demand and microeconomics, 454–456
Species specific conservation, 9, 68 Supreme Being and Hinduism, 432
Species specific responses to climate change, 160 Surplus-yield model, 346, 347
Species Survival Plan (SSP), 197 Surveys and research on attitudes and values, 419–420
Spotted owl (Strix occidentalis) Sustainability and ecosystem management paradigm, 112
ecosystem management, 362–364 Sustainable development, 24, 30, 485, 500, 510–513, 516
Endangered Species Act, 364 Sustainable Development Goals (SDG), 517
Mexican (Strix occidentalis lucida), 289, 290 Sustainable use, 14, 96, 355, 452, 494, 511, 517, 524
northern (Strix occidentalis caurina), 361–364 Sustained yield forestry, 15
percolation theory, 289 Swaisgood, R., 32
Squinting bush brown butterfly (Bicyclus anynana), 186 Swamp wallaby (Wallabia bicolor), 374–376, 389
Stage-based deterministic models, 243 Sweden, 21, 117, 190, 512
Stage-based (Lefkovitch) model, 243 Switchgrass (Panicum virgatum), 391
Stakeholders Sylvatic plague (Yersinia pestis), 236, 237
knowledge-deficient model, 404 Sympatric populations, 203
Starfield, A.M., 88, 90 Sympatric species, 203
Stated preference methods and ecosystem valuation, 460–461 Systematic conservation planning (SCP)
Statement on Forest Principles, 518 ecological uniqueness, 70
Steelhead trout (Oncorhychus mykiss), 196 informed opportunism, 70
Stegner, W.E., 10 opportunism, 70
Steller sea lion (Eumetopias jubatus), 345 relative threat of loss, 70
Stochastic models, 241, 246, 268, 270, 289
Stochastic population viability analysis, 241
Stochastic variation T
demographic, 214, 215, 240, 246, 256, 345 Taking, of endangered species, 494, 506
environmental, 214, 215, 220, 240, 246, 256, 289, 345 Tanzania, 61, 395, 480
genetic, 214 Tapanuli orangutan (Pongo tapanuliensis), 442
Stock-flow resources, 450–452 Taq polymerase, 179
Stockholm Conference, and international conservation law, 512 Tattoni, C., 266, 267
Stoll, M.R., 428, 430 Tawhid (unity), and Islam, 431
Stone, C., 425, 427, 430 Taxation, 460, 463, 464
Stoppani, A., 83 Taxation and environmental regulation, 144, 456, 526
Stratified diffusion models, 103 Taxon-based indicator species, 69
Stratigraphic markers, 82, 83 See also Taxon surrogates
Index 611
Taxonomic assessment, 203 Tribe, L., 412, 413, 435

