FUNDAMENTALS

OF ECOSYSTEM SCIENCE
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FUNDAMENTALS
OF ECOSYSTEM
SCIENCE
KATHLEEN C. WEATHERS, DAVID L. STRAYER, AND GENE E. LIKENS

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 Contents

Preface ix 3. Secondary Production and Consumer

Energetics 53
DAVID L. STRAYER

I Introduction 53
Consumer Energetics 53
INTRODUCTION Secondary Production 58
Definition of Secondary Production 58
1. Introduction to Ecosystem Science 3 Methods to Estimate Secondary Production 59
KATHLEEN C. WEATHERS, DAVID L. STRAYER, Controls and Prediction of Secondary
AND GENE E. LIKENS Production 61
Production of an Individual Species of
What Is an Ecosystem? 3 Consumer 62
What Are the Properties of Ecosystems? 8 Production of a Guild of Consumers 64
Why Do Scientists Study Ecosystems? 11 Production of the Entire Community of
How Do Ecosystem Scientists Learn about Consumers 66
Ecosystems? 12 Conclusion 69
From There to Here: A Short History of the References 72
Ecosystem Concept in Theory and Practice 19
References 22 4. Organic Matter Decomposition 75
STUART E.G. FINDLAY

Introduction 75
Decomposition of Plant-Derived Particulate
II Matter 78
Organisms Responsible for Decomposition 81
ECOLOGICAL ENERGETICS Controls on Decomposition 84
Interactions with Other Element Cycles 90
2. Primary Production: The Foundation Summary 91
of Ecosystems 27 References 92
MICHAEL L. PACE AND GARY M. LOVETT

Introduction 27
III
Components of Primary Production 28 BIOGEOCHEMISTRY
Measuring Primary Production 33
Regulation of Primary Production 36 5. Element Cycling 97
Rates and Patterns of Primary Production 42 KATHLEEN C. WEATHERS AND HOLLY A. EWING
Fates of Primary Production 46
A Tale of Scale 48 What Is an Element Cycle? 98
Summary 49 The Importance of Chemical Properties 101
References 50 Move, Stick, and Change: A Simple Framework for
References for Table 2.1 51 Elemental Cycling 104

v
vi CONTENTS

What Kinds of Questions Are Associated with Summary 177

Element Cycles? 107 References 177
Some Characteristics Important for Understanding
an Element’s Role in Ecosystems 107
References 108 IV
6. The Carbon Cycle 109 SYNTHESIS
JONATHAN J. COLE
9. Revisiting the Ecosystem Concept:
Why Study the Carbon Cycle? 109 Important Features That Promote
Biogeochemistry of Carbon 109
The Carbon Atom and Its Chemistry 110
Generality and
The Present-Day Global Carbon Cycle and the Understanding 181
Greenhouse Effect 116 MICHAEL L. PACE
The Holocene Preindustrial Global Carbon
Budget 119 Introduction 181
The Carbon Cycle in Selected Ecosystems 123 Budgets and Boundaries 182
Concluding Remarks 132 Inclusiveness and Flexibility 187
References 132 Generality and Prediction 189
References 190
7. The Nitrogen Cycle 137
PETER M. GROFFMAN AND EMMA J. ROSI-MARSHALL
10. Ecosystems in a Heterogeneous
Introduction 137
World 191
The Global Picture 137
STEWARD T.A. PICKETT AND MARY L. CADENASSO
Nitrogen Cycle Processes 140
Nitrogen Cycling in Terrestrial Ecosystems 148 Introduction 191
Nitrogen Cycling in Aquatic Ecosystems 152 The Nature of Heterogeneity 192
Nitrogen Balances: The Enigma of Missing Toward a Framework for Space and Time
Nitrogen 156 Heterogeneity 201
References 157 Internal and External Heterogeneity 205
First Principles for Assessing
8. The Phosphorus Cycle 159
Heterogeneity 208
ELENA M. BENNETT AND MEAGAN E. SCHIPANSKI
Conclusions: Ecosystems in Time
Introduction 159 and Space 209
Background 159 References 211
The Importance of Phosphorus in Terrestrial
Ecosystems 160
The Importance of Phosphorus in Agricultural 11. Controls on Ecosystem Structure
Ecosystems 161 and Function 215
The Importance of Phosphorus in Aquatic KATHLEEN C. WEATHERS, HOLLY A. EWING,
Ecosystems 161 CLIVE G. JONES, AND DAVID L. STRAYER
The Global Phosphorus Cycle 163
Human Alteration of the Global Phosphorus What Do We Mean by “Control”? 215
Cycle 165 Why Do We Care about Controls
The Phosphorus Cycle at the Watershed Scale 171 on Ecosystems? 216
The Phosphorus Cycle at the Local Scale 172 How Are Ecosystems Controlled? 216
Managing Human Interaction with the Phosphorus Complications 228
Cycle 176 References 230
 CONTENTS vii

