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Essentials of Conservation Biology
Fifth Edition
Sinauer Associates, Inc., Publishers
Sunderland, Massachusetts U.S.A.
About the Cover
Brown pelicans (Pelecanus occidentalis) represent a conservation success story.
Pelican numbers had declined in the United States, with the pesticide DDT
identified as the cause of thinning eggshells and a lack of reproduction. Since its
listing under the U.S. Endangered Species Act in 1970 and the banning of DDT,
brown pelican numbers have increased substantially. Because of this recovery,
the brown pelican was removed from listing under the Act in 2009. (Photograph
© Tom Vezo/Minden Pictures.)
Printed in China
5 4 3 2 1
To my family, Margaret, Dan, Will, and Jasper,
and the teachers who inspired me,
Carroll E. Wood Jr. {1921-2009} and Janis Antonovics
Brief Contents
PART I Major Issues That Define the Discipline 1
1 What Is Conservation Biology? 3
2 What Is Biological Diversity? 23
3 Where Is the World's Biological Diversity Found? 51
Habitat Destruction 177 BOX 9.1 Pesticides and Raptors: Sentinel Species
Warn of Danger 198
Threatened Rain Forests 180
Water Pollution 198
Other Threatened Habitats 184
Air Pollution 201
Marine Coastal Areas 185
Desertification 187 Global Climate Change 204
, -
Changes in Temperate and Tropical Climates 208
Habitat Fragmentation 189
Plants and Climate Change 209
Edge Effects 193
Rising Sea Levels and Warmer Waters 209
Two Studies of Habitat Fragmentation 195
The Overall Effect of Global Warming 211
Habitat Degradation and Pollution 196
Minimizing Edge and Fragmentation Effects 374 Landscape Ecology and Park Design 382
Networks of Protected Areas 375 Conclusion 386
Habitat Corridors 375
Contents xi
Appendix 539
Glossary 545
Bibliography 553
Index 587
Preface
After decades of public interest in nature and the environment, the United Nations
focused worldwide attention on conservation by declaring 2010 to be the Interna-
tional Year of Biodiversity. The general public has absorbed this message and is ask-
ing its political leaders to provide the policy changes needed to address this issue.
Conservation biology is the field that seeks to study and protect the living world
and its biological diversity (or biodiversity in its shortened form). The field emerged
during the last 35 years as a major new discipline to address the alarming loss of
biological diversity. The threats to biodiversity are all too real, as demonstrated by
the recent recognition. that fully one-third of amphibian species are in danger of ex-
tinction. At the same time, our need to remain hopeful is highlighted, for exam-
ple, by increasing sea turtle populations at many locations throughout the world
following comprehensive conservation efforts. Many examples described in this
book show that governments, individuals, and conservation organizations can work
together to make the world a better place for nature.
Evidence of the explosive increase of interest in conservation biology is shown
by the rapidly increasing membership in the Society for Conservation Biology, the
great intellectual excitement displayed in many journals and newsletters, and the
large numbers of new edited books and advanced texts that appear almost week-
ly. International conservation organizations have emerged to tackle conservation
issues with a multi-disciplinary approach, and an Encyclopedia of Life is being de-
veloped as an online resource to provide the needed information for conservation
issues.
University students continue to enroll enthusiastically and in large numbers in
conservation biology courses. Previous editions of Essentials of Conservation Biolo-
gy have provided a comprehensive textbook for this subject. (The Primer of Conser-
vation Biology, in its Fourth Edition, continues to fill the need for a "quick" guide
for those who want a basic familiarity with conservation biology.) The Fifth Edition
of Essentials provides a thorough introduction to the major concepts and problems
of the field. Like its predecessors, it is designed for use in conservation biology
courses, and also as a supplemental text for general biology, ecology, wildlife biol-
ogy, and environmental policy courses. The book is also intended to serve as a de-
tailed guide for professionals who require a comprehensive background in the sub-
ject. Readers should enjoy and benefit from the updated full-color illustration and
photo program. Highlighted synopses of major points in the text have been added
as sidebars and serve as useful study aids.
This Fifth Edition reflects the excitement and new developments in the field. It
provides coverage of the latest information available on a number of topics, includ-
ing the expanding system of marine protected areas and linkages between conser-
vation and global change. It also highlights new approaches culled from the liter-
ature on topics such as species reintroductions, population viability analysis,
protected areas management, and payments for ecosystem services. Also new to
this edition is an Instructor's Resource CD, available to qualified adopting instruc-
tors of the text. This IRCD includes electronic versions of all the figures, photos,
and tables from the textbook.
xiv Preface
Acknowledgments
I sincerely appreciate the contribution of everyone who helped make this book ac-
curate and clear. Individual chapters in this edition were reviewed by Dana Bauer,
Patrick Bohlen, Katrina Brandon, Sue Bratton, Phil Cafaro, Linus Chen, Richard Cor-
lett, Chris Elphick, Richard Frankham, Elizabeth Freeman, Richard Griffiths, Susan
Jacobson, Christopher Johnson, Jeff McNeely, Michael Reed, Tom Ricciardi, Marcos
Robles, Eric Seabloom, Jodi Sedlock, Howard Snell, and Navjot Sodhi. Les Kaufman
of Boston University provided expertise on marine systems in all chapters.