Tejadhammo, P., 441 Trillium (Trillium spp.), 421
Tellico Dam controversy (Tennessee, USA), and Endangered Species Tropholytic zone, 309
Act, 501 Tropic of Capricorn, 351
Temperate regions and biodiversity, 59 Tropical Conservation and Development (TCD) Program, and The
Temperature University of Florida, 543, 545
air, 136, 137, 275 Tropholytic zone, 309
animal distribution and extinction, 66, 126, 128, 136, 138, 145, 149, Trust
153, 266, 385 components of, 404
Arctic, 88, 135, 147 establishment with stakeholders, 403–406, 408
climate change, 32, 82, 88, 92, 129, 139–141, 144, 145, 319, 393 importance and role of in ecosystem-based management, 405–407
coral bleaching, 145 Tubeworms, 307, 332
diagnostic fingerprint, 135 Tull, J., 564, 565
geological cycles of, 127 Tully-Murray floodplain, 398
ice cores, estimation of, 129–131 Tully-Murray Water Quality Improvement Plan (WQIP), 396
ice extent, 135 Tully-Murray Watershed (Catchment), 396, 397, 402
ice thickness, 147 Tully River, 396
invasive species, effects on, 85, 90, 111, 125, 141 Tuna, and Marine Mammal Protection Act, 522, 523
keystone species, effects on, 141 Turchin, P., 218
ocean, 145, 340 Turkey (Meleagris gallopavo), 196
polar, 54, 57, 133, 147, 148 Turtle excluder devices (TED), 349, 523
rate of increase, 130 Turtles, see Sea turtles
timing events, 134 (see also Climate) Typological species concept, 45
Temple, S.A., 2, 277 Tzu Chi, 441
Temporal scales, 267, 268, 270, 272, 367–369, 377, 427
Tennessee Valley Authority, 464, 501
Tennessee Valley Authority v Hill, 501 U
Texas (US), xi, 102, 113, 173, 174, 189, 190, 192, 216, 522, 536 Umbrella species, 59
Texas State Bison Herd (TSBH), 173, 174 Underwood, B., 414
Thailand, 433, 440, 441 Ungulates (hooved mammals), 42, 382, 391
The Nature Conservancy (TNC), 75, 109, 115, 117, 119, 289, 302, 320, United Kingdom, 6, 8, 52, 134, 190
321, 341, 361, 385, 469–471, 534, 550, 565 United Nations
Theodore Roosevelt National Park, 174 Charter of the, 510
Thermal expansion, of warming oceans, 133 Conference on Environment and Development, 515
Thermus aquaticus, 179 Conference on the Global Environment, 512
Thistle, 8 Conference on the Human Environment, 512
Thomas, C., 158 Convention on the International Trade in Endangered Species
Thompson, B., 491, 492, 522, 523, 525 (CITES), 206, 489
Thoreau, H.D., 12, 13, 432 Convention to Combat Desertification (UNCCD), 511, 518
Threatened species and Endangered Species Act, 76, 500, 501, 503, 505 Declaration on the Rights of Indigenous Peoples, 524
Threats, as criteria for protected area selection, 30, 71, 106, 125, 240, Development Programme (UNDP), 351, 353, 512, 519
241, 252, 264, 309–312, 332, 335, 337, 339, 340, 347, 351, 521 Educational, Scientific and Cultural Organization (UNESCO), 19,
Timber Culture Act, 13 20, 24, 350, 513, 521
Timber industry, 15, 21, 23, 283, 362, 375 Environmental Programme (UNEP), 19, 24, 441, 442, 510, 512,
Time-control substitutes, and undisturbed ecosystems, 388 513, 518, 519
Timing limitations, and mitigation, 300 Food and Agriculture Organization (FAO), 518, 524
Tingley, M., 159–161, 164 Framework Convention on Climate Change (UNFCCC), 164, 510,
Tlingit Tribe, 4 517, 518
Torres Strait, 351 General Assembly, 510, 518
Total allowable catch levels (TACs), 344 Sustainable Development Goals Third Conference on the Law of the
Tourist-recreation marine reserves, 334, 367 Sea, 513
Tradable property rights, 465–466, 468 Universities and conservation biology as academic discipline, 29, 411
Tragedy of the commons, 468 See also Conservation Education
Transcendentalism, and environmental values, 12 University of British Columbia’s (Canada) Institute for Resources,
Transects, 225, 246, 382, 383 Environment and Sustainability, 549
Transition matrix, 244, 245, 269 University of California (USA), 221, 341, 552
Travel cost method (TCM), and microeconomics, 458, 459 University of Florida (USA), 543, 545
Treaties, see Global issues, and international cooperation with University of Maryland, College Park (USA), 547, 548, 557
conservation law and treaties University of New South Wales Canberra (Australia), 560
Treaty Concerning the Regulation of Salmon Fishery in the Rhine River University of Oklahoma (USA), 536
Basin, 18 University of Oklahoma Botanical Society, 536
Tree frog University of Sustainable Development Eberswalde, 73
Copes Gray (Hyla chrysoscelis), 46 University of Wisconsin at Madison (USA), 16
Gray (Hyla versicolor), 46 Urban adapters, 111
Trend analysis and population restoration, 233–236, 240, 256 Urban avoiders, 111
Treves, A., 461 Urban biodiversity, 64, 65, 110, 111, 113–114, 117
612 Index
Urban conservation, 112–116 Values

