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FUNDAMENTALS
OF ECOSYSTEM SCIENCE
FUNDAMENTALS
OF ECOSYSTEM
SCIENCE
SECOND EDITION
Edited by
KATHLEEN C. WEATHERS
DAVID L. STRAYER
GENE E. LIKENS
Academic Press is an imprint of Elsevier
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than as may be noted herein).
Notices
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understanding, changes in research methods, professional practices, or medical treatment may become
necessary.
Practitioners and researchers must always rely on their own experience and knowledge in evaluating and using
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Library of Congress Cataloging-in-Publication Data
A catalog record for this book is available from the Library of Congress
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A catalogue record for this book is available from the British Library
ISBN 978-0-12-812762-9
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Publisher: Candice Janco

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Contributors
Elena M. Bennett Bieler School of Environment Gary M. Lovett Cary Institute of Ecosystem
and Department of Natural Resource Sciences, Studies, Millbrook, NY, United States
McGill University, Montreal, QC, Canada Pamela A. Matson Stanford University,
Mary L. Cadenasso Department of Plant Stanford, CA, United States
Sciences, University of California, Davis, Judy L. Meyer Odum School of Ecology,
United States University of Georgia, Athens, GA, United
Cayelan C. Carey Virginia Tech, Blacksburg, States
VA, United States Richard S. Ostfeld Cary Institute of Ecosystem
Jonathan J. Cole Cary Institute of Ecosystem Studies, Millbrook, NY, United States
Studies, Millbrook, NY, United States Michael L. Pace Department of Environmental
Holly A. Ewing Bates College, Lewiston, ME, Sciences, University of Virginia, Charlottesville,
United States VA, United States
Stuart E.G. Findlay Cary Institute of Ecosystem Steward T.A. Pickett Cary Institute of
Studies, Millbrook, NY, United States Ecosystem Studies, Millbrook, NY, United
Robinson W. Fulweiler Department of Earth States
and Environment, Department of Biology, Emma J. Rosi Cary Institute of Ecosystem
Boston University, Boston, MA, United States Studies, Millbrook, NY, United States
Peter M. Groffman Cary Institute of Ecosystem Meagan E. Schipanski Department of Soil and
Studies, Millbrook; City University of New Crop Sciences, Colorado State University, Fort
York, Advanced Science Research Center at Collins, CO, United States
the Graduate Center and Brooklyn College Christopher T. Solomon Cary Institute of
Department of Earth and Environmental Ecosystem Studies, Millbrook, NY, United
Science, New York, NY, United States States
Stephen K. Hamilton Cary Institute of Emily H. Stanley University of Wisconsin-
Ecosystem Studies, Millbrook; Kellogg Madison, WI, United States
Biological Station and Department of
David L. Strayer Cary Institute of Ecosystem
Integrative Biology, Michigan State
Studies, Millbrook, NY; Graham Sustainability
University, Hickory Corners, MI, United States
Institute, University of Michigan, Ann Arbor,
Oleksandra Hararuk University of Central MI, United States
Florida, Orlando, FL, United States
R. Quinn Thomas Virginia Tech, Blacksburg,
Clive G. Jones Cary Institute of Ecosystem VA, United States
Studies, Millbrook, NY, United States
Kathleen C. Weathers Cary Institute of
Gene E. Likens Cary Institute of Ecosystem Ecosystem Studies, Millbrook, NY, United
Studies, Millbrook, NY; University of States
Connecticut, Storrs, CT, United States
ix
Preamble to the 2nd Edition
As we write this, the world is gripped by a ecosystems in the form of changes in eco-
devastating pandemic (COVID-19). The nomic demand, trade patterns, pollutant re-
socioecological causes, responses to, leases, and so on. Thus, the COVID-19
and ramifications of COVID-19 have been pandemic reminds us that diseases are an
and will be profound, and they will play ecological problem as well as a medical or
out over short- and long-term scales. Because veterinary problem, and their prevention
we are writing in the midst of this pandemic, and solution will require understanding
the specifics about its ultimate resolution and application of ecological ideas
and effects are far from clear. What is clear and principles. Although pandemics such
from the COVID-19 and previous pan- as COVID-19 have had analogs in the
demics, though, is that ecosystems can affect past (e.g., Black Death in the 1300s, influenza
the emergence and spread of disease (as in 1918), conditions are far different now
discussed by Rick Ostfeld in his essay in this than then—from the number
book), and infectious diseases can have enor- and distribution of people on the planet,
mous effects on ecosystems. Human impacts to the degradation and change of ecosys-
such as habitat alteration, rapid global move- tems worldwide from human activities
ment of plants, animals, people, and the interconnectedness of natural
and pathogens, and the size and density and human systems. These factors may tend
of the human population itself favor the to make future pandemics more frequent
emergence and spread of disease. In turn, and more severe, and lend urgency to the
as we are seeing in this time of COVID-19, effective application of ecosystem science
diseases have such strong effects on human and other sciences to understanding as
activities that they can feed back onto well as the management of our world.
xi
Preface
This book provides an introduction to the “voices” will be evident throughout the
content, ideas, and major findings of contem- book. We believe that this diversity reflects
porary ecosystem science. We wrote the book some of the myriad perspectives and ap-
primarily for beginning graduate students proaches that are fruitfully brought to bear
and advanced undergraduates, but it should on the field of ecosystem science.
also be useful to a broad range of academic The book contains six major sections. The
scientists and resource managers, and even opening chapter introduces the concept of
to dedicated amateurs who seek an intro- the ecosystem, explores some of the conse-
duction to the field. Ecosystem science is a quences of this concept, describes the intel-
rigorous, quantitative science; we assume lectual tools of the science, and briefly
that readers of the book will have had an reviews the history of this young science.
introductory class in ecology and basic Chapters 2–9 lay the foundation for the study
understanding of chemistry and math. The of ecosystems, and cover the two major
book deliberately covers multiple approa- branches of ecosystem science: energetics
ches to understanding ecosystems (e.g., the (Chapters 2–5) and biogeochemistry
use of experiments, theory, cross-system (Chapters 5–9). These chapters present the
comparisons), in multiple environments (ter- core content of ecosystem science—the
restrial, freshwater, and marine; managed, movement and fate of energy and materials
built, and natural ecosystems), across all parts in ecosystems—in some detail. In the syn-
of the world (although many examples come thetic Chapters 10–12, we revisit major
from the authors’ experience in North themes that cut across multiple areas of
America). study in ecosystem science. The authors of
The origins of this book stem from an these chapters review the power and utility
intensive 2-week Fundamentals of Ecosystem of the ecosystem concept, the roles of hetero-
Ecology class (the FEE class) that we have geneity in space and time, and the impor-
taught to graduate students from around tance of various types of controls in
the world every year or two at the Cary Insti- ecosystems. Chapters 13–18 take ecosystem
tute of Ecosystem Studies since 1989. We, and science into the real world by illustrating,
many of the chapter authors, have played cen- through six case studies, the value of ecosys-
tral roles in the development, evolution, and tem science in identifying and solving
running of the FEE class since its origin. a range of environmental problems. The
We decided upon an edited book for sev- book closes with Chapter 19, which lays
eral reasons, not the least of which was its out some challenges and needs for the future.
genesis in this team-taught course. While Today’s ecosystem science is evolving rap-
we shepherded and integrated the chapters idly, with major new discoveries and ideas
and their contents, we also deliberately emerging every year. The ultimate shape
allowed—and even encouraged—multiple and contributions of this science remain to
approaches, and as a result, multiple be discovered.
xiii
xiv Preface
This book benefited from the persistent their scholarship, patience, goodwill, and
and hard work of the Academic Press team, commitment to bringing this project to fru-
especially Jill Cetel, Candice Janco, and ition. The Cary Institute’s assistant, Matt
multiple graphic artists. We were also fortu- Gillespie, was an enormous help as well.
nate to have received helpful and critical Finally, generations of FEE students were
reviews of chapters from colleagues, includ- and continue to be an impetus and inspira-
ing Alexandra Ponette-González, Clifford tion to us and the field of ecosystem science.
Ochs, and several anonymous reviewers
who teach ecosystem science; their com- Kathleen C. Weathers
ments substantially improved the book. We David L. Strayer
thank the authors of various chapters for Gene E. Likens
C H A P T E R
1
Introduction to Ecosystem Science
Kathleen C. Weathersa, David L. Strayera,
and Gene E. Likensa,b
a
Cary Institute of Ecosystem Studies, Millbrook, NY, United States bUniversity of Connecticut,
Storrs, CT, United States
Humans have devised many intellectual systems to understand and manage the
complicated world in which we live, from physics to philosophy to economics. In this book,
we present one such intellectual system, ecosystem science, which tries to make sense of the
complex natural world and helps us manage it better. As we will see, ecosystems can be
highly varied in size and character, from a little pool of water in a tree cavity, to a redwood
forest, to a neighborhood in a city, to a frigid river, to the entire globe (Figure 1.1).
Nevertheless, a common set of tools and ideas can be used to analyze and understand these
varied and complicated systems. The results of these analyses are both intellectually
satisfying and useful in managing our planet for the benefit of nature and humankind.
Indeed, because of the growing demands placed on living and nonliving resources by
humans, it has been argued that ecosystem science is one of the essential core disciplines
needed to understand and manage the modern planet Earth (Weathers et al. 2016).
This book defines the ecosystem, illustrates the ecosystem approach, describes the chief
characteristics of ecosystems and the major tools that scientists use to analyze them, and
presents important discoveries that scientists have made about ecosystems. It also lays out
some critical questions for the future. Although the book is not focused on the management
of ecosystems, several management implications of ecosystem science are described and
illustrated.
What Is an Ecosystem?
An ecosystem is the interacting system made up of all the living and nonliving objects in a
specified volume of space.
This deceptively simple definition both says much and leaves out much. First, as with other
systems (Box 1.1), ecosystems contain more than one object, and those objects interact. More
Fundamentals of Ecosystem Science 3 Copyright # 2021 Elsevier Inc. All rights reserved.
https://doi.org/10.1016/B978-0-12-812762-9.00001-0
4 1. Introduction to Ecosystem Science
FIGURE 1.1 Some examples of ecosystems: (A) the frigid Salmon River, Idaho; (B) a residential neighborhood in
Baltimore, Maryland; (C) a biofilm on a rock in a stream; (D) a section of the southern ocean containing a phytoplank-
ton bloom; (E) a redwood forest in the fog in California; (F) a tree cavity; (G) the Earth. (Photocredits: A—John Davis;
B—Baltimore Ecosystem Study LTER; C—Colden Baxter; D—US government, public domain; E—Samuel M. Simkin;
F—Ian Walker; G—Source: https://www.publicdomainpictures.net/en/view-image.php?image=86448&picture=
planet-earth.)
surprisingly, living and nonliving objects are given equal status in ecosystem science.
A particle of clay and the plant drawing its nutrition from that clay particle are both parts
of an ecosystem, and therefore equally valid objects of study. This viewpoint contrasts with
physiology and population ecology, for example, in which the organism is the object of study,
I. Introduction
What Is an Ecosystem? 5
BOX 1.1
Some Nonecological Systems

