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LANDSCAPE
ECOLOGY IN
THEORY AND
PRACTICE
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M O N I C A G . T U R N E R
R O B E R T H . G A R D N E R
R O B E R T V . O ' N E I L L
LANDSCAPEECOLOGY INTHEORY ANDPRACTICEP a t t e r n a n d P r o c e s s
Springer
Monica G. TurnerDepartment of ZoologyBirge HallUniversity of WisconsinMadison, WI [email protected]
Robert H. GardnerAppalachian LaboratoryUniversity of Maryland301 Braddock RoadFrostburg, MD [email protected]
Robert V. O’NeillEnvironmental Sciences DivisionOak Ridge National LaboratoryOak Ridge, TN [email protected]
© 2001 Springer-Verlag New York, Inc.
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Library of Congress Cataloging-in-Publication DataLandscape ecology in theory and practice : pattern
and process / Monica G. Turner, Robert H. Gardner,Robert V. O’Neill.
p. cm.Includes bibliographical references (p. ).ISBN 0-387-95122-9 (alk. paper)ISBN 0-387-95123-7 (softcover : alk. paper)1. Landscape ecology. I. Gardner, R.H.
II. O’Neill, R.V. (Robert V.), 1940– III. Title.QH541.15.L35T87 2001577—dc21 00-047094
Printed in the United States of America.
9 8 7 6 5 4 3 2 1
ISBN 0-387-95122-9 SPIN 10778231 (hardcover)ISBN 0-387-95123-7 SPIN 10778249 (softcover)
Springer-Verlag New York Berlin HeidelbergA member of BertelsmannSpringer Science+Business Media GmbH
Preface
Landscape ecology is not a distinct discipline or simply a branch of ecology, but rather is the synthetic
intersection of many related disciplines that focus on the spatial-temporal pattern of the landscape.
Risser et al., 1984
The emergence of landscape ecology as a discipline has catalyzed a shift in paradigms among ecologists,
. . . resource managers and land-use planners. Having now seen the faces of spatial pattern and scale
. . . we can never go back to the old ways of viewing things. Wiens, 1999
This book presents the perspective of three ecologists on the concepts andapplications of landscape ecology, a discipline that has shown expansive
growth during the past two decades. Although landscape ecology is a multidisci-plinary subject involving components as diverse as economics and sociology, theearth sciences and geography, remote sensing and computer applications, we fo-cus here on what ecologists need to know about landscapes.
Landscape ecology served as the integrating theme of our collaborative researchfor nearly 15 years, including a 7-year period during which we worked togetherat Oak Ridge National Laboratory. We became acquainted in January 1986 atthe first annual United States Landscape Ecology symposium held at the Univer-sity of Georgia and organized by Monica Turner and Frank Golley. Landscapeecology was, at that time, a new subject in the United States. The first U.S. work-shop on landscape ecology, organized by Paul Risser, Richard Forman, and JimKarr, had occurred less than 3 years prior (Risser et al., 1984). One of us (O’Neill)was a participant in that workshop, and two of us (O’Neill and Gardner) wereresearch scientists in the Environmental Sciences Division of Oak Ridge National
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Laboratory (ORNL) who had collaborated for several years on many aspects ofecosystem ecology and ecological modeling. Turner had a newly minted Ph.D. andwas continuing as a postdoctoral research associate at the University of Georgia,where Frank Golley (also a participant at the 1993 workshop and Turner’s Ph.D.advisor) was actively engaging his colleagues in the developing ideas in landscapeecology. The mutual interests shared by Turner, Gardner, and O’Neill, coupledwith the excitement and challenge of working in a newly emerging branch of ecol-ogy, led to Turner’s move to ORNL in July 1987. The subsequent seven years inwhich we collaborated so closely were among the most exciting times that any ofus have had in our careers. There are times and places at which creativity seemsto be fostered more than others, and that time and place had it. The writing ofthis book was precipitated, in part, by the fact that although we are now locatedat different institutions, we shared the desire to provide a synthesis of the field inwhich we have worked so closely together.