Numerous people offered specialized input that helped make the boxes and case
studies current and accurate. I would particularly like to recognize the contribu-
tions of Kamal Bawa, Steve Bousquin, Marlin Bowles, David Bray, Jim Estes, Ed
Guerrant, Shen Guozhen, Kayri Havens, Rob Horwich, Daniel Janzen, Lukas Keller,
Cheryl Knot, Tom Kunz, Kerry Lagueux, Laurie Bingaman Lackie, Rodrigo Gamez
Lobo, Kathy MacKinnon, Elizabeth Marquard, Carlos Peres, Tom Power, Robert
Simmons, Lisa Sorenson, Michael Thompson, Sebastian Troenig, David Western,
Tony Whitten, Peter Wrege, Miriam Wyman, and Truman Young.
Rachel Morrison was the principal research assistant and organizer for the proj-
ect, with additional help from Jin Chung, Libby Ellwood, Elysia Heilig, Heather
Lieb, Farah Mohammedzadeh, Rebecca Norklun, Caroline Folgar, and Lily Smith.
Sydney Carroll and Kathaleen Emerson provided invaluable help in the produc-
tion of the book, with numerous suggestions on how to make the book friendlier
to student readers. Andy Sinauer, Chris Small, David McIntyre, Joan Gemme and
the rest of the Sinauer staff helped to transform the manuscript into a finished book.
Special thanks are due to my wife Margaret and my children Dan, Will, and Jasper
for encouraging me to fulfill an important personal goal by completing this book.
I would like to recognize Boston University for providing me with the facilities and
Preface xv
environment that made this project possible and the many Boston University stu-
dents who have taken my conservation biology courses over the years. Their en-
thusiasm and suggestions have helped me to find new ways to present this mate-
rial. And lastly, I would like to express my great appreciation to my coauthors in
other countries who have worked with me to produce conservation biology text-
books in their own languages, which are critical for spreading the message of con-
servation biology to a wider audience.
Richard Primack
Boston, Massachusetts
April, 2010
is so altered from its original condition that a site may no longer be suitable for
the continued existence of certain species.
There are many other threats facing modem ecosystems, including climate change
and invasive species. Efforts to protect a species in one area may be severely crip-
pled as a result of a rapid climate change to which the species cannot adapt (see
Chapter 9). Also. biol_ogica) cornmunities have been particularj.y,devastated by the
introduction of exotic species, some of which have been deliberately brought in
from other areas and established by people, such as domesticated animals and or-
namental plants, and some of which have been brought in accidentally, such as
weed species, insect pests, and new diseases. In many cases, particularly on islands,
these species have become invasive (see Chapter 10) and have displaced and elim-
inated native species.
Another major threat to biological diversity is the use of modem technolo to
over arvest anirna s an pants for loca an mtemationa markets. Hunters in trop-
ical forests now use guns and motorized vehicles, where before they used bows and
arrows and walked on foot. Fishing has changed from small wind- and hand-pow-
ered boats to large motorized fleets with freezers that can stay at sea for weeks or
months at aJirne. Entire forest, grassland, and ocean communities have been emp-
tied of their animal life and, in many cases, cleared of their plant life as well.
Pgwerful te_chnolo.gies allow alteration of the environment on a regional and even
a global scale. S_2tne ..9f t~ese transformations are inteptional, §llCh as the creatio:n
of darns and th~development of n_ew agrkultural land✓ but other change~, such as
iiirpolfution, strip-mining of entire hills, and damage to seabed habitats during
fishing, are by-prod~~ts of our 9-ctivities. Unregulated dumping ofchernicals and
sewage into streams, rivers, and lakes has polluted major freshwater and coastal
marine systems throughout the world and has driven significant numbers of species
toward extinction. Pollution has reached such high levels that even large marine
environments, such as the Mediterranean Sea, the Gulf of Mexico, and the Persian
Gulf, which were once assumed to be able to absorb pollution with no negative ef-
fects, are threatened with the loss of whole suites of formerly common species. Some_
ip.land water b~c!ies, such <!._S the Aral S~a, have been completely d~~d, along
with the many unique fish species that lived in them. Air pollution from factories
and cars has turned rainwater into an acid solution that weakens and kills moun-
tain trees downwind of industrial centers and, in tum, removes habitat for the an-
imals that depend on those plants. Scientists have warned that levels of air pollu-
tion have become severe enough to alter global climate patterns and strain the
capacity of the atmosphere to filter out harmful ultraviolet radiation. The impacts
of these events on ecosystems are enormous and ominous; they have also stimulat-
ed the growth of conservation biology.
Scientists now realize that man of the threats to biological diversit ares ner-
gis.tic-t a 1s, e negative e fects of several independent factors such as logging,
fire, poverty, and overhunting combine additively or even multiplicatively. Scien-
tists also know that the threats to biological diversity directly threaten human pop-
ulations because people are dependent on the natural environment for raw mate-
rials, food, medicines, and even the water they drink. And the poorest people are
the ones who will experience the greatest hardship from damaged environments.
how many of the world's species and natural areas will survive. People may some-
day look back on the first decades of the twenty-first century as an extraordinari-
ly exciting time, when a collaboration of determined people acting locally and in-
ternationally saved large numbers of species from extinction and even entire
ecosystems from destruction. Examples of such conservation efforts are described
later in this chapter and throughout this book.