Urban ecological research, 107 alternative measurements of, and ecological economics, 474, 475
Urban ecological systems, 107, 108, 110, 112, 117, 122 bequest, 419
See also Urban ecology, urban ecosystems biodiversity, 31, 62, 120
Urban exploiters, 111 categories of, 411, 417–421, 477
Urbanization, effects on stream communities, 111, 325 decay (K), 274, 275
US Agency for International Development (USAID), 479, 481 definition of, 415
US Army Corps of Engineers, 326 demand, 421
US Bureau of Land Management, 239, 362, 404, 496, 534 ecological, 28, 63, 427, 466, 477
US Catholic Conference, 437 economic valuation methods, 420–421
US Congress, 13, 60, 344 ecosystem management, 359, 373
US Court of Appeals, 495 ecosystem management paradigm, 360
US Court of International Trade, 206, 489, 513, 523 existence, 367, 419, 460
US Department of Agriculture (USDA), 14, 15, 117, 496 experiential, 421
US Department of Commerce, 507 hedonic, 457–458
US Department of the Interior, 14, 425 ignition, 422
Use/availability data, and preferred habitats, 256, 362 instrumental, 76, 418–419, 444, 477
US Environmental Protection Agency (USEPA/EPA), xi, 131, 455, 456, intrinsic, 3, 5, 12, 13, 18, 28, 29, 32, 239, 411, 413, 418, 421–429,
463 444, 450, 482, 493, 509, 510
User fees, 459, 460 moral (see Intrinsic)
User satisfaction and values, 420 option, 419, 467
Use values, 367, 419 outcomes as expressions of, 416
US Fish and Wildlife Service (USFWS) pluralism, in ecosystem valuation, 476
black-footed ferret, 424, 546 quasi-option, 419
California condor, 424 science and, 412, 431
ecosystem management, 363, 494, 509 sociocultural, 477
Endangered Species Act, 193, 494, 500, 501 utilitarian, 3, 28, 29, 411, 417, 419 (see also Ethics)
government-market coordination, 466–467 Van Dyke, F., 28, 31, 49, 58, 93, 225, 229–231, 245, 262, 264, 269, 287,
history of conservation, 3 300, 391, 415, 416, 442, 444, 459, 567
Kirtland’s warbler, 190, 424 Van Houtan, K.S., 32
US Forest Service Vaquita porpoise (Phocoena sinus), 348
biodiversity management, 17, 289 Variable number tandem repeats (VNTRs), 181
ecosystem management, 298, 389, 427 Variegated habitat and landscape, 271
habitat management on non-reserve lands, 17, 254, 284, 289, 299 Vaughn, D., 341
history of conservation, 14, 17, 254, 255, 299, 420, 427 Veery (Catharus fuscescens), 287
Interagency Scientific Committee (ISC), 363 (see also Northern Vegetation Management Act (VMA), 398
spotted owl (Strix occidentalis caurina)) Vernal pools, 325, 326
invasive species, 103 Vernal Pool Working Group (VPWG), 326
multiple use management, 14, 16 Viability, as criteria for protected area selection, 289
National Environmental Policy Act (NEPA), 360 Victoria, Australia, 223, 375
Smokey Bear, 420 Victorian Department of Sustainability and Environment (DSE), 375
spotted owl, 289, 361–364 Vidya, T.N.C., 204
US Geological Survey (USGS), 11, 46, 66, 96, 138, 148, 229, 230, 236, Vienna Convention of the Law of Treaties, 510
240, 375, 415 Villaseñor, J.G., 69
US Homestead Act of 1862, 12 Virtues and conservation ethics, 433
US House Committee on Natural Resources, 499, 507 Visual minimization, and mitigation, 300
US National Academy of Sciences, 132, 343 Vitousek, P.M., 91, 96, 104, 126
US National Aeronautics and Space Administration (NASA), 66, 328, Vocation, 113, 429, 531–569
386, 457 Vocational experience, in conservation biology, 534–535
US National Science Foundation (NSF), 107, 534, 543 Vocational settings and career in conservation biology, 551–563
US Northeast Shelf Ecosystem, 343, 345 Voluntary programs, of environmental regulation, 463
US Pacific Northwest, 283, 288, 362–364, 393, 406 Von Ribbentrop, B., 21
US Pacific Rivers Council, 406 VORTEX and population viability analysis, 206
US Supreme Court, 425, 426, 501
US Sustainable Fisheries Act, 344
US Wilderness Act of 1964, 17 W
Utah (USA), 290, 385, 495 Walker, S., 85, 278
Utilitarianism, 421 Waller, D.M., 183, 187, 188
Utilitarian values, 3, 28, 29, 417, 419 Walton, J.H., 429
Utility and neoclassical economics, 454, 476 Washburn expedition, 9–11, 16