Thinking about some of the many familiar with other banks. Systems may be described
examples of nonecological systems may help according to their controls as well. Gravity
you understand how ecosystems are de- and rotational dynamics control the motions
scribed and compared. A system is just a col- of the planets, and the copper atom is con-
lection of two or more interacting objects. A trolled by strong and weak atomic forces,
few familiar systems include the group of whereas the Federal Reserve System is con-
planets rotating around the sun as a system trolled by the decisions of its Board of Gover-
(the solar system); the group of electrons, nors (who, in turn, are chosen by a president
protons, and neutrons forming an atom; who is elected by the voters of the United
and the system of banks that controls the States). All of these descriptions allow us to
money supply of the United States (the Fed- understand how each system works. Perhaps
eral Reserve System). Just as with eco- more importantly, they let us compare one
systems, we can describe these systems by system to another—our solar system with
their structures, their functions, and the fac- those of other stars; the copper atom with
tors that control them. the cadmium atom; the current banking
A description of system structure often be- system in the United States with that of
gins with the number and kinds of objects in France, or with that of the United States in
the system. Thus, we might note that our so- the 19th century. Ecosystem scientists like-
lar system contains eight or nine planets; or wise describe ecosystems in various ways to
that the copper atom has 29 electrons, 29 pro- understand them better, and to allow com-
tons, and 35 neutrons; or that the Federal Re- parisons across ecosystems.
serve System contains a seven-member Board Systems science, the general field of
of Governors, 12 banks, and 26 branch banks. understanding all kinds of systems, is well
Systems have functional properties as well— developed. Many of the conceptual frame-
the copper atom exchanges electrons with works for ecosystem science are those of
other atoms in chemical reactions, and the system science (e.g., Hogan and Weathers
Federal Reserve System exchanges money 2003; Mobus and Kalton 2015).
and the nonliving environment is conceived of as an external influence on the object of study.
Finally, the definition implies that ecosystems have definite boundaries, but does not tell us
how we might go about setting or finding the boundaries to an ecosystem.
There are some unexpectedly powerful advantages to this simple definition. First, by in-
cluding all living and nonliving objects in a specified space, it is possible to use the tool of
mass balance to follow the movement and fate of materials (Box 1.2). Material that comes into
an ecosystem must either stay in the ecosystem or leave—there is no other possible fate for the
material. Mass balance offers a convenient quantitative tool for measuring the integrated
activity of entire, complicated systems without having to measure the properties and inter-
actions of each of its parts. It also allows ecosystem scientists to estimate the size of a single
unknown flux by difference. Consequently, it will become evident throughout the book that
ecosystem scientists often use the powerful tool of mass balance.
I. Introduction
BOX 1.2
Ecosystem Goods and Services

Ecosystems provide many valuable goods in dollars, whereas others are more elusive.
and services to people. People have recog- Thus, it seems easy to place a value on X
nized for a long time that ecosystems provide board feet of oak timber sold on the open
physical, marketable products such as timber market in the year 2015 for Y dollars, but
and fish, and have often managed ecosystems how does one quantify the spiritual satisfac-
to protect or increase the supply of these tion that arises from contemplating a flowing
goods. As economics and ecology have devel- river? Nevertheless, economists have devel-
oped, it has become apparent that ecosystems oped methods to estimate the value of even
provide many things other than marketable the most elusive of ecosystem services (EPA
goods that are of value to people. For in- Science Advisory Board 2009). Third, it is a
stance, ecosystems may remove pollutants, fundamental and serious mistake to assume
reducing the cost of providing drinking water that ecosystem services that are hard to quan-
or clean air to breathe. They may protect us tify are trivially small, and can be left out of
from diseases, or protect our infrastructure analyses. For instance, cultural ecosystem
from flooding. People may get peace of mind, services typically are difficult to quantify,
write songs (Coscieme 2015), or even heal and so are often omitted from estimates of
faster (Ulrich 1984) when they have access ecosystem services. Yet Carson et al. (2003)
to natural ecosystems. estimated that the existence value associated
One commonly used framework for enu- with the Exxon Valdez oil spill in Prince Wil-
merating and organizing these diverse bene- liam Sound was at least USD $2.8 billion (in
fits (Millennium Ecosystem Assessment 2003, 1990 dollars), for example. Fourth, if we are
2005) organizes the benefits that ecosystems to use an ecosystem services framework to
provide to people into four broad classes guide the management of ecosystems and
(Table 1.1). We offer several observations aid in environmental decision-making so that
about this list (or indeed any list of ecosystem aggregate ecosystem benefits to humans are
services). First, the list of goods and services maximized, then it is essential to include all
that ecosystems provide to people is long and kinds of ecosystem services in the analysis.
varied, ranging from tangible goods sold on It’s easy to see that deciding whether to build
the open market to the least tangible of bene- a dam by considering only the hydroelectric-
fits, and including everything from physical ity to be generated, but not the effects of the
and biogeochemical characteristics of ecosys- dam on navigation, fisheries, or recreation,
tems to specific parts of specific plants and is likely to lead to a poor decision. Similarly,
animals. Attempts to put a dollar value on considering any subset of ecosystem services
these services often result in very large esti- instead of the entire array of services is likely
mates. For example, a famous early analysis to result in a decision that does not maximize
by Costanza et al. (1997) estimated the global benefits to people. Finally, many ecologists
value of ecosystem services to be USD $16–54 object to any attempt at reducing the value
trillion/year. Second, some of these ecosys- of nature to a dollar value, considering it in-
tem services are easy to quantify and value appropriate.
Continued
I. Introduction
What Is an Ecosystem? 7
BOX 1.2 (cont’d)
TABLE 1.1 Categories of ecosystem services.

Type of Service Examples
Provisioning Provision of food, fresh water, wood, fiber, and biochemicals such as natural medicines
directly to humans
Regulating Regulation of climate, air quality, diseases, erosion, or natural hazards such as floods;
pollination
Cultural Provision of recreational, aesthetic, religious, spiritual, or educational opportunities to
people
Supporting Where an ecosystem provides structures or functions that support any of the other three
classes of services; examples include soil formation and nutrient cycling
Modified from Millennium Ecosystem Assessment (2003, 2005) and Costanza et al. 2017.
Second, defining an ecosystem as we have done makes it possible to measure the total ac-
tivity of an ecosystem without having to measure all the parts and exchanges within the eco-
system. This advantage is sometimes referred to as a “black-box” approach, because we can
measure the function (input and output) of a box (the ecosystem) without having to know
what is in the box (Figure 1.2). Sometimes ecologists debate whether it is philosophically pos-
sible to predict the properties of a complex system by studying its parts (reductionism) or
whether it is necessary to study intact systems (holism). It is not necessary to accept the phil-
osophical claims of holism, though, to recognize that studies of whole systems may be a much
more efficient way than reductionism to understand ecosystems. Such a holistic approach to
ecosystems is a powerful tool of ecosystem science, and is often combined with reductionist
approaches to develop insights into the functioning and controls of ecosystems.
ECOSYSTEM BOUNDARY
ORGANIC COMPARTMENT
ATMOSPHERIC Litter
COMPARTMENT Biological uptake
Meteorologic Meteorologic
of gases and aerosols Herbicore Geologic Geologic
Windblown particulates Living Dead
and gases above and
Biological release of Plant
Carnivore
Biomass
Biologic Biologic
below ground
gases and organic aerosols Omnivore
Biomass
INPUT INPUT
Detritivore
W
et
leaching, throughfall,
an
Dry deposition
stemflow, exudation
d
dr
Mineralization,
Inorganic
aerosols
yd
Re ep
Biological
leas os BIOSPHERE BIOSPHERE

uptake
e itio
of n
ga
ses
OUTPUT OUTPUT
AVAILABLE NUTRIENT
SOIL AND ROCK Weathering COMPARTMENT
MINERAL Meteorologic Meteorologic
COMPARTMENT
on in Geologic Geologic
Formation of exchange soil Biologic Biologic
secondary minerals sites solution
INTRASYSTEM CYCLE
FIGURE 1.2 Two views of the same ecosystem. The left side shows some of the parts inside an ecosystem and how
they are connected, as well as the exchanges between the ecosystem and its surroundings, whereas the right side
shows a black-box approach in which the functions of an ecosystem (i.e., its inputs and outputs) can be studied with-
out knowing what is inside the box. (Modified from Likens 1992.)
I. Introduction
Third, the definition gives the investigator complete flexibility in choosing where to set the
boundaries of the ecosystem in time and space. The boundaries of an ecosystem (i.e., size,
location, and timescale) can therefore be chosen to match the question that the scientist is try-
ing to answer. Boundaries often are drawn at places where fluxes are easy to measure (e.g., a
single point on a stream as it leaves a forested, watershed-ecosystem) or so that fluxes across
the boundary are small compared to cycling inside the ecosystem (e.g., a lake shore). Never-
theless, boundaries are required to make quantitative measures of these fluxes. It is true that
ecosystems frequently are defined to be large (e.g., lakes and watersheds that are hectares to
square kilometers in size) and are studied on the scale of days to a few years, but there is noth-
ing in the definition of an ecosystem that requires ecosystems to be defined at this scale. In-
deed, as we will see, an ecosystem may be as small as a single rock or as large as the
entire Earth (see Chapter 7), and can be studied for time periods as long as hundreds of mil-
lions of years.
Fourth, defining an ecosystem to contain both living and nonliving objects recognizes the
importance of both living and nonliving parts of ecosystems in controlling the functions and
responses of these systems. There are examples throughout the book in which living organ-
isms, nonliving objects, or both acting together determine what ecosystems look like (struc-
ture) and how they work (function). Furthermore, the close ties and strong interactions
between the living and nonliving parts of ecosystems are so varied and so strong that it would
be very inconvenient if not misleading to study one without the other. Thus, the inclusion of
living and nonliving objects in ecosystems has practical as well as intellectual advantages.
Finally, we note one further property of ecosystems: they are open to the flow of energy
and materials. It might be theoretically possible to define particular examples of ecosystems
that are closed systems, not exchanging energy or materials with their surroundings, but
nearly all ecosystems as actually defined have important exchanges of energy and materials
with their surroundings. Indeed, such exchanges are one of the central subjects of ecosystem
science. We note in particular that most ecosystems depend on energy inputs from external
sources, either as energy from the sun or as organic matter brought in from neighboring
ecosystems.
Now consider briefly what is missing from the definition. We have already noted that the
definition does not specify the time or space scales over which an ecosystem is defined, or
where exactly the boundaries are placed. Ecosystems are not required to be self-regulating, per-
manent, stable, or sustainable. They are not required to have any particular functional prop-
erties. For example, they need not be in balance or efficient in the way that they process
materials. Our definition does not require ecosystems to have a purpose. Although ecosystems
change over time, the basic definition does not suggest anything about the nature or direction of
that change. It might seem like a shame not to include such interesting attributes in a definition
of ecosystem (O’Neill 2001), and indeed some ecologists have incorporated such attributes
in their definitions, but we think it is neither necessary nor helpful to include them in a basic
definition. They may, however, be useful hypotheses and the subject of fruitful research pro-
jects. For instance, we might hypothesize that as forest ecosystems recover from disturbances
like fire or clear-cutting, they retain a higher proportion of nutrient inputs from precipitation
or release from weathering substrates. This viewpoint is quite different than saying that
ecosystems are systems that tend to maximize efficiency of use of limiting nutrients.
I. Introduction
What Are the Properties of Ecosystems? 9
What Are the Properties of Ecosystems?