As ecologists embraced the challenges of understanding spatial complexity,landscape ecology moved from being a tangential subdiscipline in the early 1980sto one that is now mainstream. Indeed, a landscape approach, or the landscapelevel, is now considered routinely in all types of ecological studies. It is our hopethat this text will provide a synthetic overview of landscape ecology, including itsdevelopment, the methods and techniques that are employed, the major questionsaddressed, and the insights that have been gained. The enclosed CD contains allfigures for this book, including the color images. We hope that this will enhancethe utility of the book, especially for teaching. The companion volume (Gergeland Turner, 2001) provides opportunities for hands-on learning of many of themethods and concepts employed by landscape ecologists. It is our hope that ourbooks might serve to inspire others to embark on landscape ecological studies, forthere is much yet to be learned. As we begin this new century, we look forwardto the many contributions that landscape ecologists will make in the future andto the continued growth of this exciting discipline.
A c k n o w l e d g m e n t s
Research that we have conducted over the past 15 years and that led to the de-velopment of this book has been funded by a variety of agencies, and we grate-fully acknowledge research support from the National Science Foundation (Long-term Ecological Research, Ecosystem Studies, and Ecology programs), Departmentof Energy, USDA Competitive Grants Program, National Geographic Society,
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Environmental Protection Agency (EMAP and STAR programs), and the Univer-sity of Wisconsin-Madison Graduate School.
Our ideas have evolved over the years and been shaped by fruitful and oftenspirited discussions with many colleagues. Among the most memorable of thesewere discussions with Don DeAngelis, Jeff Klopatek, John Krummel, and GeorgeSugihara in the “prelandscape” years at ORNL that crystallized much of the phi-losophy and approach adopted in our research. Virginia Dale, Kim With, ScottPearson, and Bill Romme have been regular collaborators as well as supportivefriends. Although it is impossible to mention everyone at ORNL who contributedideas and assisted us with their expertise, we would be remiss not to acknowledgethe valuable contributions of Steve Bartell, Antoinette Brenkert, Carolyn Hun-saker, Tony King, and Robin Graham. While at Oak Ridge we hosted a numberof visitors from other institutions, including Bill Romme, Linda Wallace, BruceMilne, Tim Kratz, Sandra Lavorel, Tim Allen, Eric Gustafson, and Roy Plotnick.These colleagues made substantial contributions to and lasting impacts on ourideas, and we thank them all for engaging interactions and fruitful collaborations.
Special thanks are due to Richard Forman, Frank Golley, and John Wiens forlongstanding collegial relationships, the sharing of their ideas (and students!) thatoften challenged our thinking, and their invaluable reviews and critiques over theyears. Turner also sincerely thanks Hazel Delcourt (University of Tennessee) andDavid Mladenoff (University of Wisconsin), with whom she has jointly taughtlandscape ecology courses over the past decade; co-teaching has been inspiringand fun and has certainly helped shape her thinking.
This book benefited tremendously from valuable critical comments providedby numerous colleagues. We especially thank David Mladenoff and Sarah Gergel,who both read nearly the entire manuscript and provided constructive criticismthat has been enormously helpful. David Mladenoff actually read the whole man-uscript twice, and Turner especially thanks him for being such a good colleague.In addition, we are grateful to the following friends and colleagues for reviewingone or more chapters: Jeff Cardille, Steve Carpenter, F. S. (Terry) Chapin, MarkDixon, Tony Ives, Dan Kashian, Jim Miller, Bill Romme, Tania Schoennagel, SteveSeagle, Emily Stanley, Dan Tinker, Phil Townshend, and Kim With. Commentson draft chapters from the students in Principles of Landscape Ecology, taught atthe University of Wisconsin-Madison during the spring 1999 semester, were alsovery helpful.
The graphics and illustrations for this book were prepared by Michael Turner,and we are indebted to him for greatly improving the visual communication of
viii
the concepts and examples in this book. We are delighted with the clarity andconsistency of the figures throughout the text. We thank Kandis Elliot (Universityof Wisconsin) and Michael Mac (Biological Resources Division, U.S. GeologicalSurvey) for sharing visual resources. Sandi Gardner and Sally Tinker providedvaluable editorial assistance in the final stages of manuscript preparation. Finally,we thank the two editors with whom we worked at Springer-Verlag, initially RobGarber and then Robin Smith, for their patience and support of this effort, espe-cially given the time it has taken us to complete it.