Washburn, H., 10
Washington (USA), 138, 322, 324, 361, 363, 440, 513, 526, 536, 547,
V 548, 566
Value judgements, 412–418, 564 Washington DC – Baltimore Corridor (US), 106
Value rational approaches, in conservation, 436, 437 Washington Department of Fish and Wildlife, 566
Value-laden nature, of conservation biology, 28, 45, 411–412 Water conservation areas (WCAs), 377
Index 613
Water flow, and ecosystem management, 392, 393 Willow (Salix spp.), 323, 394
Water regimes, and edge environments, 276 Wilson, E.O., 44, 58, 117–119, 222, 295, 422
Waters, C., 82, 84 Wind, as a factor in edge environments, 273–274
Watershed-based ecosystem management, 381 Wisconsin (USA)
Watershed coalitions, councils and partnerships, 408 ecosystem management, 379
Wavelengths, of light, 308 wolf restoration, 212
Weather, 94, 127 With, K.A., 103, 282, 283, 290
Weiss, E.B., 519–521 Wolf spider
Western prairie fringed orchid (Platanthera praeclara), 242–246, 248 Pardosa milvina, 279–281
Wetland banking, 465, 466 Tigrosa helluo (formerly Hogna helluo), 279
See also Tradable permits Wolverine (Gulo gulo), 502
Wetlands Women conference organizers, and professional societies, 560
Aammiq wetland, 437, 438 Wood Buffalo National Park, 173
biodiversity, contribution to, 59, 326 Wood of the Hague, 9
coastal, 396 Wordsworth, W., 7
constructed, 242, 317 World Bank
definition, 322 Global Environmental Facility, 351, 519
ecosystem management, 379 World Commission on Protected Areas (WCPA), 61
Everglades, 11, 375 World Conservation Congress, 39
functions and services, 323 World Heritage Centre, 521
invasive species, 311 World Heritage List (WHL), 521
management, 324, 325 World Heritage Sites, 197, 351, 511
Ramsar Convention, 326, 511 World Resources Institute, 450
restoration, and vocational experience, 379 World Trade Organization (WTO), 522–524, 526
soils, periodically saturated, 323 World Vision, 437
soils, permanently saturated, 323 World Wildlife Fund (WWF), 72, 349, 361, 420, 440–442
species, 242, 325 Wright, S., 175
Weyerhaeuser Company v. United States Fish and Wildlife Service, 505 Wunder, S., 482–484
Whale feces, role in ecosystem production, 346 Wyoming (USA), 10, 51, 141, 237, 299, 386, 496, 504
White, G., 7, 8, 32
White-footed mouse (Peromyscus leucopus), 186, 269
White-headed vulture (Trigonoceps occipitalis), 230–232 X
White House Council of Environmental Quality (CEQ), 494, 497, 499 Xapuri Rural Workers’ Union, 23
White, L. Jr., 429
White rhino (Ceratotherium simum), 514, 515
White-tailed deer (Odocoileus virginianus), 271, 471 Y
Whittaker, R.H., 49, 50 Yandle, B., 468
Whittaker’s Measure, 49 Yangtze dolphin (Lipotes vexillifer), 348
Whitten, T., 441 See also Beiji
Whole catchment management (WCM), 319, 320 Yellow-bellied marmots (Marmota flaviventris), 136
Widespread distribution, of species, 54–57 Yellowstone Ecosystem, 10
Wildebeest (Connochaetes spp.), 480 Yellowstone model, 10, 17, 21–25
Wilderness Act, 17 Yellowstone National Park (YNP), 10, 11, 22, 51, 116, 141, 173, 174,
Wilderness Ideal, 16–17, 21, 22 218, 219, 382, 383, 386, 388, 428, 461
Wilderness lodge, Walt Disney World, 414 Yellowstone River, 10, 466
Wilderness Society, 16, 17 Yield-based management, 15
Wildlands, 65, 83, 85, 90, 98, 107, 116, 117, 121, 122 Yisatii and Tlingit and Haida Tribes, 4
Wildlife ecology, 26–28, 552 Yohe, G., 134, 135
Wildlife food supplementation, 65 Yosemite National Park, 22
Wildlife management, 16, 25–27, 35, 224 Yosemite Valley, 14, 22
Wildlife Management Institute (WMI), 195
Wildlife Preservation Society (WPS) (Australia), 351
Wildlife sanctuaries, 14, 471–472 Z
See also Nature reserves Zebra mussel (Dreissena polymorpha), 96, 100
Wildlife Society, The Zheng, D., 274, 275
Annual Conference, 536 Zilly, J.T., 363
Wild rice (Zizania spp.), 324 Zimbabwe, 61
Wilhere, G.F., 491, 566, 567 Zonation management, 309
William of Normandy, 6 Zoning, 65, 116, 117, 314, 402, 403, 469–471, 561
William of Orange, 9 ZooRisk, 206
Willingness to accept compensation (WTA), 460, 461 Zooxanthellae, 145, 334, 335, 340
Willingness to pay (WTP), 459–461