All systems have characteristic properties that allow us to describe them and compare
them with other similar systems (Box 1.1). How might we describe the properties of
ecosystems?
What Is in an Ecosystem?
We might begin simply by listing the contents of an ecosystem. Plants and animals occur in
most ecosystems. As we will see later in the book, the number and kinds of plants and animals
can have a strong influence on ecosystem function. Many ecosystems also contain people.
Historically, many ecologists treated humans as outside of the ecosystem, or deliberately
studied ecosystems without people, but it is now common to treat people and our institutions
as parts of ecosystems (e.g., Pickett et al. 2001, 2011; McPhearson et al. 2016). Certainly the
structure and function (and change) of many modern ecosystems cannot be understood with-
out considering human activities.
Almost all ecosystems contain microbes (bacteria and fungi); although these are not as con-
spicuous as plants and animals, their activities are vital to ecosystem functioning. Viruses oc-
cur in most ecosystems, and may play important roles as regulators of plant, animal, and
microbial populations. Ecosystems also contain water and air, which are themselves re-
sources for many organisms and also serve as media in which organisms and nonliving ma-
terials can be transported, both within and across the boundaries of ecosystems. Finally,
ecosystems contain an enormous variety of nonliving materials, organic and inorganic, solid
and dissolved. These nonliving materials, including such disparate items as dead wood, clay
particles, bedrock, oxygen, and dissolved nutrients, interact with the living biota and exercise
strong influences on the character and functioning of ecosystems. Thus, the total inventory of
an ecosystem can be very long; it might contain thousands or millions of kinds of items, living
and nonliving, and countless numbers of individuals in these “kinds.”
Ecosystems Have Structure

This complexity allows for an essentially infinite number of possible descriptions of eco-
system structure. Nevertheless, only a few descriptions of ecosystem structure are commonly
used by the scientists who study ecosystems. Often ecosystems are described by the numbers
and kinds of objects that they contain, focusing on key materials or organisms. Thus, we may
describe an ecosystem as having a plant biomass of 300 g/m2, or a deer population of 5/km2,
or a nitrogen content of 200 kg/ha. Sometimes ecosystem scientists describe ecosystems by
the ratios of key elements such as the nitrogen:phosphorus ratio of a lake ecosystem. If we
were interested in the role of biological communities in regulating ecosystem function, we
would refer to the biodiversity (especially the species richness) of the organisms in the eco-
system. We may be interested in the spatial variation, as well as the mean value, of any such
key variables (see Chapter 11). Thus, we may describe ecosystems as being highly patchy as
opposed to relatively homogeneous in nitrogen content or biodiversity. Finally, scientists
I. Introduction
often describe ecosystems by their size or location (e.g., latitude, altitude, biogeographic
realm, or distance from the coast).
Ecosystems Perform Functions

In the broadest sense, ecosystems consume energy and transform materials. As with all
systems subject to the second law of thermodynamics, some of the useful energy that comes
into ecosystems (crossing the ecosystem’s boundary) in forms such as solar radiation, chem-
ical energy (e.g., organic matter), or mechanical energy (e.g., wind) is degraded to heat and
becomes unable to perform further work. In particular, living organisms need a continual
source of energy to maintain biochemical and physiological integrity, as well as to perform
activities such as swimming, running, or flying. Curiously, although these biological energy
transformations are only a part of the energy transformations that occur in an ecosystem, most
studies of energy flow through ecosystems treat only forms of energy that can be captured
and used by living organisms (i.e., solar radiation and chemical energy), and ignore purely
abiotic processes such as the conversion of kinetic energy to heat by flowing water. Organisms
can capture solar energy or chemical energy from inorganic compounds (photosynthesis and
chemosynthesis, respectively), store energy, obtain energy from other organisms (e.g., preda-
tion), or convert energy into heat (respiration). Patterns of energy flow through ecosystems
can be of direct interest to humans who harvest wild populations, and can tell ecosystem sci-
entists a good deal about how different ecosystems function.
Ecosystems also transform materials in various ways. Materials that come into the ecosys-
tem may be taken up by some part of the ecosystem and accumulate. In some cases, this ac-
cumulation may be temporary so that the ecosystem acts as a sort of capacitor, releasing the
material at a later time. The lag time between atmospheric deposition of sulfate onto a terres-
trial ecosystem and its export in stream water from that system is an example. Ecosystems
may also be a source of material, releasing their internal stores to neighboring systems. Meth-
ane gas flux from a wetland to the atmosphere is an example. Finally, and perhaps most in-
teresting, ecosystems transform materials by changing their chemical and physical states
(Chapter 6). Nitric acid contained in rainwater falling on a forest soil may react with the soil
and form calcium nitrate in soil water. The nitrate in the solution may then be taken up by a
plant and incorporated into protein in a leaf. At the end of the growing season, the leaf may
fall into a stream where it is eaten by an insect and chopped into small leafy bits, which then
wash out of the ecosystem. The description of chemical and biological transformations by eco-
systems forms the field of biogeochemistry (e.g., Schlesinger and Bernhardt 2013; see
Chapter 7), which is a major part of modern ecosystem science (and this book). Many biogeo-
chemical functions are important to humans (e.g., the removal of nitrate by riparian forests in
the Mississippi River basin; see Chapter 19, Figure 19.2), as well as essential to understanding
how different ecosystems work.
Ecosystems often are described by their functions as well as their structures. One of the most
common functional descriptions of ecosystems is whether the system is a source or a sink of a
given material—that is, whether the inputs of that material to the ecosystem are less or more,
respectively, than the outputs of that material from the ecosystem. In the special case of energy
flow through ecosystems, the degree to which an ecosystem is a source or a sink is described by
I. Introduction
What Are the Properties of Ecosystems? 11
the P/R (gross photosynthesis to respiration) ratio for the system. At a steady state, ecosystems
with a P/R ratio less than 1 must import chemical energy (usually organic matter) from neigh-
boring ecosystems and are called heterotrophic; those with a P/R ratio greater than 1 export
chemical energy to neighboring ecosystems and are called autotrophic. Another useful
functional description is the residence time of a given material in an ecosystem—that is, the
average amount of time that a material spends within an ecosystem. Residence time is cal-
culated by dividing the standing stock of the material in the ecosystem by its input rate.
Ecosystem structures and functions can have economic value. For instance, ecosystems
provide lumber, they purify water and air, they regulate the prevalence of human diseases,
and they provide pollination for crop plants. These and many other goods and services
provided by ecosystems are commonly called “ecosystem services”—the benefits that people
derive from ecosystem structures and functions (e.g., Daily 1997; Millennium Ecosystem
Assessment 2005; Kareiva et al. 2011, Box 1.2). Developing ways to estimate quantitatively
the value of ecosystem services is an important and growing field at the intersection of
ecology, sustainability science, and economics.
Ecosystem Structure and Function Are Controlled by Many Factors

Unlike systems such as the solar system, the dynamics of which are controlled by just a few
factors, ecosystem structure and function depend on many factors. Ecosystem scientists have
learned much about how ecosystems are controlled, and much of the remainder of this book
will be concerned with this subject. Ecosystem structure and function often are affected by or-
ganisms (including humans), either through trophic activities such as herbivory, predation, and
decomposition, or through engineering activities ( Jones et al. 1994) such as burrowing, shelter
construction (e.g., beaver dams), and the like (see Box 12.1 in Chapter 12). Likewise, the
nonliving parts of ecosystems often control ecosystems by determining supplies and movement
of air, water, key nutrients, and other materials. Temperature is another abiotic factor that has
strong effects on ecosystems. Finally, because most ecosystems are open and exchange energy
and materials with the ecosystems that surround them or that preceded them, the structure and
function of an ecosystem can be strongly affected by its spatial and temporal context.
Ecosystems Change Through Time

Ecosystems change through time (see Chapters 11 and 12). These changes may be gradual
and subtle (the millennial releases of nutrients from a weathering soil) or fast and dramatic
(a fire sweeping through a forest). Both external forces (changes in climate or nutrient inputs)
and internal dynamics (aging of a tree population, accumulation or depletion of materials in a
soil or a lake) are important in driving temporal changes in ecosystems. In some cases,
changes are directional and predictable (e.g., soil weathering, the filling of a lake basin), while
in other cases changes may be idiosyncratic and difficult to predict (e.g., the arrival of an in-
vasive species, disturbance by a hurricane). Understanding and predicting how ecosystems
change through time is of great theoretical and practical interest, and is a major part of con-
temporary ecosystem science.
I. Introduction
How Do We Classify or Compare Ecosystems?

Thus, ecosystem scientists use structure, function, control, and temporal dynamics to clas-
sify and compare ecosystems. For instance, it is common to see ecosystems described as rich
in nitrogen (structure), sinks for carbon (function), fire-dominated (control), or recently dis-
turbed (dynamics). All of these attributes of ecosystems can provide useful frameworks to
classify ecosystems, and ultimately to organize and interpret the vast amount of information
that scientists have collected about ecosystems. Similar descriptions and classifications are
evident throughout the book.
Why Do Scientists Study Ecosystems?
Scientists have been motivated to study ecosystems for several reasons. To begin with, if
ecosystems truly are the “basic units of nature” on Earth (Tansley 1935), any attempt to
understand our planet and the products of evolution on it must include ecosystem science
as a central theme. Indeed much study of ecosystem science has been motivated by simple
curiosity about how our world and how systems—whether ecological, social, or socio-
ecological—work. Many salable products such as timber and fish are taken directly from
“wild” ecosystems, so many early ecosystem studies were carried out to try to understand
better the processes that supported these products and ultimately increase their yields. Espe-
cially in the past 30 years, we have come to realize that the valuable products of nature include
far more than obviously salable products like timber and fish. Ecosystems also provide us
with clean air and water, opportunities for recreation and spiritual fulfillment, protection
from diseases, and many more “ecosystem services” (Box 1.2). Human economies and
well-being are wholly embedded in and dependent on wild ecosystems. Thus, many contem-
porary ecosystem studies are concerned with how ecosystems provide this broad array of
services, how human activities reduce or restore the ability of ecosystems to provide these
services, and ultimately how to reconcile the growing demands of human populations with
the needs of both nature and ourselves for functioning ecosystems.
How Do Ecosystem Scientists Learn about Ecosystems?
Depending on the problem that they are studying, ecosystem scientists use a wide variety
of approaches and an array of simple to sophisticated tools to measure different aspects of
ecosystem structure and function. We offer a few examples here; however, new approaches
and tools emerge every year, and with them come more ways to open black boxes in ecosys-
tem science (see Chapter 19).
Approaches for Learning about Ecosystems

There are multiple approaches by which scientists can understand ecosystem structure,
function, and development, both qualitatively and quantitatively. Five approaches (modified
from the lists of Likens 1992; Carpenter 1998) are especially important in ecosystem science,
I. Introduction
How Do Ecosystem Scientists Learn about Ecosystems? 13
TABLE 1.2 Strengths and limitations of approaches to understanding ecosystems. Natural history observations
and understanding underpin all of these approaches.
Approach Some Strengths Some Limitations
Theory Flexibility of scale; integration; deduction of Cannot develop without linkage to
testable ideas observation and experiment
Long-term observation Temporal context; detection of trends and Potentially site-specific; difficult to
surprises; test hypotheses about temporal determine cause; costly and difficult to
variation maintain
Comparison Spatial or interecosystem context; detection of Difficult to predict temporal change or
spatial pattern; test hypotheses about spatial response to perturbation
variation
Ecosystem experiment Measure ecosystem response to perturbation; Potentially site-specific; potentially
test hypothesis about controls and difficult to rule out some explanations;
management of ecosystem processes hard to do; expensive
After Carpenter (1998).
including: (1) natural history or observation; (2) theory and conceptual models; (3) long-term
study; (4) cross-ecosystem comparison; and (5) experiments. These approaches are comple-
mentary to one another (Table 1.2), and are best used in combination. Almost every scientific
question of any complexity or importance in ecosystem science requires the use of two or
more of these approaches to get a robust answer.
Natural History
A good deal can be learned about ecosystems simply from watching them and
documenting what is observed in some fashion. Do fallen leaves decay in place, wash away
into a stream, or burn in episodic fires? Is the soil deep and rich, or shallow and rocky? Does it
freeze in the winter? As a result, our understanding of an ecosystem often is based on simple
observations of its natural history. Indeed, without such careful observations, even the most
sophisticated studies can go astray by formulating nonsensical questions or omitting key ob-
servations or measurements. Not surprisingly, careful natural history studies (such as Forbes’
“The Lake as a Microcosm,” discussed later) were important precursors to modern ecosystem
science. Although these forerunners of ecosystem science often included speculation about
ecosystem processes, they did not have the technical means to measure easily such functions
as net ecosystem productivity or nutrient cycling, or to quantify trophic transfers.
Long-Term Studies
Long-term studies (i.e., those lasting for more than 10 years—longer than the tenure of
most grants or the time it takes to earn a PhD!) are relatively rare in ecology as a whole
(Lindenmayer and Likens 2018). However, long-term studies are especially good at providing
insight into slow processes (e.g., changes associated with forest succession), subtle changes
(e.g., changing chemistry of precipitation), rare events (e.g., the coldest winter, effects of hur-
ricanes or insect outbreaks), or processes controlled by multiple interacting factors (e.g., fish
I. Introduction
recruitment; Likens 1989; Lindenmayer and Likens 2018; and see the Long-Term Ecological
Research Program (LTER) of the National Science Foundation, http://www.lternet.edu).
Sometimes long-term understanding can be obtained by short-term analyses of materials that
record history, such as soil or sediment cores, otoliths (fish ear-stones), or written historical
records. For example, analysis of pollen, diatoms, pigments, or geochemistry in lake sediment
can reveal the millennium-long history of terrestrial vegetation, phytoplankton, soils, and
lake level—in short, the history of the development of the linkages between terrestrial and
aquatic ecosystems. It is from long-term studies or their surrogates that scientists have
documented climatic, atmospheric, geochemical, and organismal changes over decades to
billions of years.
Cross-Ecosystem Comparison
Comparative studies have served two important roles in ecosystem science. Most simply,
scientists often have measured some variable associated with ecosystem structure or function
across a series of ecosystems to identify typical values of that variable, show how it varies
among types of ecosystems, and generate hypotheses about what factors might control that
variable. An example of such an analysis is shown in Table 2.1 in Chapter 2. Alternatively,
scientists often test whether some factor controls an ecosystem by comparing ecosystems that
differ in that factor and not (to the extent possible) in any other relevant characteristic. For
instance, if we wanted to test whether phosphorus inputs control primary production in
lakes, we might try to measure primary production in a series of 10 lakes of similar size,
depth, species composition, and terrain that differ in their phosphorus inputs. In practice,
it often is difficult to find such a perfect series of well-matched ecosystems that could serve
as study sites.
Experiments
Experiments, whether conducted in the laboratory or in the field, are powerful ways to re-
veal controls on ecosystem structure and function (Likens 1985; Weathers et al. 2016). There are
no rules about the size of experimental units: manipulations have been made across hundreds
of square kilometers (e.g., iron fertilization experiments conducted in the ocean) and within
square centimeters. Often, the goal of experiments is to measure an ecosystem’s response to
a change in a single variable while holding all others as constant as possible. For example,
to understand whether phytoplankton in lakes were controlled by phosphorus or by other nu-
trients such as nitrogen and carbon, scientists in the Experimental Lakes Area of Canada
added phosphorus, nitrogen, and carbon to one-half of a lake (cut in two by a massive
curtain) and just nitrogen and carbon to the other half. They then compared responses—such
as the amount of primary production—in each half of the lake to see the effects of the
treatments. This whole-lake experiment helped to demonstrate that phosphorus was a major
factor controlling algal productivity in the lakes being tested (but see Chapter 9).
Theory and Conceptual Models