~::> Conservation biology is an integrated, multidisciplinary scientific field that has
developed in response to the challenge of preserving species and ecosystems. It has
~egoals:
• to document the full range of biological diversity on Earth
• to investigate human impact on species, genetic variation, and ecosystems
The first two of these goals involve the dispassionate search for factual knowledge
typical of scientific research. The third goal, however, defines conservation biology
as a normative discipline-that is, it embraces certain values and attempts to apply
scientific methods to achieving those values. Like medical science, which applies knowl-
edge gleaned from physiology, anatomy, biochemistry, and genetics to the goal of
achieving human health and eliminating illness, conservation biologists intervene to
prevent the human-enhanced loss of biodiversity, because they believe the preserva-
tion of species and ecosystems to be an ultimate good (Nelson and Vucetich 2009).
BOX 1 .1 (continued)
Information from the TAMAR survey was critical to legis- facilities and small medical and dental clinics. Villagers are
lation passed in 1986 in Brazil that led to the complete pro- employed in making turtle-themed crafts to sell to tourists.
tection of sea turtles and the establishment of two new bi- To increase awareness of the program at the local level ,
ological reserves and a marine national park to protect TAMAR personnel give talks about marine conservation in
important nesting beaches. While creating protected areas village schools and organize hatch ling release ceremonies.
is important in conservation efforts, ongoing management The project reaches a wide audience in Brazil through
activities are also needed. Projeto TAMAR chose an innova- coverage in popular articles and on television programs. In
tive and comprehensive approach to protecting the turtles addition, TAMAR operates sea turtle educational centers
on the ground. They established conservation stations at where hundreds of thousands of tourists, most of whom
each of 21 main nesting beaches. The Brazilian government are from Brazil, visit each year. The tourists get to see con-
grants TAMAR complete responsibility for and control of the
beaches within these stations. Each station has a manager,
several university interns, and local employees. More than
85% of TAMAR's 1000 employees live on the coast; many are
former fish_ermen wh_o bring their knowledge of sea turtles
to bear on conservation . These local employees have be-
come strong advocates for the turtles because their wages
from Projeto TAMAR and the related tourist industry are
linked to the continuing presence of these animals.
The stations' personnel regularly patrol the conservation
areas on foot and by vehicle, measuring turtles for size and
permanently flipper-tagging all adults observed on the beach .
In places where predators are abundant, some nests are cov-
ered with wire mesh fitted with small gaps to protect the eggs
and then allow movement of the baby turtles after they hatch.
Alternatively, the eggs are collected and brought to nearby
Projeto TAMAR generates publicity for sea turtle conservation
hatchery areas, where they are reburied (Almeida and Mendes
by staging festive events involving tourists, school groups,
2007). These measures allow baby turtles emerging from pro- and local people, such as this release of hatchlings that were
tected nests or hatcheries to enter the ocean just as if they incubated in the safety of a protected hatchery. (Courtesy of
had emerged from natural nests. TAMAR protects over 4000 Projeto TAMAR Image Bank.)
turtle nests each year and has protected around 100,000 nests
and approximately 7 million hatch lings in the years since its
inception. On average, the number of turtle nests on the servation in action and receive a large dose of conservation
beaches has also been increasing by an impressive 20% a year education; in turn, they support the project through their
(Marcovaldi and Chaloupka 2007). purchase of souvenirs.
TAMAR is also working with the Brazilian government Projeto TAMAR has tried to involve the next generation
to protect and manage the nesting beaches on the offshore of concerned conservationists in current projects, helping
islands. The project has extended its mission to include pre- student interns experience success with a real-life conser-
venting turtles from getting caught in fishing nets while feed- vation project. Hopefully, the awareness raised by Projeto
ing in coastal waters. TAMAR provides fishermen with infor- TAMAR will extend gradually to other conservation programs.
mation about the importance of turtles and about fishing As a result of Projeto TAMAR's efforts in protecting thou-
gear designed to prevent turtle capture. Fishermen are also sands of adult turtles, tens of thousands of nests, and mil-
taught techniques for reviving turtles caught in their nets lions of hatchlings, sea turtle numbers in Brazil have stabi-
so the turtles will not suffocate. Their increasing apprecia- lized and even show signs of increasing. The project has
tion of turtles and their awareness of the new laws lead most changed people's attitudes, both in coastal villages and in the
fishermen to cooperate with these policies. However, acci- wider Brazilian society. By integrating conservation goals with
dental capture remains a leading cause of turtle mortality. community education and development, Projeto TAMAR has
Projeto TAMAR plays a positive role in the villages where improved the future for sea turtles and for local people in-
it operates. In many areas, TAMAR is the primary source volved with their conservation.
of income for the local people, often providing child care
10 Chapter 1
• The diversity of species and ecosystems should be preserved. The rich diversity of
iife should be protected. In general, most people agree with this principle
because they enjoy biological diversity. The hundreds of millions of visi-
tors each year to zoos, national parks, botanical gardens, and aquariums tes-
tify to the general public's interest in observing different species and ecosys-
tems (Figure 1.3). Genetic variation within species also sparks popular interest,
as shown by the wide appeal of pet shows, agricultural expositions, flower
exhibitions, and large numbers of specialty clubs (African violet societies,
rose societies, etc.). Home gardeners pride themselves on how many types
of plants they have in their gardens, while bird-watchers compete to see how
many species they can identify in one day or in their lifetimes. It has even
been suggested that humans may have a genetic predisposition to like bio-
logical diversity, called biophilia, from the Greek root words bio or "life" and
philia or "loving"; that is, to love living things (Kellert 1997; Corral-Ver-
dugo et al. 2009).