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Fred Van Dyke · Rachel L.

ISBN 978-3-030-39532-2 ISBN 978-3-030-39534-6 (eBook)

# Springer Nature Switzerland AG 2008, 2020

Mancelona, MI, USA Fred Van Dyke

Writing acknowledgments is a necessary pleasure for authors, an opportunity to say the

Third Edition Reviewers

Katie DeVoss, Editorial and Research Assistant

Charles Acosta, Northern Kentucky University

Second Edition Reviewers

Michael J Bigelow, Illustrator and Permissions Assistant

Charles Acosta, Northern Kentucky University

Colleen Lynch, University of South Dakota

First Edition Reviewers

Krista Peppers, Editorial Consultant

Vickie L. Backus, Middlebury College

1 The History and Distinctions of Conservation Biology . . . . . . . . . . . . . . . . . . 1

2 Biodiversity: Concept, Measurement, and Management . . . . . . . . . . . . . . . . . 35

3.2 Understanding and Managing the Novel Ecosystem . . . . . . . . . . . . . . . . . 88

4.3.3 Dynamic Global Vegetation Models: Process-Based Approaches to

5.9.4 Genetic Analysis Can Determine Rates of Gene Flow Among

7 The Conservation of Terrestrial Habitat and Landscape . . . . . . . . . . . . . . . . 261

8.1.7 Dams as Barriers to Population Persistence and Reproduction . . . . 316

9.2.3 Evaluating Ecosystem-Based Management as a

10.5.6 Conservation Teachings in Buddhism . . . . . . . . . . . . . . . . . . . . . 432

12.3.3 Protection of Endangered Species: The Convention

13.6.5Making Advocacy Intentional – Avoiding Inadvertent

Keywords 65 million years, it was time for academics and conserva-

# Springer Nature Switzerland AG 2020 1

persons and groups, the worst biological disaster in the last

effective combinations of ﬁre and cultivation could create

Before Plato, Judaism extended the principle of Sabbath,

1.2.5 Conservation as Knowledge – The

Although such an idealized view of nature did not stimu-

Information Box 1.1 (continued) Information Box 1.1 (continued)

The formation of the APSB marked the beginning of the

Fig. 1.5 Castle Geyser, Upper

Northern Paciﬁc Railroad, the falls of the Yellowstone and the

extraction of its resources as commodities for human use and

Fig. 1.7 Walden Pond, a 26 ha

Fig. 1.9 Gifford Pinchot, American forest scientist and administrator

use of nature. They believed that all interests in resource use,

the present generation, but of the generations to come. Such

1.5.2 Forums for International Conservation –

Although nations increasingly began entering into bilateral

non-indigenous Brazilians to move to and occupy the Ama-

representing indigenous people are now increasingly present

Fig. 1.15 The endangered red-crowned crane (Grus japonensis),

the function of conservation biology as a professional scien-

Fig. 1.17 Three species, snow

Table 1.2 Seven core disciplinary distinctions of conservation biology

Conservation 153: 25–31. https://doi.org/10.1016/j.biocon.2012.

Keywords of natural habitats to systems designed to maximize human

# Springer Nature Switzerland AG 2020 35

Fig. 2.2 The Black-footed Ferret

Wild) (Fig. 2.5). Twenty percent (one in ﬁve) vertebrate

has yet been signiﬁcantly affected by conservation initiative.

Many articles and textbooks on conservation biology provide

Fig. 2.6 Relationship between

Fig. 2.8 (a) The Cope’s gray tree

Table 2.2 Eight conceptual definitions of species