As in other sciences, ecosystem scientists routinely use theory and conceptual models.
Such theories and models are highly varied in structure and purpose (Canham et al. 2003;
Pickett et al. 2007; Childers et al. 2014). Models may be as simple as a statistical regression
(see Chapter 12) or a box-and-arrow diagram drawn on a napkin, or as complex as a detailed
I. Introduction
Land
management,
natural
disturbance Atmospheric Deposition
Cations and Anions
Atmospheric CO2
SUBLIMATION
TRANSLOCATION Snowpack
CO2
N2O LITTERFALL
NOx ET INFILTRATION
RUNOFF
PHREEQC soil reactions
UPTAKE SOM DECOMP.
MINERALIZATION Cation Exchange
Soil Organic Aqueous reactions
C,N,P,S NO3– NITRIF./DENITRIF.
Mineral denudation
NH4+ CO2 CO2 dissolution
DOC, Cations, Anions, CEC
LEACHING ANC, pH, BC, Cl, Al, SO42–
Aquifer
PHREEQC stream reactions
BASEFLOW Aqueous reactions

Mineral denudation
CO2 dissolution
Stream Flow
CO2
FIGURE 1.3 DayCent-Chem model processes. DayCent-Chem was developed to address ecosystem responses to
combined atmospheric nitrogen and sulfur deposition. DayCent-Chem operates on a daily time step and computes
atmospheric deposition, soil water fluxes, snowpack and stream dynamics, plant production and uptake, soil organic
matter decomposition, mineralization, nitrification, and denitrification (left side of figure) while utilizing PHREEQC’s
(an aqueous geochemical equilibrium model) low-temperature aqueous geochemical equilibrium calculations, in-
cluding CO2 dissolution, mineral denudation, and cation exchange, to compute soil water and stream chemistry (right
side of figure), ANC, acid neutralizing capacity; BC, base cations (Ca, Mg, K, Na); CEC, cation exchange capacity; DOC,
dissolved organic carbon; and evapotranspiration; ET, evapotranspiration. The model requires considerable site-
specific environmental data to run. (From Hartman et al. 2009, Figure 1.3.)
simulation model (Figure 1.3). Models are highly flexible, can cover scales of time and space
that are difficult to study using other approaches, and often can provide quick answers at low
cost. They also are very useful as a way to organize facts and ideas, to generate, sharpen, or
narrow hypotheses, and to guide research activities. Scientists often make rapid progress by
tightly coupling theory and models to other approaches.
What Do Ecosystem Scientists Measure?

Ecosystem scientists are inherently interested in the connections between structure and
function of ecosystems, and how these develop over time. Thus, many of the examples of
measurements or values in this book are related to structure and function, such as biomass
I. Introduction
of a species, or rates of carbon cycling. They are what is often found on the x or y axes of
graphs, or are used as treatments or are measured as responses in experiments. Ecosystem
structure is sometimes measured by variables such as leaf area index or the number of trophic
levels in a lake (see Chapter 11). Productivity (Chapters 2 and 3), rates of decomposition
(Chapter 4) or mineralization (Chapter 7) over time or space, or the accumulation of some
element of interest can be indicators of ecosystem function. Ecosystem development is often
described by changes in structure, function, and their relationship over time (e.g., linked
changes in soil and vegetation over millennia).
Many, if not most, of our measurements of ecosystem function are indirect. Sometimes we
can measure function directly, such as gas exchange, but these measurements are almost in-
evitably made on a tiny fraction of the ecosystem (e.g., individual leaves within a grassland or
bottle of water from a lake). To estimate a flux over a larger area of a grassland, for example,
an ecosystem scientist might deploy eddy covariance instruments that measure carbon diox-
ide, water, temperature, and wind speed and direction continuously at a place within the
grassland (see Chapter 2). From these measures, a model can be used to infer carbon dioxide
flux into or out of the ecosystem.
Scientists often choose indirect measures because they are easier to obtain across larger
parts of a system or across more systems. As another example, the measurement of
chlorophyll-a is often used as an indicator of primary productivity in aquatic ecosystems.
However, chlorophyll-a is not a direct measure of productivity, but a measure of the presence
of a pigment used in photosynthesis, and the photosynthetic process builds biomass. Like-
wise, the carbon:nitrogen (C:N) ratio in soil is often used as an indicator of litter or soil quality,
and is used to predict decomposition rates, or rates of nitrogen cycling (see Chapters 4 and 8).
To make these indirect measures useful, empirical relationships between direct and
surrogate measures must be established—quantifying these relationships is an active area
of research.
Some Tools in the Ecosystem Scientist’s Toolbox

Ecosystem scientists try to answer a diverse range of questions about a wide array of char-
acteristics of the most varied kinds of ecosystems, using many scientific approaches. It will
therefore come as no surprise that ecosystem scientists use an enormous number of specific
scientific techniques in their investigations, some simple, some sophisticated, some devel-
oped within the discipline, and some borrowed and adapted from other disciplines. These
techniques are far too numerous to list and discuss here. Nonetheless, several tools are worth
introducing because they are characteristic of ecosystem science and will appear repeatedly in
the coming chapters; some exciting, emerging tools are noted in the last chapter.
Balances: Mass and Charge

Mass balance (Box 1.3) is a major tool in ecosystem science, especially for ecosystems in
which the boundaries are defined by their watersheds. Mass balances are how all ecosystems
are connected: outputs from one system are inputs for another. The First Law of Thermody-
namics tells us that matter and energy are not created or destroyed. When both inputs and
outputs of energy or matter can be measured relatively completely and accurately, it is
I. Introduction
possible to construct a mass balance and infer processes. For example, a watershed analysis
that does not balance (quantitatively) suggests that either the element of interest is being
retained in (inputs > outputs) or leaking from (outputs > inputs) the ecosystem (see
Chapters 6 and 10). The watershed mass balance approach was pioneered in the 1960s by
BOX 1.3
Mass Balance
To see just how useful the tool of mass bal- that was bounded by the lake shore, the over-
ance can be, suppose we are trying to evalu- lying air, and the bedrock deep beneath the
ate whether a lake ecosystem is taking up or lake sediments. Using mass balance, we note
releasing phosphorus. We could try to mea- that the amount of phosphorus being retained
sure all the exchanges between parts of the by the lake ecosystem is simply the amount of
ecosystem (e.g., the uptake of phosphorus phosphorus going into the lake minus the
by phytoplankton and rooted plants; the con- amount that is leaving the lake. Now we just
sumption and excretion of phosphorus by the have to measure the exchanges across the eco-
animals that eat phytoplankton and plants; system boundary (stream water and ground-
the release of phosphorus during the decay water going into and out of the lake; rain,
of phytoplankton, plants, and animals; and snow, and particles falling on the lake; and
dozens of other exchanges), then simply any animals entering and leaving the lake—
sum up all of these measurements. It would hard enough!) to calculate whether the lake is
take an enormous amount of work to mea- taking up or releasing phosphorus. In the case
sure all the exchanges, and our final answer of Mirror Lake, New Hampshire (Figure 1.4),
would be fraught with large uncertainties. Al- almost 40% of incoming phosphorus is
ternatively, we could define a lake ecosystem retained by the lake.
Precipitation FIGURE 1.4 Average phosphorus inputs and

3.7 outputs in kilograms/year to Mirror Lake, NH. To-
t tal average inputs ¼ 6.7 kg/year; total average
Tr
le
ib 1
in outputs ¼ 4.1 kg/year. Inputs outputs ¼ 2.7 kg/

ut .2
y
ar 1
ar
t year or 39.7% retention of phosphorus in the lake.

y
u .
in
ib 0
Tr (Data from Winter and Likens 2009.)
le
t
Tributary inlet
1.4
Outflow
Ground water in 1.7
0.3
Ground water out

2.4
I. Introduction
scientists at the Hubbard Brook Experimental Forest, New Hampshire (Bormann and Likens
1967; Likens 2013), and has been used powerfully around the world to understand the abiotic
and biotic movement of a suite of elements through ecosystems.
The other powerful “balance” tool that ecosystem scientists use is charge balance. In water,
the charges held by positive ions (cations) and negative ions (anions) must balance each other.
That is, for every anion (such as chloride) in an aqueous solution, there must be a
corresponding cation (such as sodium). Why is this tool so useful? Charge balance tells us,
for example, that when an anion moves through a forest soil from groundwater into a stream,
it must be accompanied by a corresponding cation (see Chapter 6). The sum of all the negative
charges brought by anions must be balanced by the same number of positive charges from
cations. Charge balance also makes it possible to check whether the major ions in a water sam-
ple have been measured correctly; a charge imbalance tells us that a measurement error has
been made or that we have not quantified all the cations or anions that are important in a
sample.
Tracers
As useful as balances are as tools, they tell us about the bulk (or net) movements of mate-
rials through ecosystems, and rarely allow us to distinguish among different pathways of ma-
terial movement within ecosystems. All nitrogen atoms look alike to a mass balance. Tracers
are tools that allow ecosystem scientists to distinguish among particular pathways of material
movement by labeling just some of the atoms or molecules of interest. Ecosystem scientists
have used several tracer methods, which have been enormously powerful in understanding
how ecosystems work.
Radioisotopes (Box 1.4) were some of the first tracers used in ecosystem science. Radioiso-
topes can be detected and quantified at very low concentrations, so they make excellent
tracers, and have had many applications in ecosystem science. In the mid-20th century, eco-
system scientists added small amounts of radioisotopes to ecosystems to trace the movement
of water and the uptake and movement of carbon and limiting nutrients, such as phosphorus,
through ecosystems. Radioisotopes are no longer added to natural ecosystems as tracers be-
cause of associated health risks, but they continue to be used widely in laboratory studies and
measurements (e.g., to measure microbial production; see Chapter 3). They also are used in
“natural abundance” studies where ecosystem scientists use the very low natural abundance
of radioisotopes to trace the movement of materials through ecosystems, rather than adding
radioisotopes to ecosystems. For example, Caraco and her colleagues (2010) observed that the
concentration of 14C in organic matter that washed into the Hudson River from the soils of its
watershed was very different than that of organic matter produced by photosynthesis within
the river. They could therefore use 14C to trace movement of terrestrial organic matter through
the Hudson River food web, and show that modern zooplankton were being supported in
part by carbon that was captured by primary production thousands of years ago (Figure 1.5).
Stable isotopes have largely taken the place of radioisotopes as tracers outside the labora-
tory (Box 1.4). Although much more difficult to measure and often expensive to use, stable
isotopes do not present a health risk to humans and wildlife. Stable isotopes are available
for many elements of ecological interest, including hydrogen, nitrogen, carbon, oxygen,
and sulfur. Stable isotopes often are added to ecosystems (or to laboratory experiments)
and followed as they move through the system. For example, Templer and her colleagues
I. Introduction
BOX 1.4
Ecological Tracers: Isotopes