• The untimel extinction o opulations and species should be prevented. The ordi-
-nary extinction of species and populations as a result o natural processes
is an ethically neutral event. Ihrough the millennia of geological time, the
n!_tural extinction of each species has tended to be balanced by the evolu-
tion of new species. The local loss of a population of a species likewise is u~u-
ally offset by the establishment of a new population through dispersal. How-
ever, as a result of human activity, the rate of extinction has increased by
more than a hundredfold (see Chapter 7). Virtually all of the hundreds of
vertebrate species-and the presumed tens of thousands of invertebrate
species-that have gone extinct in the last few centuries have been wiped
out by humans. Many people now recognize their 'role and responsibility
in causing and, more important, in preventing extinctions.
FIGURE 1.3 People enjoy seeing the diversity of life, as shown by the growing popularity
of butterfly gardens. (Photograph by Richard B. Primack.)
What Is Conservation Biology? 11
FIGURE 1.4 Tanah Lot Temple is a Hindu temple on the island of Bali in Indonesia. Its
coastal setting allows worshippers to experience the connection of the human spirit with
the natural world. (Photograph© Hemis/ Alamy.)
In an ecological and cultural history of the Indian subcontit)ent, Gadgil and Guha
(1992) argue that the belief systems, religions, and myths of hunter-gatherer soci-
eties and stable agricultural societies tend to emphasize conservation themes and
the wise use of natural resources because these groups have learned over time to
What Is Conservation Biology? 13
live within the constraints of a fixed resource base. In contrast, the belief systems
of communities that raise livestock, and rapidly expanding agricultural and indus-
trial societies, emphasize the rapid consumption and destruction of natural resources
as a way to maximize growth and assert control over other groups. These groups
move to new localities when the resources of any one place are exhausted. Modern
industrial states represent the extreme of such societies. Their excessive and waste-
ful consumption requires the transportation of resources to urban centers in ever-
widening circles of resource depletion. However, what will we do when the re-
sources are all gone?
European Origins
To the European mind, the prevalent view has been that God created nature for
humans' use and benefit. In Genesis, the first book of the Bible, God instructs Adam
and Eve to "be fruitful and multiply and fill the Earth and subdue it; have domin-
ion over every living thing that moves upon the Earth." The biblical instruction sup-
ports a dominant tenet of Western philosophy: Nature should be converted into
wealth as rapidly as possible and used for the benefit of humans. This point of view
justifies neatly all land uses and implies that to leave land unused is to misuse God's
gift-a foolish, if not downright sinful, mistake. In medieval Europe, wilderness gen-
erally was perceived to be useless land and was often believed to be inhabited by
evil spirits or monsters, in contrast to the orderly qualities and appearance of agri-
cultural landscapes. This perspective of nature was not true in all places and in every
period, but it describes a general perception that is different from our view today.
This anthropocentric (human-centered) view of nature led to the exploitation
and degradation of vast resources in the regions colonized by European countries
from the sixteenth century onward (Diamond 1999). In practice, the wealth and ben-
efits that came from this policy accrued primarily to the citizens of the colonial pow-
ers, while the needs of non-European native peoples were largely disregarded. The
long-term ramifications for the forests, fisheries, and other natural resources them-
selves were not considered at all; the unexplored territories of the Americas, Asia,
Africa, and Australia seemed so vast and rich that it was inconceivable to the colo-
nial powers that their natural resources could ever be depleted.
An important element of the conservation movement did develop in Europe,
however, based on the experiences of scientific officers-often imbued with Roman-
tic idealism-who were sent to assist in the development of colonies in the eigh-
teenth and nineteenth centuries (Subashchandran and Ramachandra 2008). These
scientists were trained to make detailed observations on the biology, natural his-
tory, geography, and anthropology of the colonial regions. Many of them expected
to find the indigenous people living in wonderful harmony with nature. Instead,
they found devastated forests, damaged watersheds, and newly created poverty.
In European colonies throughout the world, perceptive scientific officers came to
see that protection of forests was necessary to prevent soil erosion, provide water
for irrigation and drinking, maintain wood supplies, and prevent famine. Some colo-
nial administrators also argued that certain intact forests should remain uncut be-
cause of their necessary role in ensuring a steady supply of rainfall in adjacent agri-
cultural areas-foreshadowing modern concern with global climate change. Such
arguments led directly to conservation ordinances. On the Indian Ocean island of
Mauritius, for example, the French colonial administration in 1769 stipulated that
25% of landholdings should remain forested to prevent erosion, degraded areas
should be planted with trees, and forests growing within 200 meters of water should
be protected. In order to prevent water pollution and the destruction of fish popu-
·lations, various colonial governments passed laws in the late eighteenth century reg-
ulating the.Pollutants being discharged by sugar mills and other factories. On a larg-
er scale, British scientists working in India issued a report in 1852 urging the
14 Chapter 1
(A) (B)
FIGURE 1.5 (A) Roland Savery's figure of the dodo in his picture of the Fall of Adam, in
the Royal Gallery at Berlin. This illustration was painted using a live dodo which was
brought to Europe in the early seventeenth century before the species went extinct. (B)
One of Europe's first nature reserves was established to protect the wisent in Poland. (B,
photograph© Liz Leyden/istockphoto.com.)
What Is Conservation Biology? 15
*For an explanation of the term hectare and other measurements, see Table 1.1.