V 16. Streams and Their Valleys 265

JUDY L. MEYER
CASE STUDIES
References 275
12. From Global Environmental Change
to Sustainability Science: Ecosystem VI
Studies in the Yaqui Valley,
Mexico 233 FRONTIERS
PAMELA A. MATSON
17. Frontiers in Ecosystem Science 279
The Yaqui Valley Case Study 234 HEATHER A. BECHTOLD, JORGE DURÁN, DAVID L.
Lessons from the Yaqui Valley Ecosystems STRAYER, KATHLEEN C. WEATHERS, ANGELICA P.
Study 239 ALVARADO, NEIL D. BETTEZ, MICHELLE A. HERSH,
References 240 ROBERT C. JOHNSON, ERIC G. KEELING, JENNIFER L.
MORSE, ANDREA M. PREVITALI, AND ALEXANDRA
RODRÍGUEZ
13. Ecology of Lyme Disease 243
RICHARD S. OSTFELD Introduction 279
Pressures and Pace of Environmental Change 280
Discovery 243 Conceptual Advances 284
It’s the Deer 244 Technological Advances 288
References 250 The Changing Culture of Science 291
Conclusion 294
14. Understanding Ecosystem References 294
Effects of Dams 253
EMILY H. STANLEY
Appendix: A Primer on Biologically
References 257 Mediated Redox Reactions in
Ecosystems 297
15. Acid Rain 259 STUART E.G. FINDLAY AND DAVID L. STRAYER
GENE E. LIKENS
Glossary 303
References 264 Index 307
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 Preface
This book provides an introduction to
the content, ideas, and major findings of
contemporary ecosystem science. We wrote
the book primarily for beginning graduate
students and advanced undergraduates but
it should also be useful to a broad range of
academic scientists and resource managers,
and even to dedicated amateurs who seek
an introduction to the field. Ecosystem
science is a rigorous, quantitative science;
we assume that readers of the book will
have had an introductory class in ecology
and basic understanding of chemistry and
math. The book deliberately covers multi-
ple approaches to understanding ecosys-
tems (e.g., the use of experiments, theory,
cross-system comparisons), in multiple
environments (terrestrial, freshwater, and
marine; managed, built, and natural ecosys-
tems), across all parts of the world
(although many examples come from the
authors’ experience in North America).
The origins of this book stem from
an intensive two-week Fundamentals of
Ecosystem Ecology class (the FEE class)
that we have taught to graduate students
from around the world every year or two
at the Cary Institute of Ecosystem Studies
since 1989. We, and many of the chapter
authors, have played central roles in the
development, evolution, and running of the
FEE class since its origin.
We decided upon an edited book for sev-
eral reasons, not the least of which was its
genesis in this team-taught course. While
we shepherded and integrated the chapters
and their contents, we also deliberately
allowed—and even encouraged—multiple
ix
approaches, and as a result, multiple
“voices” will be evident throughout the
book. We believe that this diversity reflects
some of the myriad perspectives and
approaches that are fruitfully brought to
bear on the field of ecosystem science.
The book contains six major sections.
The opening chapter introduces the concept
of the ecosystem, explores some of the
consequences of this concept, describes
the intellectual tools of the science, and
briefly reviews the history of this young
science. Chapters 2 through 8 lay the foun-
dation for the study of ecosystems, and
cover the two major branches of ecosystem
science: energetics (Chapters 2
4) and bio-
geochemistry (Chapters 5
8). These chap-
ters present the core content of ecosystem
science—the movement and fate of energy
and materials in ecosystems—in some
detail. In the synthetic Chapters 9
11,
we revisit major themes that cut across mul-
tiple areas of study in ecosystem science.
Authors of these chapters review the power
and utility of the ecosystem concept, the
roles of heterogeneity in space and time,
and the importance of various types of con-
trols in ecosystems. Chapters 12
16 take
ecosystem science into the real world by
illustrating, through five case studies, the
value of ecosystem science in identifying
and solving a range of environmental pro-
blems. The book closes with Chapter 17,
co-authored by several current Cary Institute
postdoctoral associates, which lays out
some challenges and needs for the future.
Today’s ecosystem science is evolving rap-
idly, with major new discoveries and ideas
x PREFACE
emerging every year. The ultimate shape
and contributions of this science remain to
be discovered.
This book benefited from the persistent
and hard work of the Academic Press
team, especially Jill Cetel, Candice Janco,
and multiple graphic artists. We were
also fortunate to have received helpful
and critical reviews of chapters from
colleagues, including Clifford Ochs and
several anonymous reviewers who teach
ecosystem science; their comments sub-
stantially improved the book. We thank
the authors of various chapters for their
scholarship, patience, goodwill, and
commitment to bringing this project to fru-
ition. The Cary Institute’s assistant, Matt
Gillespie, was an enormous help as well.
Finally, generations of FEE students were
and continue to be an impetus
and inspiration to us and the field of eco-
system science.