34
Most elements exist in several forms that S, although many other isotopes have been
contain different numbers of neutrons (but used in specialized studies.
the same number of protons and electrons, The concentration of stable isotopes is
and basically the same chemical properties). usually expressed in a “del” (δ) notation that
For example, about 99% of the carbon on compares the abundance of the heavier iso-
Earth is 12C, which contains six protons, six tope to that of the lighter isotope.
electrons, and six neutrons, but about 1% of Thus, the abundance of 13C in a sample is
the carbon is 13C, which contains seven neu- expressed as:
trons. A tiny amount (0.0000000001%) of the 13 !
carbon is 14C, which has eight neutrons. Some C 12 C sample
δ Cð%Þ ¼ 13 12
13
1 1000
isotopes are stable, while others are radioac- C C standard
tive (i.e., they spontaneously decay into other
elements or isotopes). In the case of carbon, The standard in this case is Vienna Pee
12
C and 13C are stable isotopes, whereas 14C Dee Belemnite (a particular kind of fossil).
is a radioisotope that decays into nitrogen Negative δ values indicate that the heavier
(14N) with a half-life of 5730 years. Some iso- isotope is less abundant in the sample
topes that commonly make an appearance in than in the standard, while positive δ values
ecosystem science include the radioisotopes indicate that the heavier isotope is more
3
H (tritium), 14C, 32P, and 35S, and the stable abundant in the sample than in the
isotopes 2H (deuterium), 13C, 15N, 18O, and standard.
(2005) added a stable isotope of nitrogen, 15N, to forest plots in the Catskill Mountains, NY,
and then followed it into soil, microbial biomass, understory plants, tree roots, wood, and
leaves, and found that most of the nitrogen stayed in the soil. Alternatively, ecosystem scien-
tists often use natural abundance studies of stable isotopes to follow the movement of mate-
rials through ecosystems.
Substances other than isotopes can be used as tracers as well. For instance, certain fatty
acids cannot be synthesized by animals and are made only by particular kinds of algae. By
analyzing the fatty acid content of zooplankton and fish, the contribution of different kinds
of algae throughout the food web can be traced. Caffeine, which is not readily degraded in
conventional sewage treatment plants, is sometimes used as a tracer for sewage. The kinds
of substances that can be used as tracers are highly varied, limited only by the ingenuity
and analytical capabilities of the investigator.
Spatial Data
Where are the regions of high and low productivity around the globe? How do they change
over the seasons? These are questions that can now be answered largely as a result of the
availability of remote sensing tools and spatially explicit data. The ability to collect, represent,
and analyze spatially explicit data has risen exponentially over the past decade (Weathers
I. Introduction
100
Modern
Modern FIAV FIAV
terrestrial
terrestrial
0
SAV
SAV
Phytoplankton Phytoplankton
–100
δ14C (‰)
–200
Zooplankton Zooplankton
–300
Aged Aged
terrestrial terrestrial
–400
–35 –30 –25 –20 –250 –200 –150 –100
δ13C (‰) δ2H (‰)
FIGURE 1.5 Use of stable and radioisotopes to determine the source of organic matter supporting zooplankton in
the Hudson River. Isotope bi-plots show 14C vs. 13C (left side) and 14C vs. 2H (right side). Sources of carbon from modern
primary production are shown near the tops of the graphs (FlAV, floating-leaved aquatic vegetation; SAV, submersed
aquatic vegetation). If zooplankton were composed of carbon and hydrogen from these sources, then the data for
isotopic composition of zooplankton would fall in the same region of the graph as the sources. Instead, zooplankton
fall far outside this region of the graph, showing that they must be composed of organic matter from both modern and
“aged” sources (i.e., organic matter thousands of years old from the soils of the Hudson River’s watershed). (From
Caraco et al. 2010.)
et al. 2016). Remote sensing and the georeferencing of basic data on landscape characteristics
such as elevation, water bodies, land cover, and geological materials have opened the door to
a description of ecosystem structure over large areas (see Holmes and Likens 2016; Boucher
et al., 2018). Geographic information systems (GISs) allow analysis of the relationships be-
tween these structures and fluxes in or out of these systems. For example, the variation in
atmospheric deposition across the mountainous terrain of Acadia National Park or Great
Smoky Mountains National Park can be described by a GIS model that links empirical mea-
surements to landscape features that are described in the GIS (Figure 1.6). Such spatially ex-
plicit models greatly enhance our ability to identify places on the landscape and times that
may be subject to particularly high levels of atmospheric deposition (Weathers et al. 2006).
GIS and other technologies are being used creatively and hold tremendous potential for un-
derstanding ecosystem processes across heterogeneous landscapes.
I. Introduction
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sonado allí. Óyelo el de las gafas, y, clavándolas en usted, le
pregunta, con una voz muy dulce y una cara muy risueña:
—¿Verduguillos ha dicho usted, caballero?
—Verduguillos, sí señor,—responde usted parándose en firme.
—¿Sabe usted—insiste el otro,—(y usted perdone si le interrumpo
un momento), si ese señor de Verduguillos tiene parientes en
Cuzcurrita de Río Tirón?
—¿Por qué he de saber yo eso, si jamás allá estuve, ni conozco á
ese señor más que de vista?—replica usted con el sosiego y la
amabilidad que eran de esperarse.
—Perdone usted, caballero—dice el intruso hecho unas mieles,—y
verá por qué me he tomado la libertad de interrumpirle.
Y en esto, deja la silla, sale al centro, encárase con el grupo
principal, afirma las gafas en el entrecejo, carraspea, sonríese y
dice:
—Pues, señor, verán ustedes por qué me ha interesado tanto el oir
á este caballero nombrar á ese señor de Verduguillos. Por el mes de
septiembre del año treinta y ocho, salí yo de Zamora (donde nací y
me crié y radican los pocos ó muchos bienes que heredé á la
muerte de mis padres, y los que he podido adquirir después acá con
el fruto de mis especulaciones modestas), con el propósito de hacer
un largo viaje, por exigirlo así los asuntos de la familia, y también, si
he de ser franco, el estado de mi salud...
Así comienza este señor la relación de un viaje por media España,
con largas detenciones en todos los puertos y plazas del tránsito, y
minuciosas observaciones estadísticas y climatéricas, sin pizca de
interés, ni método, ni estilo, ni substancia, hasta venir á parar, al
cabo de tres mortales cuartos de hora, á Logroño, en la cual ciudad
conocía al comerciante don Fulano de Tal; y decirnos que, yendo á
visitarle á su escritorio, hallóse allí con un caballero, muy amigo
también del don Fulano, el cual don Fulano le dijo á él al despedirse
el otro:
—Este señor que acaba de salir, es don Pacomio Verduguillos,
natural y vecino de Cuzcurrita de Río Tirón.
Al llegar aquí con el cuento el de las gafas, espera usted el toque de
efecto, el desenlace sorprendente, la gracia del suceso; porque es
de saberse que el narrador se ha quedado en silencio y mirando de
hito en hito á los resignados oyentes. Pero el silencio sigue y la
sorpresa no asoma. Alguien se aventura, y pregunta al del bastón:
—Pero ¿por qué le chocó á usted tanto el oir nombrar á este
Verduguillos?
—Hombre—responde el interpelado, con candidez angelical,—
porque podía muy bien ser pariente del otro Verduguillos que yo
conocí en Logroño.
¡Y para eso interrumpió un animado y sabrosísimo debate; y estuvo,
durante cerca de una hora, ensartando insulsez tras de insulsez,
simpleza tras de simpleza, adormeciendo á unos, quemando la
sangre á otros y aburriéndolos á todos! Y usted llevó la cruz con
paciencia, y yo también; y lo mismo al día siguiente, porque el
bueno del zamorano, desde que pierde la cortedad con el primer
relato, ya no cierra boca en la tertulia, y siempre tan ameno,
divertido y oportuno. Pero nos permitimos los dos un desahoguillo
en un aparte.
—Amigo—dije, ó me dijo usted,—¡este hombre es insufrible:
estando él no se puede venir aquí! Y se oyó el rumor del desahogo,
y ¡qué caras nos pusieron los señores tolerantes, que estaban tan
aburridos como nosotros!
Al día siguiente asoma usted la cabeza á la puerta, ve al de las
gafas en el uso de la palabra, retrocede y no vuelve; ni yo tampoco.
Y porque no volvemos, y además decimos lo que mejor nos parece
del motivo, ¡qué ponernos de intolerantes y hasta de inciviles!...
¡Caramba, protesto contra la enormidad de esta injusticia! En este
caso no hay más intolerantes que el señor de Zamora, que
interrumpe toda conversación racional y obliga á hombres de buen
sentido á que oigan las interminables boberías que él enjareta sin
punto de reposo, y los forzados tolerantes que le escuchan con
paciencia, y no la tienen para oir que otros carecen de ella.
Trátase ahora de un embustero, que un día y otro día le abruma á
usted con narraciones autobiográficas, sin principio ni fin, como la
eternidad de Dios; pero muy punteadas, muy comeadas y con más
espacios que un libro de malos versos. Oye usted una historia, y
dos, y tres, ya con mala cara; pero, al fin, se acaba la paciencia, y
un día interrumpe usted al sujeto de los á propósitos, y le dice:
—Mire usted, hombre: en primer lugar, la mayor parte de lo que
usted me cuenta se lo he contado yo á usted en cuatro palabras; en
segundo lugar, le sucedió á un condiscípulo mío en Oviedo, y no á
un amigo de usted en Zaragoza; en tercer lugar, no pasó como
usted lo refiere, sino del modo contrario: mi condiscípulo no adquirió
una capa aquella noche, sino que perdió la que llevaba, y, además,
el juicio, con costas, á los pocos días...
—Pues lo mismo da...
—Justo: media vuelta á la derecha es lo mismo que media vuelta á
la izquierda, sólo que es todo lo contrario.
—¡Caramba, es usted lo más intolerante!... No se puede hablar con
usted...
¡Todavía le parece poco, al ángel de Dios, la tolerancia que se ha
tenido con él!
Media docena de mujeres, ó menos, si á usted le parecen muchas
seis, se pasan una tarde entera desollando con la lengua al lucero
del alba. ¡Eso sí, con las mejores formas y la intención más santa!
De una dirán que es un dolor que, siendo tan bonita, sea tan charra
en el vestir, tan tosca en el hablar, tan inconsecuente en sus
amistades, tan desleal en sus amores; de otra, que es mordaz y
maldiciente, en lo cual se perjudica mucho, porque teniendo esta
falta, y la otra, y la de más allá, da pie para que cualquiera que se
estime en tan poco como ella, se las saque á relucir; de otra, que es
una desgraciada, porque el marido la ha puesto á ración, así en el
vestir como en el bailar, á causa de que fué algo despilfarrada
siempre en estos dos ramos de buena sociedad; de otra, que ya no
halla modista que la haga un traje si no paga adelantadas las
hechuras, y que no le venden nada en las tiendas, sino con el dinero
en la mano, etc., etc., etc... En esto, entra usted (es un suponer) y,
continuando el desuello, llegan á preguntarle si conoce á cierta
señora de éstas ó las otras señas; y como la tal es mujer de historia,
y usted la sabe de corrido, repítela allí con comentarios, creyendo
hacer á su auditorio un señalado servicio. Yo creo también que
usted se le hace, pues no fué á humo de pajas la preguntita; pero es
lo cierto que todas aquellas señoras, después de oirle á usted,
exclaman, con el más sincero de los asombros:
—¡Jesús!... Con razón dicen que es usted temible.
—¡Yo temible, señoras mías?—responde usted.—¿Y por qué?
—¡Porque es usted lo más intolerante y lo más!...
¡Vaya usted á convencer á aquellas damas de que viven
constantemente encenagadas en el pecado que á usted le cuelgan!
No hay inconveniente en que, abandonando estos tiquis-miquis que
ocurren en el ordinario trato social, dirijamos el anteojo unos grados
más arriba.
Todos los días halla usted en periódicos, en folletos y en libros,
sátiras, burlas y disertaciones en serio contra ideas, sentimientos y
hasta personas muy de la devoción de usted. Ocúrresele mirar al
campo de donde parten tantos proyectiles, y le ve usted sembrado
de ridiculeces, farsas y toda clase de miserias; saca usted al palo
media docena de ellas, por vía de muestra, en un papel, en un
folleto ó en un libro; y ¡Virgen María, cómo le ponen á usted de
intolerante y de mordaz, los mismos que tienen la mordacidad y la
intolerancia por oficio!
Así andan, amigo, las cosas de justicia en el ordinario comercio de
las gentes; así se ataja al más inofensivo en el trayecto social en
que pasea su nombre, y así se pretende conducirle al extremo á que
no llegan en el mundo más que las bestias... y los que tienen la
manía de la tolerancia (siendo lógicos en ella): á ver, oir y callar... es
decir, á matar la sed con petróleo, allí donde haya un extravagante
que tal haga delante de usted.
Usted es hombre de sencillas y ordenadas costumbres (es también
un suponer): ni el mundo le tira, ni sus pompas y algaradas le
seducen. Éstos son gustos lícitos y racionales. Ajustándose á ellos,
en paz y en gracia de Dios, se da usted con un baile en los ojos:
tuerce usted el camino; tropieza usted más allá con una mascarada
de calaveras del gran mundo: echa usted por otro lado; allí topa
usted con la misma gente haciendo cuadros plásticos y animados
acertijos: cambia usted de rumbo; aquí asaltos, en el otro lado
conciertos... pues á la otra acera. Ni usted apedrea á los que bailan,
ni apostrofa á los que jiran, ni se ríe de los que se descoyuntan para
remedar á Cristo en la agonía, ni silba á los que reciben una
sorpresa, anunciada quince días antes, ni influye con el Gobernador
para que meta en la cárcel á toda esa gente: limítase á huir de lo
que le aburre, y á hacer lo que más le divierte ó menos le incomoda.
No haría otra cosa un santo.
Pero es el caso que los señores tolerantes no se conforman con
esto, y quieren que les diga usted por qué no concurre á los bailes, y
á las jiras, y á los cuadros vivos, y á los asaltos... y aquí está el
intríngulis precisamente; y si estos rasguños que trazo no fueran,
como he dicho, un inocente desahogo entre nosotros dos, y en
reserva, me atrevería á llamar la atención del lector hacia el
aparente fenómeno, cuya explicación es sencillísima, por lo cual, no
es fenómeno, aunque por tal le toman algunos.
Cuando á usted se le pregunta por qué no piensa como su vecino
sobre determinados puntos de transcendencia, á buen seguro que
se le ocurra á nadie que oiga la respuesta, agarrarse á ella para
llamarle á usted intolerante; pero que se le pregunte por qué no
baila, por qué no jira, etc., etc... y no bien ha contestado usted, ya
tiene encima el Inri de la intolerancia. Y ¿por qué en este caso y en
el otro no? Porque no está el intríngulis en la persona, ni en sus
razones, ni en el modo de exponerlas, sino en la cosa de que se
trata, que, muy á menudo, es, de por sí, ridícula, ó impertinente, ó
pueril cuando menos, y no resiste, sin deshacerse entre las manos,
el análisis de un hombre de seso; al cual hombre, no pudiendo
replicársele en buena justicia, en venganza se le pone un mote.
Por eso llevan el de intolerantes tantos caracteres dóciles, y creen
poner una pica en Flandes, y hasta se llaman guapos chicos y
excelentes sujetos en la sociedad, los que en ella entran con todas,
como la romana del diablo, menos con el sentido común. Quod erat
demonstrandum.
Á pesar de ello, y aun de la mucha saliva, que al propio asunto
hemos consagrado en nuestras conversaciones verbales, júzgole
apenas desflorado. ¡Cuánto me queda todavía que oir de los
inofensivos labios de usted!
Entre tanto, y dicho lo dicho, despidámonos por hoy, con la íntima
satisfacción, bien añeja en nosotros, de haber pasado juntos, en
espíritu, un agradable rato, sin murmurar de nadie ni ofender al
prójimo con hechos, con dichos ni con deseos.
1880.
EL CERVANTISMO
El Diccionario de la Academia no contiene este vocablo; pero es uno