Area
square meter (m2) Area encompassed by a square, each side of
which is 1 meter
1 hectare (ha) 1 ha = 10,000 m 2 = 2.47 acres
100 ha = 1 square kilometer (km2)
Mass
1 kilogram (kg) 1 kg = 2.2 pounds
1 gram (g) 1 g =1/1000 kg= 0.035 ounce
1 milligram (mg) 1 mg= 1/1000 g = 0.000035 ounce
Temperature
°C = 5/9(°F - 32)
degree Celsius (°C) 0°C = 32° Fahrenheit (the freezing point of water)
100°C = 212° Fahrenheit (the boiling point of water)
20°C = 68° Fahrenheit ("room temperature")
16 Chapter 1
American Origins
Among the first major intellectual figures in the United States arguing for the pro-
tection of natural areas were the nineteenth-century philosophers Ralph Waldo
Emerson and Henry David Thoreau (Callicott 1990). Emerson, in his transcenden-
talist writings, saw nature as a temple in which people could commune with the
spiritual world and achieve spiritual enlightenment (Emerson 1836). Thoreau was
both an advocate for nature and an opponent of materialistic society, believing that
people needed far fewer possessions than they sought. To prove his point, he lived
simply in a cabin near Walden Pond, writing about his ideas and experiences in a
book Walden , publishPd in J854 that has had a si nificant impact on man en-
erations of students and environmentalists. Thoreau believe t at the experience
of nature was a necessary counterweight to the weakening tendencies of civiliza-
tion. In his collection of essays (1863) he argued emphatically that
[in] wilderness is the preservation of the world .... The story of Romulus and
Remus [the founders of the Roman Empire] being suckled by a wolf is not a
meaningless fable. The founders of every state which has risen to eminence have
drawn their nourishment and vigor from a similar wild source.
This concern for preserving wilderness, large areas that remain essentially un-
occupied, unmanaged, and unmodified by human beings, is a continuing and dom-
inant theme in the American conservation movement up to the present time (Con-
gressional Research Service and Saundry 2009). It contrasts sharply with the
traditional European view that because landscapes developed over thousands of
years of human interaction, further management is appropriate in attempts to reach
conservation objectives (Cooper 2000).
Eminent American wilderness advocate John Muir used the transcendental
themes of Emerson and Thoreau in his campaigns to preserve natural areas. Ac-
cording to Muir's reservationist ethic, natural areas such as forest groves, moun-
tamto s, and w iritual values that are enera superior to t e tan-
gi le material gain obtained by their exploitation (Muir 1901). T is philosop y
emphasized the needs of pfolosophers, poets, artists, and spiritual seekers-who
require the beauty and stimulus of nature for their development-over the needs
of ordinary people, who require jobs and material goods from the natural environ-
ment. Some see this view as undemocratic and elitist, arguing that it disregards
the very real material needs of food, clothing, shelter, and employment, which may
require economic exploitation of the wilderness. Yet one does not have to be a mem-
ber of the elite in order to appreciate natural beauty: All human beings share these
impulses, and Muir's arguments for the spiritual and artistic value of nature did
not limit its accessibility or its benefits to a single stratum of society. That wilder-
JOHN MUIR ness can benefit all of society can be seen today in special programs, such as Out-
(1838-1914) ward Bound, that use experiences with nature and wilderness to challenge and en-
rich the character development and self-confidence of teenagers and young adults,
some of whom might otherwise succumb to drugs, crime, despair, or apathy.
In addition to advocating the preservation of nature on the grounds of human spir-
itual needs, Muir WgS among the first American conservationists to explicitly state
that nature has intrinsic value-value in and of itself, apart from its value to human-
.~ Muir argued on biblical grounds that because God had created nature and indi-
vidual species, to destroy them was undoing God's work. In Muir's view, species
have an equal place with people in God's scheme of nature (Muir 1916, p. 139):
Why should man value himself as more than a small part of the one great unit of
creation? And what creature of all that the Lord has taken the pains to make is
not essential to the completeness of that unit-the cosmos? The universe would
be incomplete without the smaHest transmicroscopic creature that dwells beyond
our conceitful eyes and knowledge.
What Is Conservation Biology? 17
Leopold eventually came to the conclusion that the most important goal of conser- ALDO LEOPOLD
vation is to maintain the health of natural ecosystems and ecological processes (Leopold (1887-1948)
18 Chapter 1
2004). As a result, he and many others lobbied successfully for certain parts of nation-
al forests to be set aside as wilderness areas (Shafer 2001). He also considered humans
part of the ecological community rather than standing apart from
and exploiting nature, as the proponents of the resource conserva-
Discussions qf natural resources, ecosystem
tion ethic argued. Despite Leopold's philosophical shift, he remained
management, and sustainable development committed to the idea that humans should be involved in land man-
are major themes throughout the field of agement, seeking a middle ground between overexploitation and
conservation biology. total control over nature, on the one hand, and complete preserva-
tion of land with no human presence or activity, on the other.