Monica G. TurnerMadison, Wisconsin
Robert H. GardnerFrostburg, Maryland
Robert V. O’NeillOak Ridge, Tennessee
� R E F E R E N C E S
GERGEL, S. E., AND M. G. TURNER, editors. 2001. Learning Landscape Ecology: A
Practical Guide to Concepts and Techniques. Springer-Verlag, New York.
RISSER, P. G., J. R. KARR, AND R. T. T. FORMAN. 1984. Landscape Ecology: Directions
and Approaches. Special Publication Number. Illinois Natural History Survey,
Champaign, Illinois.
WIENS, J. A. 1999. The science and practice of landscape ecology, in J. M. Klopatek
and R. H. Gardner, eds. Landscape Ecological Analysis: Issues and Applications,
pp. 371–383. Springer-Verlag, New York.
Contents
Preface v
Chapter 1 � Introduction to Landscape Ecology 1
What Is Landscape Ecology? 2
Why Landscape Ecology Has Emerged as a Distinct Area of Study 7
The Intellectual Roots of Landscape Ecology 10
Objectives of This Book 20
Summary 21
Discussion Questions 22
Recommended Readings 23
Chapter 2 � The Critical Concept of Scale 25
Scale Terminology and Its Practical Application 27
Scale Problems 32
x
Scale Concepts and Hierarchy Theory 34
Identifying the “Right” Scale(s) 38
Reasoning About Scale 40
Scaling Up 40
Summary 43
Discussion Questions 44
Recommended Readings 45
Chapter 3 � Introduction to Models 47
What are Models and Why Do We Use Them? 47
Steps in Building a Model 56
Landscape Models 64
Caveats in the Use of Models 66
Summary 67
Discussion Questions 68
Recommended Readings 69
Chapter 4 � Causes of Landscape Pattern 71
Abiotic Causes of Landscape Pattern 73
Biotic Interactions 83
Human Land Use 86
Disturbance and Succession 90
Summary 90
Discussion Questions 92
Recommended Readings 92
Chapter 5 � Quantifying Landscape Pattern 93
Why Quantify Pattern? 93
Data Used in Landscape Analyses 95
Caveats for Landscape Pattern Analysis, or “Read This First” 99
Metrics for Quantifying Landscape Pattern 108
Geostatistics or Spatial Statistics 125
xi
Summary 132
Discussion Questions 133
Recommended Readings 134
Chapter 6 � Neutral Landscape Models 135
Random Maps: The Simplest Neutral Model 138
Maps with Hierarchical Structure 147
Fractal Landscapes 149
Neutral Models Relating Pattern to Process 153
General Insights from the Use of NLMs 153
Summary 155
Discussion Questions 156
Recommended Readings 156
Chapter 7 � Landscape Disturbance Dynamics 157
Disturbance and Disturbance Regimes 159
Influence of the Landscape on Disturbance Pattern 162
Influence of Disturbance on Landscape Pattern 174
Concepts of Landscape Equilibrium 188
Summary 196
Discussion Questions 198
Recommended Readings 199
Chapter 8 � Organisms and Landscape Pattern 201
Conceptual Development of Organism-Space Interactions 204
Scale-Dependent Nature of Organism Responses 221
Effects of Spatial Pattern on Organisms 229
Spatially Explicit Population Models 240
Summary 243
Discussion Questions 246
Recommended Readings 247
xii
Chapter 9 � Ecosystem Processes in the Landscape249
Spatial Heterogeneity in Ecosystem Processes 251
Effects of Landscape Position on Lake Ecosystems 261
Land–Water Interactions 265
Linking Species and Ecosystems 280
Searching for General Principles 284
Summary 285
Discussion Questions 287
Recommended Readings 288
Chapter 10 � Applied Landscape Ecology 289
Land Use 290
Forest Management 307
Regional Risk Assessment 314
Continental-Scale Monitoring 319
Summary 321
Discussion Questions 324
Recommended Readings 325
Chapter 11 � Conclusions and Future Directions327
What Have We Learned? 328
Research Directions 329
Conclusion 331
Discussion Questions 332
Recommended Readings 332
References 333
Index 389