de los propuestos por el último de los individuos del insigne cuerpo
literario para la edición que está imprimiéndose. Por si la Academia
no le acepta, conste que entiendo yo por
Cervantismo: La manía de los cervantistas; y por
Cervantista: El admirador de Cervantes, y el que se dedica á
ilustrar y comentar sus obras.
En rigor, pues, estos párrafos debieran haberse incluido entre los
que, bajo el rótulo de Manías, quedan algunas páginas atrás; pero
son tantos, y de tal índole la enfermedad á que se refieren, que bien
merecen vivir de cuenta propia y establecerse capítulo aparte.
Dice Chateaubriand, hablando de los españoles como soldados, que
nuestro empuje en el campo de batalla es irresistible; pero que nos
conformamos con arrojar al enemigo de sus posiciones, en las
cuales nos tendemos, con el cigarrillo en la boca y la guitarra en las
manos, á celebrar la victoria.
Si despojamos á esta pintura del colorido francés que la califica, nos
queda en ella un exactísimo retrato del carácter español, no sólo en
la guerra, sino en todas las imaginables situaciones de la vida.
Ya que no la guitarra, la pereza nacional nos absorbe los cinco
sentidos, y sólo cuando el hambre aprieta, ó la bambolla empuja, ó
la curiosidad nos mueve, sacudimos la modorra. Entonces
embestimos con el lucero del alba para estar donde él estuvo,
medrar de lo que medró y hacer todo cuanto él hizo. Pero de allí no
pasamos. Nuestra política, nuestra industria y nuestra literatura
contemporáneas lo declaran bien alto. Todo el mundo nos lleva la
delantera, y siempre estamos imitando á todo el mundo, menos en
andar solos y por delante; vivimos de sus desechos, y cada trapo
que le cogemos nos vuelve locos de entusiasmo, como si se hubiera
cortado para nosotros. Así estamos llenos de conquistas y de
«títulos á la admiración de las naciones extranjeras»; todos somos
ilustres estadistas, invictos guerreros, sabios hacendistas, insignes
literatos, laboriosos industriales y honrados obreros; hemos tenido
códigos á la francesa, códigos á la inglesa, códigos á la americana;
revoluciones de todos los matices, reacciones de todas castas,
triunfos de todos calibres, progresos de todos tamaños; y á la
presente fecha, el ciudadano que tiene camisa propia se cree muy
rico; la escasa industria desaparece antes que la Hacienda la
devore; los bufos imperan en el teatro; el hijo de Paul de Kock en la
novela; los Panchampla en desfiladeros y caminos reales, y la
navaja del honrado menestral desbandulla en las plazas públicas, á
la luz del mediodía, las víctimas á pares. De manera que quien nos
comprara por lo que decimos y nos vendiera por lo que hacemos,
buen pelo iba á echar con el negocio. Á hacer cosas nuevas y útiles
nos ganará cualquiera; pero á ponderar lo que hacemos no hay
quien nos eche la pata, ni á hacerlo mal y fuera de sazón, tampoco.
—Pero ¿qué tiene que ver todo esto con el cervantismo?—
preguntará el lector, oliéndole lo dicho á artículo de oposición más
que á otra cosa.
—No sé—respondo—por cuál de los lados encajará mejor en el
asunto prometido; pero lo cierto es que á las mientes se me ha
venido con él y como eslabón de la misma cadena de ideas. Acaso
en el cervantismo vea yo algo de la intemperancia, que, entre
nosotros, lleva en todo lo demás hasta el ridículo las cosas más
serias y respetables; quizá esa manía me ha hecho recordar la
tendencia española á perder en escarbar el huerto del vecino, el
tiempo que necesitamos para cultivar el propio; quizá me asaltó las
mientes el dicho de Chateaubriand pensando en los valientes que
conquistan el Quijote, y no pasan de allí, y allí se quedan,
rebuscando hasta las polillas, como si ya no hubiera otra cosa que
leer ni que estudiar en el mundo; acaso coinciden los dos asuntos
por el lado de la facilidad con que pasamos de la apatía al asombro,
de la indiferencia al entusiasmo, de la fiebre al delirio... ¡Quién
sabe? Pero el hecho existe, y ya no borro lo escrito, aunque el lector
me diga que soy uno de tantos como en España malgastan sin fruto
la hacienda, echando siempre los garbanzos fuera de la olla... Y
vamos al caso.
Y el caso es que ya estaba el mundo cansado de admirar á
Cervantes y de reproducir las ediciones del Quijote en todas las
lenguas que se hablan sobre la haz de la tierra, y aún eran muy
contadas en España las librerías en que se vendiera la obra inmortal
del ilustre soldado, que vivió de las limosnas de los próceres y fué
enterrado de caridad. Conocíanla los literatos, poseíanla los menos
de ellos, y veíase de vez en cuando en los mezquinos estantes de
algún particular, al lado de Bertoldo cuyos chistes saboreaba con
preferencia la patriarcal familia. Los nombres de don Quijote y
Sancho Panza eran populares; pero contadísimas las personas que
conocían á estos personajes más que de oídas: teníanlos unas por
históricos, las menos por novelescos; pero ni unas ni otras habían
oído jamás el nombre del padre que los engendró en su fantasía.
De pronto, ayer, como quien dice, alguien, y no español ciertamente,
nos aguija y nos apunta el Quijote con el dedo; sacudimos la
tradicional modorra, y allá vamos en tropel, y caemos como espeso
granizo sobre la obra señalada; las prensas gimen vomitando
ediciones populares del libro insigne, entre los cuadernos de Jaime
el Barbudo y Las cavernas del crimen, y aunque las masas de levita
siguen prefiriendo estas creaciones para solaz del espíritu, el
nombre de Cervantes suena en todas partes y á todas horas, y las
plumas y las lenguas ya no saben decir sino «el Cautivo de Argel» y
«el Manco de Lepanto».
¡Qué baraúnda! ¡Qué vocerío! Hay hombre, ya con canas, que
acaba de leer á saltos el Quijote, y se escandaliza de buena fe al
saber que un mozo imberbe no le conoce todavía; otro no le ha visto
ni por el forro, y mira con lástima á quien declara noblemente que no
ha podido adquirir un ejemplar para leerle... ¡Y cómo abunda esta
clase de admiradores!
—«Pero ¡qué hombre!... Pero ¡qué libro!... Pero ¡qué tiempos
aquéllos en que se morían de hambre tan preclaros ingenios! Como
esa obra no hay otra... El mundo la admira, y España no necesita
más que ella para su gloria... ¡Ah, Cervantes! ¡Ah, el Manco de
Lepanto!... ¡Ah, el Cautivo de Argel!».
Verdades como puños, enhorabuena; pero que tienen suma gracia
dichas por una generación, ya vieja, que no ha reparado en ellas
hasta que se las han metido por los ojos; y aun así no las ha visto
bien.
Y sigue el estrépito, y llena los ámbitos de la patria, y se conmueven
los poetas de circunstancias y los periodistas de afición y hasta los
filántropos de la usura; y allá van odas Al Manco de Lepanto, y
sonetos Al Cautivo de Argel, y llega á verse el nombre de Cervantes
en la popa de un falucho carbonero, y en el registro de una mina de
turba, y en los membretes de una sociedad anónima, y hasta en la
muestra de una zapatería; y hoy se celebra el aniversario de su
muerte, y mañana el de su nacimiento, y al otro día el de su
redención por los frailes trinitarios, y al otro, el de su casamiento; y
aquí brota una Academia cervantina, y allí un Semanario cervantino
y un Averiguador cervantesco; y en los unos y en los otros, y acá y
allá, no se trata sino de Cervantes y sus obras; y Cervantes aparece
en discursos, en gacetillas, en charadas, en rompe-cabezas y en
acertijos.
Lo que era de temer, sucede al cabo: la fiebre se propaga, hácese
peste asoladora, y no se libran de ella ni los que tienen el juicio más
aplomado; caen hasta los cervantistas de buena casta, y caen sobre
el Quijote y sobre la memoria de su autor, como antes cayera el
servum pecus, y allí se están cual si hubieran jurado, en el
paroxismo de su manía, gastar en la empresa hasta el último soplo
de la vida; porque cada cual cree encontrar en aquellas páginas
inmortales lo que más se acomoda á sus deseos y aficiones.
Imagínomelos yo como aquellos sabios resucitados de que nos
habla Balmes, husmeando el amplísimo establecimiento, y tráenme
á la memoria el caso de Mabillon despistojándose sobre un viejo
pergamino para descubrir algún renglón medio borrado, cuando
llega un naturalista y tira hacia sí del pergamino, para ver si halla en
él huevos de polilla.
Merced á estas faenas sobrehumanas, sabemos hoy, por otros
tantos señores cervantistas, cuyas plumas lo han afirmado en
sendos escritos, á cual más serio y pespunteado, que de las obras
de Cervantes resulta que fué éste sobresaliente
Teólogo,
Jurisperito,
Cocinero,
Marino,
Geógrafo,
Economista,
Médico,
Liberal (¡patriotero!)
Administrador militar (!!!!),
Protestante (¡¡¡!!!),
Viajero, etc., etc., etc.
Es decir, Cervantes omniscio, y sus obras la suma de los humanos
conocimientos.
Pero ni con todo esto, ni con más de otro tanto por el estilo, que no
hay para qué mentar, ni con el pintoresco catálogo de los
cervantómanos que han contado las veces que dice sí don Quijote,
ó Sancho vuesa merced, y otros admiradores de parecida ralea,
hemos llegado al delirium tremens de la enfermedad; puesto que
hay un español que ha dicho, y dice sin tregua ni descanso, porque
sospecho yo que por eso y para eso alienta y ha nacido:
—Caballeros, nada de lo que el mundo ha leído en el Quijote es la
obra de Cervantes.
Asombró el aserto, y preguntósele:
—Pues ¿qué otra cosa puede ser?
—Quiero decir—repuso el crítico,—que hasta ahora nadie ha sabido
leer el Quijote. No hay tal Dulcinea, ni tal Sancho Panza, ni tales
molinos, ni tales yangüeses, ni tal Ínsula Barataria, ni nada de lo que
allí aparece tal como suena. El Quijote, en suma, es una alegoría.
—¡Canastos! Y ¿quién se lo ha dicho á usted?
—Me lo han dicho treinta años de estudio incesante de esa obra
maravillosa, y lo demuestro en catorce volúmenes de comentarios,
que he escrito y tengo en casa esperando un editor que se atreva
con ellos.
—¡Tendrán que leer! Y diga usted, señor sabio, ¿qué especie de
alegoría es ésa que usted ha visto en el famoso libro?
—Es, como si dijéramos, el siglo xix hablando en profecía en el siglo
xvii; la luz de nuestras libertades columbrada por un ojo sutil, á tan
larga distancia; la protesta de un alma generosa contra la cadena de
la tiranía y las mazmorras de la Inquisición.
—¡Cáspita! Luego Cervantes...
—Cervantes fué un libre-pensador; un demócrata que nos precedió
cosa de tres siglos.
—Pero, hombre, aquellas declaraciones terminantes de neto y
fervoroso católico, que á cada instante hace; aquél su único
propósito, que jamás oculta, de escribir el Quijote para matar los
libros de caballerías...
—No hagan ustedes caso de ello. También dice (no lo niega al
menos) que lo de cabalgar Sancho en el Rucio después de
habérsele robado Ginés de Pasamonte, fué un lapsus de su
memoria, si no descuido del impresor, y, sin embargo, se le ha
demostrado todo lo contrario... Á Cervantes hay que saber leerle,
desengáñense ustedes.
—Corriente; pero ¿cómo teniendo ese hombre tanto talento no logró
hacerse entender de sus lectores?
—Porque temía á la Inquisición y al tirano.
—Callárase entonces, y ahorrárase el riesgo y la fatiga.
—No debía callar, porque había nacido para escribir.
—Pero no alegorías; pues, por las trazas, no le daba el naipe para
ellas.
—¡Cómo que no?
—Hombre, me parece á mí que una alegoría que no halla en cerca
de tres siglos más que un sabio que la desentrañe, no es cosa
mayor que digamos.
—¿Y qué son tres siglos en la vida de la humanidad?
—Trescientos años nada más; y aunque á usted le parezcan pocos,
pienso yo que, para desentrañar un libro, sobran de ellos casi todos,
aunque el libro esté en vascuence, cuanto más en neto castellano...
No se eche á broma el precedente diálogo, porque es la quinta
esencia de las polémicas sostenidas en la prensa, todos los días,