Leopold's synthesis has been termed the land ethic. In his writings and in prac-
tice at his talllly farm, Leopold advocated a land use policy in which human use
of natural resources was compatible with, or even enhanced, biological diversity
(Leopold 1939b, 1949). Integrating human activity into preservationist philosophy
also makes practical sense because complete exclusion of human impact from nat-
ural reserves has always been very difficult and is now becoming impossible be-
cause of human population growth, air pollution, and global climate change. An
approach that combines ideas of both Leopold and Pinchot has been developed,
known as ecosystem management, which places the highest management priority
on cooperation among businesses, conservation organizations, government a en-
cies, private citizens, an o er meres e parties to prov1 e or uman needs and
t~ aintam the health of wild species and ecosystems. -
Development of these philosophies has taken place alongside the growth of many
U.S. conservation organizations, such as the Wilderness Society, the Audubon So-
ciety, Ducks Unlimited, and the Sierra Club; the development of the national and
state park systems; and the passing of numerous environmental laws. Elements of
each of these differing philosophies are present in contemporary writings, the stat-
ed goals of conservation organizations, and government policy in both the United
States and other countries. Disagreements over policy and practice among and with-
in conservation organizations, individual conservationists, and government depart-
ments continue to reflect these long-term philosophical differences. This continu-
ing debate over elements of conservation philosophy and ethics is necessary in
deciding how to balance the long-term needs of protecting biological diversity with
the more immediate needs of modern society for natural resources.
Environmental activists, writers, and educators have applied these diverse
philosophies in ways that have benefited and transformed society. Ellen Swallow
Richards (1842-1911) was one such influential individual, though she had great dif-
ficulty obtaining a professional position as a chemist, a field not open to women
at that time. After being appointed as chemistry instructor at the Massachusetts In-
stitute of Technolo , she develo ed the first course in the new subject of ecolo .
In her many public activities she emphasized the nee to protect t e natural envi-
ronment as a key element in maintaining public health. Richards was particularly
concerned with how water quality was affected by sewage and industrial wastes,
and she began to test the quality of water in rivers and lakes. Her procedures led
to the first water quality standards in the country and eventually to the develop-
ment of modern sewage treatment plants that help protect public drinking supplies
ELLEN SWALLOW
as well as the natural environment.
RICHARDS
Another key figure was.Rachel Carson (1907-1964). In her widely read book Silent
(1842-1911) Spring (1962), she documented the role of pesticides and the chemical industry in
the loss of bird populations. At first she was heavily criticized by representatives
of the chemical industry. However, her tireless campaigning led to bans on DDT
in many countries and to better regulation of other toxic chemicals, and it was
crucial to the development of the modern environmental movement. The recov-
ery of numerous bird species, such as falcons, eagles, and ospreys, in the years fol-
What Is Conservation Biology? 19
lowing the ban on DDT proved that her observations were correct (see Box 9.1).
Carson was especially effective in changing public opinion through writing popu-
lar books, some specifically written for children.
Within the American conservation movement, other writers have prophetically
warned about the increasing destruction of biological diversity and the natural
environment (Meine 2001). Key authors extend from G. P. Marsh, with his Man and
Nature: Or, Physical Geography as Modified by Human Action (1864), and Fairfield
Osborn, author of Our Plundered Planet (1948), up to former U.S. Vice President
Albert Gore, author of An Inconvenient Truth: The Planetary Emergency of Global Warm-
ing and What We Can Do About It (2006), and Jared Diamond, with his decisive
historical analysis Collapse (2005). These authors have found a receptive general au-
dience and have galvanized citizens by the millions to join efforts to protect birds
and other wildlife; to conserve mountains, seashores, wetlands, and other habitats;
and to limit environmental pollution (Leisher 2008). Over the past decade, a new RACHEL CARSON
crop of writers has emerged to address growing concern with global climate change (1907-1964)
and damage to the world's oceans.
Despite the threats to biological diversity, we can detect many positive signs that
allow conservation biologists to remain cautiously hopeful. The number of people
living in extreme poverty has been in decline since the Industrial Revolution, and
the rate of human population growth has slowed (Sachs 2008). The number of pro-
tected areas around the globe continues to increase, with a dramatic expansion in
the number of marine protected areas. In just 2006, the South Pacific country of Kiri-
bati established the world's largest marine sanctuary. Moreover, our ability to pro-
tect biological diversity has been strengthened due to a wide range of local, nation-
al, and international efforts. Certain endangered species are now recovering as a
FIGURE 1. 7 The Society for
result of conservation measures. We can point to an expansion of our knowledge Conservation Biology has a
base and the science of conservation biology, the developing linkages with rural de- simple, yet powerful, logo
velopment and social sciences, and our increased ability to restore degraded envi- showing the circle of life, with-
ronments. All of these suggest that progress is being made despite the enormous in which we live. The ocean
tasks still ahead. waves in the center symbolize
the changes that lie ahead. The
logo can also be viewed as a
bird, which provides us with
Summary beauty; on closer look, we see
that its wings are really rustling
1. Thousands of species are goin extinct, genetic variation is being lost, millions of pop- leaves. (Courtesy of the Society
ulations are isappearing, and entire ecos stems are being estroyed as a result o for Conservation Biology.)
numan activities. onservation biology is a e~tic:_ dis~e combining basTcand
~p1iecfresearch to describe biological diversitict'oclirnent the threats it faces from
human activities, and develop methods to protect and restore biological diversity.
2. Conservation biology rests on a number of underlying assumptions that are accept-
ed oy~most conservat10n biologists: b1olog1cal d1vers1ty, including the range of
species, genetic vanahon, 610log1cal communities, and ec;:osystem interactions,
should be preserved; the extinction of species by human activities should be pre-
-vented; the !=Om lex inter chon of s ec1es m natural communffies sncmld be main-
tained; evolutionary change should continue; and biological diversity has va ue in
and of itself.
3. Conservation biology draws on both scientific and religious/philosophical tradi-
tions. European scientists in the eighteenth and nineteenth centuries reacted to the
aestruction of forests and water pollution in their colonies by proposing some of the
first environmental legislation. The decline and extinction of species in Europe led
to the establishment of the first nature reserves and an active popular interest in con-
servation. In the United States, Henry David Thoreau and John Muir argued for the
preservation of wilderness and the intrinsic value of species. Gifford Pinchot pro-
posed developing a balance among competing natural resource needs for present
and future societies. Aldo Leopold advocated striking a balance between managing
land for ecological processes and satisfying human needs. These philosophies -still
guide land management, and elements of them can be found in the current doctrines
of conservation organizations and government departments.
For Discussion
1. How is conservation biology fundamentally different from other branches of biolo-
gy, such as physiology, genetics, or cell biology? How is it different from environ-
mentalism?
2. What do you think are the major conservation and environmental problems facing
the world today? What are the major problems facing your local community? What
ideas for solving these problems can you suggest? (Try answering this question now,
and once again when you have completed this book.)
22 Chapter 1
3. Consider the public land management and private conservation organizations with
which you are familiar. Would you consider their guiding philosophies to be clos-
est to the resource conservation ethic, the preservation ethic, or the evolutionary-eco-
logical land ethic? What factors allow them to be successful or limit their effective-
ness? Learn more about these organizations through their publications and Web sites.
4. How would you characterize your own viewpoint about the conservation of biodi-
versity and the environment? Which of the religious or philosophical viewpoints of
conservation biology stated here do you agree or disagree with? How do you, or
could you, put your viewpoint into practice?
Suggested Readings
Chan, K. M. A. 2008. Value and advocacy in conservation biology: Crisis discipline or dis-
cipline in crisis? Conservation Biology 22: 1-3. Conservation biologists need to be more
effective advocates on behalf of biodiversity.
Czech, B. 2008. Prospects for reconciling the conflict between economic growth and biodi-
versity conservation with technological progress. Conservation Biology 22: 1389-1398. Is
it possible to balance ever greater human use of natural resources with the need to pro-
tect biodiversity?
Diamond, J. 2005. Collapse: How Societies Choose to Fail or Succeed. Viking Press, New York. An
eminent biologist describes patterns of environmental catastrophe that have destroyed
human societies in the past and that threaten us today.
Dudley, N., L. Higgins-Zogib, and S. Mansourian. 2009. The links between protected areas,
faiths, and sacred natural sites. Conservation Biology 23: 568-577. In many places, local peo-
ple are already protecting biodiversity.
Hall, J. A. and E. Fleishman. 2010. Demonstration as a means to translate conservation sci-
ence into practice. Conservation Biology 24: 120-127. Conservation biologists need to demon-
strate to the public and government how their ideas work in practice.
Leisher, C. 2008. What Rachel Carson knew about marine protected areas. BioScience 58:
478-479. The key to effective MPAs is often getting local people involved.
Leopold, A. 1949. A Sand County Almanac. Oxford University Press, New York. Leopold's
evocative essays articulate his "land ethic," defining human duty to conserve the land
and the living things that thrive upon it.
Marris, E. 2007. What to let go. Nature 450: 152-155. In this time of crisis, scientists are iden-
tifying species and ecosystems that are the highest priorities for funding and conserva-
tion action.
Millennium Ecosystem Assessment (MEA). 2005. Ecosystems and Human Well-Being. 4 vols.
Island Press, Washington, D.C. Detailed report and summary by the world's leading sci-
entists documenting the importance of ecosystem services.
Morell, V. 1999. The variety of life. National Geographic 195 (February): 6-32. Special issue
includes beautifully illustrated articles about biodiversity, threats to its existence, and key
conservation projects.
Nelson, M. P. and J. A. Vucetich. 2009. On advocacy by environmental scientists: What,
whether, why, and how. Conservation Biology 23: 1090-1101. Scientists have a responsibil-
ity to be advocates as well as researchers.
Orr, D. W. 2007. Optimism and hope in a hotter time. Conservation Biology 21: 1392-1395. Hope
means to learn the truth and then have the courage to act accordingly.
Papworth, S. K., J. Rist, L. Coad, and E. J. Milner-Gulland. 2009. Evidence for shifting base-
line syndrome in conservation. Conservation Letters 2: 93-100. People often forget what
biodiversity was like in the past.
Van Heezik, Y. and P. J. Seddon. 2005. Conservation education structure and content of grad-
uate wildlife management and conservation biology programs: An international perspec-
tive. Conservation Biology 19: 7-14. Conservation education programs are increasing and
are highly diverse.
World Resources Institute (WRI). 2005. World Resources 2005: The Wealth of the Poor-Manag-
ing Ecosystems to Fight Poverty. World Resources Institute, Washington, D.C. Massive body
of data on biodiversity and the human condition.
KEY JOURNALS IN THE FIELD Biodiversity and Conservation, Biological Conservation, Bio-
Science, Conservation Biology, Conservation Letters, Ecological Applications, Journal of Ap-
plied Ecology, National Geographic
What Is Biological Diversity?
/ I I
Community/ecosystem diversity
Species Diversity
Species diversity includes the entire range of species found on Earth. Recognizing
and classifying species is one of the major goals of conservation biology (Morell
1999). How do biologists identify individual species among the mass of living or-
ganisms on Earth, many of them small in size and with few distinguishing features?