por el desenredador único de la supuesta maraña del Quijote, contra
los defensores del servum pecus, que no ha visto ni verá jamás en
las páginas del áureo libro otra cosa ¡y no es poco, en gracia de
Dios! que lo que en ellas se dice y se enseña.
¡Ah! y si al pasar esto—porque ha de pasar como pasan las
epidemias y las tempestades—nos viéramos libres de las
extravagancias del cervantismo, pudiéramos darnos con un canto en
los pechos; pero, no obstante lo impresionables que somos y lo
propensos, por ende, á olvidar mañana lo que hoy nos alborota,
como el mal deja semillas, éstas germinarán andando los años, y,
cuando menos menos, ha de nacer de ellas una raza que,
empezando por ver zurcidos en el Quijote, acabe por negar la
existencia de su autor.
Todos los grandes hombres van teniendo, en la posteridad, su fama
roída por este género de gusanos. Yo no sé qué demonios anda por
la mollera de ciertos sabios cuando examinan las obras que admira
el mundo, que, no bien las contemplan, cuando ya exclaman: «esto
nació ello solo». ¡Como si no fueran más maravillosas estas
producciones espontáneas que la existencia de un padre que las
engendrara! Á Homero le niega ya el último zarramplín de la crítica,
y hay una Escuela antihomérica, á la cual se van arrimando todos
los catasalsas del helenismo; se está negando también á Hesiodo, y
hasta á Gutenberg y á Dante, y luego se negará la luz del mediodía.
Y ¿por qué no? ¿No hay historiador que niega toda autoridad á los
cinco siglos de Roma? Y la maña es vieja: cien años hace aseguró
el P. Harduino, y hasta intentó probarlo, que todos los libros griegos
y latinos, excepción hecha de unos pocos de Cicerón, Plinio,
Horacio y Virgilio, habían sido forjados en el siglo xii por una
comunidad de frailes.
¡Y qué luz derraman estos sabios negativos en las obscuridades con
que van topando en sus investigaciones! ¡Con qué primor
reconstruyen lo que derriban de un voleo! Paréceles mucha obra la
Ilíada para un hombre solo, de tan remotos siglos; niegan la
existencia de Homero fundándose en aquella potísima razón:
pregúntaseles entonces cómo se formó ese admirable poema, y
responde uno de ellos, Dissen, por ejemplo:
—De la manera más fácil: se reunió una especie de academia de
cantores que se propusieron hacer una epopeya; encargóse cada
cual de un canto, y el resultado de esta asociación fué la Ilíada.
De modo que nos salen, por esta cuenta, veintiséis Homeros, por lo
menos. ¡Y al sabio que los presenta le asombraba, por su grandeza,
un Homero solo!
Dos cuartos de lo mismo ocurre con los sabios de otra catadura,
cuando nos hablan del Universo. Le niegan un Autor, porque no les
cabe en la cabeza la idea de tanto poder, y se le adjudican al átomo,
y sudan y se retuercen entre los laberintos de una tecnología
convencional y de unos procedimientos fantasmagóricos, para venir
á demostrar... que no saben lo que traen entre manos, y que, á
pesar de sus humos de gigantes, no pasan de gusanillos de la tierra,
como el más indocto de los que en ella moramos.
Por eso creo yo que á los sabios de la crítica les pasa algo grave en
la mollera, cada vez que se las han con otras de gran calibre. No
diré que este algo, y aun algos, sean tufillos de la envidia; pero
tampoco aseguro que lo sean de la caridad.
Volviendo al asunto, digo que nacerá quien niegue la existencia de
Cervantes, apoyando el aserto en la autoridad, por supuesto, de otro
sabio, necesariamente francés. Este tal habrá descubierto que en el
siglo xvii no sabían leer ni escribir en España sino los frailes, á los
cuales se debió la traducción, del francés al castellano, de aquel
teatro admirable que ha estado pasando tantísimos años por
español de pura raza; que los nombres de Lope, Moreto, Tirso,
Calderón, etc., etc., no son otra cosa que seudónimos con que se
disfrazaban los traductores temiendo á la Inquisición, que prohibía el
culto de las bellas letras á la gente de cogulla.—En cuanto al Quijote
(seguirá diciendo el sabio de mañana), basta examinarle una vez
para convencerse de que no pudo ser la obra de un hombre solo. La
novela de Crisóstomo, la de Dorotea y Luscinda, la del Curioso
impertinente, la del Cautivo, la del Mozo de mulas, etc., intercaladas
violentamente en la primera parte, y desenlazadas, con otros varios
sucesos, en la Venta de Juan Palomeque el Zurdo, en una sola
noche, lo prueban hasta la evidencia. Esas historias las narrarían los
ciegos por las calles al ronco son de la guitarra, ó las recitarían los
inquisidores en las tertulias de los señores de horca y cuchillo,
mientras las segnoritas y las monjas bailaban el zapateado y el
Jaleo de Jerez. Algún fraile ingenioso las recogió, engarzólas en las
populares aventuras de un loco legendario, llamado, según doctas
pesquisiciones de un bibliómano cochinchino, don Fidalgo de la
Manga, y lo publicó todo bajo el rótulo con que se conoce la obra del
supuesto Cervantes. Por lo que toca á la segunda parte de la
misma, ¿quién ignora que se debe á los frailes Agustinos, que la
escribieron en odio al autor de otro Quijote falsificado, al P.
Abellaneda, Prior de los Jerónimos del Escorial?
Cosa parecida se dirá de las Novelas ejemplares, del Persiles y la
Galatea: tradiciones popularísimas en España, aunque de
procedencia francesa, recogidas y dadas á luz por frailes codiciosos
que explotaban el prestigio del imaginario Cervantes, hecho célebre
desde la aparición de la primera parte del Quijote.
—Pero—seguirá diciendo el futuro bibliófilo francés—¿qué mayor
prueba de la no existencia de Cervantes que la que nos dan los
cervantistas españoles del siglo xix, en el que ya comenzaba á leer
y escribir la clase media, porque se había secularizado la
enseñanza? En el último tercio de aquel siglo no trataron los
escritores de España más que de Cervantes, y, sin embargo, no
pudieron hallar un solo rastro de su persona. Quién le supuso
soldado en Lepanto; quién cautivo en Argel; quién teólogo; quién
marino; quién abogado; quién cocinero; quién médico; quién
ardiente propagandista de la Reforma; quién afirmó que había
nacido en Madrid; quién que en Alcalá; quién que estuvo preso en
Argamasilla; quién que en Valladolid; y nada se prueba en limpio, ni
nadie supo jamás en qué punto de la tierra descansan sus cenizas.
La misma confusión de pareceres se observa en lo relativo al texto
primitivo y á la intención generadora de la novela. Cada edición de
ella en aquel siglo salía ilustrada por un nuevo comentarista, que
quitaba y añadía, á su antojo, frases y períodos, so pretexto de
enmendar así los errores tipográficos del impresor Juan de la
Cuesta. Esto nos hace creer que el Quijote que salió del siglo xix no
se parece en nada al que, por primera vez, publicaron los frailes del
xvii, de cuyas ediciones no ha llegado un solo ejemplar á nuestros
días. Afortunadamente, se conservan catorce volúmenes de un
literato andaluz de aquella centuria, en cuya obra se pone de
manifiesto la verdadera importancia del libro del supuesto
Cervantes. El tal libro es una ingeniosísima alegoría, según afirma el
intérprete feliz de los catorce volúmenes; y á su parecer nos
adherimos, no sin declarar que si el perspicuo andaluz sudó tinta
para dar con la clave del enigma, nosotros hemos sudado pez para
acomodar nuestro criterio á las angosturas, nebulosidades y
retortijones de sus ingeniosos razonamientos. Pero á gimnasias más
abstrusas y complicadas nos tiene avezados el intelecto la filosofía
alemana; y al influjo de esa ciencia, madre de la actual sabiduría,
debemos este descubrimiento portentoso. De modo que bien
podemos decir, con otro ingeniosísimo comentarista,
contemporáneo del de los catorce volúmenes (el cual comentarista
se jactaba de poseer el autógrafo del famoso libro): «Ni Cervantes
es Cervantes, ni el Quijote es el Quijote».
Éstos y otros tales dichos del sabio francés de los futuros siglos,
llegarán á formar escuela; y esta escuela se acreditará en España; y
habrá españoles que se pasarán la vida cotejando el fárrago
cervantista del siglo xix con los asertos de la escuela; y al fin
perderán el juicio, y quizás den origen á una nueva orden de
cervantistas andantes, que saldrán por el mundo á buscar las
aventuras, deshaciendo escolios y enderezando notas al Quijote y á
la dudosa vida de su autor, que es cuanto queda ya que ver.
Entre tanto, cosa es que abruma el espíritu la contemplación del
cervantismo de nuestros días, malgastando lo mejor de la vida en
resobar, sin pizca de respeto, al más ilustre de los nombres y á la
más hermosa de las creaciones del humano ingenio; apesta y
empalaga ese fervor monomaníaco con que todo el mundo se da
hoy á buscar misterios en el fondo del libro, y habilidades en el
autor. Debémosle admiración, y es justo que se la tributemos; pero
no con cascabeles ni vestidos de payasos. Popularícese el Quijote,
y, si es necesario, declárese de texto en las escuelas; pero no el que
nos ofrezca, arreglado á su caletre, el cervantismo al uso.
Si las investigaciones hechas por doctos y respetables literatos,
desde Navarrete hasta Hartzenbusch, no bastan á poner en claro
cuáles son, en las primeras ediciones de Juan de la Cuesta, errores
del impresor, y cuáles descuidos de Cervantes, inténtese esa
empresa; pero una sola vez y por gentes erigidas en autoridad
literaria; y lo que resulte del expurgo, sin más notas que las precisas
para aclarar la significación de palabras poco conocidas hoy del
vulgo, ó para mostrar los pasajes en que Cervantes parodia escenas
y trozos de los libros de caballerías, algo, en suma, de lo que hizo
Clemencín (y no digo todo, porque este comentarista cayó también
en la impertinente tentación de meterse en pespuntes y reparos
gramaticales, como si quisiera enmendar la plana á Cervantes),
guárdese como oro en paño y sea el modelo á que se ajusten
cuantas ediciones del Quijote se hagan en lo sucesivo; pues el mal
no está en que un literato de autoridad y de juicio meta su escalpelo
en las páginas del áureo libro, sino el precedente que de ese modo
se sienta para que todos nos demos á expurgadores de faltas y á
zurcidores de conceptos. Y aun sin este riesgo, ¿qué se saca en
limpio de las enmiendas de los doctos, si cada uno de estos señores
está tan discorde con las de los demás, como lo están todos ellos
con el asendereado Juan de la Cuesta? Y si ya entran por miles las
confesadas alteraciones hechas en el texto de las primeras
ediciones por esos respetables literatos, ¿qué lector, al poner el
dedo sobre una palabra del Quijote, se atreve hoy á asegurar que
esta palabra sea de Cervantes y no de alguno de sus correctores? Y
¿quién se atreverá mañana si á la afición reinante no se le ponen
trabas?
Volviendo al cervantismo inconsciente é intemperante, digo que no
mezcle berzas con capachos, ni confunda tan lastimosamente lo
serio con lo bufo. Elévese una estatua en cada plaza pública
española al príncipe de nuestros novelistas, y sea cada edición de
sus obras un monumento tipográfico; pero, por el amor de Dios, no
pidamos fiestas nacionales para cada uno de sus aniversarios, ni
nos demos todos á académicos cervantinos, ni estampemos el
egregio nombre en desvencijadas diligencias, ni en sociedades de
bailes públicos, ni salgamos á la calle con cara de parientes del
ilustre difunto, ni asociemos su memoria á todas nuestras
debilidades y sandeces. Léase y estúdiese la inmortal obra, que
deleite y enseñanzas contiene para doctos é indoctos en todas las
edades de la vida; pero no pretenda cada lector imponerse á los
demás con el fruto de la tarea; pues cada hombre es un carácter, y,
como dijo un insigne escritor, disputando sobre reparos hechos, y no
del todo mal, á unas enmiendas suyas al Quijote,
«Cada uno tiene, don Zacarías,