And what is the origin of new species? Identifying the process.whereby one species
evolves into one or more new species is one of the ongoing accomplishments of
modem biology. The origin of new species is normally a slow process, taking place
What Is Biological Diversity? 25
over hundreds, if not thousands, of generations. The evolution of higher taxa, such
as new genera and families, is an even slower process, typically lasting hundreds
of thousands or even millions of years. In contrast, human activities are destroying
in only a few decades the unique species built up by these slow natural processes.
What Is a Species?
A species is generally defined in one of two ways:
1. ~ group of individuals that is morphologically,* physiologically, or biochem-
ically distinct from otner groups in some important charac_teristic is t h ~
phological definition of species.
2. A_group of individu~ls that can potentially breed among themselves_in_t&
.wild and that do not breed with individuals of other groups is the _biologi-
cal definition of species.
Because the methods and assumptions used are different, these two approach-
es to distinguishing species sometimes do not give the same results. Increasingly,
differences in DNA sequences and other molecular markers distinguish species that
look almost tdentical, _such as types of bacteria (Janzen et al. 2009).
The mor holo ical definition of species is the one most commonly used by tax-
onomists, biologists who specialize int e identification of unknowns ecimens and
the classification of species (Figure 2.2). In practice, t e biological definition of species
*An individual's morphology is its form and structure-or, to put it more simply (if not totally
accurately), its appearance.
(A) (B)
:·..=~. 1 i· •
f ~:
~
•
..
·~ ~+
'
. II i'
'· '
:'i· : ' ,·
FIGURE 2.2 (A) An entomologist collects moth specimens that land on a lighted white
sheet. (B) An ornithologist at the Museum of Comparative Zoology, Harvard University
classifying collections of orioles: Black-cowled Oriole (Icterus prosthemelas) from Mexico
and Baltimore Orioles (Icterus galbula) which occur throughout eastern North America. (A,
photograph© The Natural History Museum/ Alamy; B, photograph courtesy of Jeremiah
Trimble, Museum of Comparative Zoology, Harvard University© President and Fellows
of Harvard College.)
26 Chapter 2
Kingdom: Animalia
>1,000,000 species
\~~. •
.
:VT'l,,..~a
'
'
~.',,
. -~rz~
,/2'•,.,·/
I
~· ~- ri
. , ... '!/_,
~ • I
:.
.
,
.'.
•
Phylum: Chordata
, .. ,,,, ~~~ .. •
"'::· ';~
Class: Aves (birds) I ,;.,
, .
8600 species
'
Family: Parulidae
(New World '
warblers)
125 species
_- -<:\i.
Genus: Dendroica
(various warblers)
28 species
Blackburnian warbler
BOX 2 .1 (continued)
Similar phyla.are grouped into a kingdom: all animal naeus. In the scientific name for the Blackburnian warbler,
classes belong to the kingdom Animalia.* Dendroica fusca, Dendroica is the genus name and fusca is
Biologists throughout the world have agreed to use a the species name. The genus name is somewhat similar to
standard set of scientific, or Latin, names when discussing a person's family name in that many people can have the
species. The use of scientific names avoids the confusion same family name (Sullivan), while the species name is sim-
that can occur when using common names; the Latin names ilar to a person's given name (Margaret).
are standard across countries and languages. Scientific Scientific names are written in a standard way to avoid
species names consist of two words. This naming system, confusion. The first letter of the genus name is always cap-
known as binomial nomenclature, was developed in the italized, whereas the species name is almost always low-
eighteenth century by the Swedish biologist Carolus Lin- ercased. Scientific names are italicized in print or under-
lined when handwritten. Sometimes scientific names are
*Until recently, most modern biologists recognized five kingdoms followed by a person's name, as in Homo sapiens Linnaeus,
in the living world: plants, animals, fungi, monerans (single- indicating that Linnaeus was the person who first proposed
celled species without a nucleus and without mitochondria, such the scientific name given to the human species. When many
as bacteria), and protists (more complex single-celled species
with a nucleus and mitochondria). With the increasing sophistica- species in a single genus are being discussed, or if the iden-
tion of molecular techniques, many biologists now use a system tity of a species within a genus is uncertain, the abbrevia-
of classification with six kingdoms within three domains: Bacteria tions spp. or sp., respectively, are sometimes used (e.g., Den-
(common bacteria), Archaea (ancient bacteria that live in extreme
environments, such as hypersaline pools, hot springs, and deep droica spp.). If a species has no close relatives, it may be the
sea vents), and the Eucarya (all organisms with a membrane- only species in its genus. Similarly, a genus that is unrelat-
bound nucleus, including animals, plants, fungi, and protists). ed to any other genera may form its own family.
little as a few percent. The inability to clearly distinguish one species from anoth-
er, whether due to similarities of characteristics or to confusion over the correct sci-
entific name, often slows down efforts at species protection. It is difficult to write
precise, effective laws to protect a species if scientists and lawmakers are not cer-
tain what name should be used. At the same time, species are going extinct before
they are even described. Tens of thousands of new species are being described each
year, but even this rate is not fast enough. The key to solving this problem is to train
more taxonomists, especially for work in the species-rich tropics (Wilson 2003). We'll
return to this topic in Chapter 3.
*That evolution occurs is regarded by virtually all biologists as fact. Several popular and scholarly
books (e.g., Futuyma 2009; Shanks 2004) discuss religion-based arguments (and intelligent-design ar-
guments) against evolution and why most scientists do not accept such arguments.
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