Sus aprensiones y sus manías».
¡Y adónde iríamos á parar si se diera, como se va dando, en la

gracia de remendar é interpretar el libro, al tenor de esa suma de
aprensiones, y conforme al parecer de cada aprensivo?
Dudo mucho que el Gobierno de la nación permitiera á los
aficionados á la arquitectura poner sus manos en determinados
detalles artísticos de un monumento público, so pretexto de que así
lo quiso el arquitecto, á quien no deben achacarse los errores de los
canteros. ¿Ha habido pincel que se atreva á borrar el tercer brazo
con que aparece en el Museo uno de los mejores caballos de
Velázquez? Antes al contrario, ¿no se lleva el respeto al gran pintor
al extremo de hacerse las copias de tal cuadro hasta con ese
glorioso arrepentimiento?
¿Por qué no ha de merecernos iguales deferencias y
consideraciones el blasón de nuestra nobleza literaria?
Por lo que á mí toca, desde luego aseguro que, si tuviera poder para
ello, declaraba el Quijote monumento nacional, y no consentiría,
bajo las penas más severas, que se alterara en una sola tilde el
texto de la edición que, por los medios indicados, ó por otros
análogos que se juzgasen mejores, se hubiera declarado oficial, con
todas las solemnidades y garantías apetecibles.
¿Que tiene erratas?... Que las tenga. ¿Que lo del Rucio?... Mejor
que mejor. ¿Habrá trastrueque de párrafos, ni razonamientos que
valgan lo que dice del caso el mismo Cervantes en la segunda parte
de la novela? ¿No son estos descuidos y aquellos arrepentimientos
y los otros deslices gramaticales, el mejor testimonio de la frescura y
espontaneidad de la obra? ¿O creen los químicos del cervantismo
que un libro como el Quijote puede hacerse con regla, compás y
tiralíneas?
Si Cervantes hubiera tenido que estar atento á cuantos tiquis-miquis
le quieren sujetar sus admiradores; si lo que dijo de herir de soslayo
los rayos del sol á su personaje al lanzarse al mundo de las
aventuras, lo dijo para que la posteridad no dudara que salía de
Argamasilla de Alba y no de otro lugar manchego; si no fueron
donaires de su pluma y primores de lengua otros mil pasajes de su
libro, sino estudiados disfraces de otros tantos propósitos
transcendentales; si cada frase es un jeroglífico y cada nombre un
anagrama; si, amén de esto y mucho más, necesitó trabajar con el
calendario á la vista, y encarrilar á su caballero por cualquiera de los
itinerarios que le han trazado sus comentaristas de hogaño, y
conocer á palmos los senderos para no dar con una aventura en
martes, cuando, por el cómputo del mapa y del almanaque, podía
demostrársele que la fazaña debió tener lugar en miércoles, día de
vigilia además, con otros muy curiosos pormenores que el lector
habrá visto, tan bien como yo, en escolios, notas y folletos; si á todo
esto, y á lo de la cocina, la teología, la jurisprudencia, el
protestantismo (!!!), la economía política, etc., etc., etc... y otro tanto
más, tuvo que estar atento, repito, el glorioso novelista, más le
valiera no haber salido nunca del cautiverio de Argel; que entre
escribir un libro con tales trabas, ó arrastrar las de hierro bajo la
penca de un moro argelino, aun con el ingenio de Cervantes optara
yo por el cautiverio, y saldría mejorado en tercio y quinto.
¡Dichoso día aquél en que el cervantismo pase y vuelva á reinar el
Quijote en la patria literatura sin enmiendas, reparos ni aditamentos,
y su autor perínclito sin habilidades ni misterios! Venga, pues, la
inmortal obra sin teologías, náutica ni jurisprudencia, y, sobre todo,
sin claves ni itinerarios ni almanaques; venga, en fin, como la hemos
conocido los que peinamos ya canas, cuando en ella aprendimos á
leer, á pensar y á sentir; que así, al pie de la letra y hasta con las
erratas y garrafales descuidos de los primeros impresores, ha sido
admirada de todos los hombres y traducida á todas las lenguas, y
servido de pedestal á la fama de Cervantes, que ya no cabe en el
mundo.
1880.

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Some Nonecological Systems

Ecosystem Goods and Services

BOX 1.2 (cont’d)

TABLE 1.1 Categories of ecosystem services.

leas os BIOSPHERE BIOSPHERE

What Are the Properties of Ecosystems?

Ecosystems Have Structure

Ecosystems Perform Functions

Ecosystem Structure and Function Are Controlled by Many Factors

Ecosystems Change Through Time

How Do We Classify or Compare Ecosystems?

Why Do Scientists Study Ecosystems?

How Do Ecosystem Scientists Learn about Ecosystems?

Approaches for Learning about Ecosystems

After Carpenter (1998).

Theory and Conceptual Models

LEACHING ANC, pH, BC, Cl, Al, SO42–

BASEFLOW Aqueous reactions

What Do Ecosystem Scientists Measure?

Some Tools in the Ecosystem Scientist’s Toolbox

Balances: Mass and Charge

Precipitation FIGURE 1.4 Average phosphorus inputs and

in outputs ¼ 4.1 kg/year. Inputs outputs ¼ 2.7 kg/

t year or 39.7% retention of phosphorus in the lake.

Ground water out

Ecological Tracers: Isotopes

El Diccionario de la Academia no contiene este vocablo; pero es uno

«Cada uno tiene, don Zacarías,

¡Y adónde iríamos á parar si se diera, como se va dando, en la

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