Community Ecology, 2nd Edition

413 Pages·8.889 MB·English

by  Peter J. Morin(auth.)

Community Ecology
Community Ecology
2nd edition
Peter J. Morin
Department of Ecology, Evolution, and Natural Resources
Rutgers University
New Brunswick, New Jersey, USA
A John Wiley & Sons, Ltd., Publication
This edition fi rst published 2011 © by Peter J. Morin
© 1999 by Blackwell Science, Inc.
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Library of Congress Cataloging-in-Publication Data
Morin, Peter J. 1953-
Community ecology / Peter J. Morin. – 2nd ed.
p. cm.
Includes bibliographical references and index.
ISBN 978-1-4443-3821-8 (cloth) – ISBN 978-1-4051-2411-9 (pbk.)
1. Biotic communities. I. Title.
QH541.M574 2011
577.8'2–dc22
2011000108
A catalogue record for this book is available from the British Library.
This book is published in the following electronic formats: ePDF 9781444341935; Wiley Online Library
9781444341966; ePub 9781444341942; Mobi 9781444341959
Set in 9.5/12 pt Berkeley by Toppan Best-set Premedia Limited
1 2011
Contents
Preface to the Second Edition ix
Preface to the First Edition x
Part 1 Communities: Basic Patterns and Elementary Processes 1
1 Communities 3
1.1 Overview 3
1.2 Communities 3
1.3 Communities and their members 7
1.4 Community properties 14
1.5 Interspecifi c interactions 18
1.6 Community patterns as the inspiration for theory: alternate
hypotheses and their critical evaluation 19
1.7 Community patterns are a consequence of a hierarchy
of interacting processes 22
1.8 Conclusions 23
2 Competition: Mechanisms, Models, and Niches 24
2.1 Overview 24
2.2 Interspecifi c competition 24
2.3 Mechanisms of interspecifi c competition 26
2.4 Descriptive models of competition 27
2.5 Mechanistic models of competition 33
2.6 Neighborhood models of competition among plants 40
2.7 Competition, niches, and resource partitioning 46
2.8 The many meanings of the niche 46
2.9 Other ways of thinking about the niche 50
2.10 Guild structure in niche space 54
2.11 Conclusions 55
3 Competition: Experiments, Observations, and Null Models 58
3.1 Overview 58
3.2 Experimental approaches to interspecifi c competition 58
3.3 Experimental studies of interspecifi c competition 62
3.4 Competition in marine communities 62
3.5 Competition in terrestrial communities 65
3.6 Competition in freshwater communities 74
3.7 An overview of patterns found in surveys of published experiments
on interspecifi c competition 79
v
vi CONTENTS
3.8 Null models and statistical/observational approaches to the study
of interspecifi c competition 85
3.9 Conclusions 88
4 Predation and Communities: Empirical Patterns 90
4.1 Overview 90
4.2 Predation 90
4.3 Examples from biological control 91
4.4 Impacts of predators on different kinds of communities 93
4.5 Examples of predation in marine communities 93
4.6 Examples of predation in terrestrial communities 97
4.7 Examples of predation in freshwater communities 105
4.8 Inducible defenses 110
4.9 When is predation likely to regulate prey population size and
community structure? 111
4.10 Overviews of general patterns based on reviews of experimental
studies of predation 116
4.11 Trade-offs between competitive ability and resistance to predation 116
4.12 Conclusions 119
5 Models of Predation in Simple Communities 120
5.1 Overview 120
5.2 Simple predator–prey models 120
5.3 Models of predation on more than one prey 128
5.4 Models of intraguild predation 132
5.5 Models of infectious disease 133
5.6 Conclusions 135
6 Food Webs 136
6.1 Overview 136
6.2 Food-web attributes 136
6.3 Patterns in collections of food webs 144
6.4 Explanations for food-web patterns 147
6.5 Other approaches to modeling food-web patterns 153
6.6 Experimental tests of food-web theory 155
6.7 Omnivory, increasing trophic complexity, and stability 159
6.8 Interaction strength 162
6.9 Some fi nal qualifi cations about empirical patterns 163
6.10 Conclusions 165
7 Mutualisms 166
7.1 Overview 166
7.2 Kinds of mutualisms 166
7.3 Direct and indirect mutualisms 167
7.4 Simple models of mutualistic interactions 167
7.5 Examples of obligate mutualisms 171
7.6 Energetic and nutritional mutualisms 174
7.7 Examples of facultative mutualisms and commensalisms 179
7.8 Theories about the conditions leading to positive interactions
among species 181
CONTENTS vii
7.9 Integrating positive interactions into ecological networks 183
7.10 Conclusions: Consequences of mutualism and commensalism
for community development 186
8 Indirect Effects 187
8.1 Overview 187
8.2 Types of indirect effects 187
8.3 Apparent competition 190
8.4 Indirect mutualism and indirect commensalism 194
8.5 Trophic cascades, tri-trophic interactions, and bottom-up effects 196
8.6 Interaction modifi cations: Higher-order interactions, non-additive
effects, and trait-mediated indirect effects 201
8.7 Indirect effects can complicate the interpretation of manipulative
community studies 206
8.8 Conclusions: Factors contributing to the importance of indirect effects 210
Part 2 Factors Infl uencing Interactions Among Species 213
9 Temporal Patterns: Seasonal Dynamics, Priority Effects, and Assembly Rules 215
9.1 Overview 215
9.2 The importance of history 215
9.3 Interactions among temporally segregated species 217
9.4 Consequences of phenological variation: case studies
of priority effects 224
9.5 Assembly rules 229
9.6 Examples of assembly rules derived from theory 229
9.7 Conclusions 237
10 Habitat Selection 238
10.1 Overview 238
10.2 Features of habitat selection 238
10.3 Correlations between organisms and habitat characteristics 239
10.4 Cues and consequences 241
10.5 A graphical theory of habitat selection 247
10.6 Conclusions 249
11 Spatial Dynamics 251
11.1 Overview 251
11.2 Spatial dynamics in open systems 251
11.3 Metapopulations and metacommunities 252
11.4 Interspecifi c interactions in patchy, subdivided habitats 253
11.5 Competition in spatially complex habitats 253
11.6 Predator–prey interactions in spatially complex habitats 255
11.7 Habitat fragmentation and dispersal corridors affect diversity
and movement among patches 266
11.8 Recruitment-limited interactions – “supply-side ecology” 269
11.9 Large-scale spatial patterns: island biogeography and macroecology 271
11.10 Conclusions 280
viii CONTENTS
Part 3 Large-Scale, Integrative Community Phenomena 281
12 Causes and Consequences of Diversity 283
12.1 Overview 283
12.2 Equilibrium and non-equilibrium communities 284
12.3 Experimental studies of community stability and alternate stable states 290
12.4 Examples of stable community patterns 292
12.5 Equilibrium explanations for diversity 292
12.6 Situations where diversity may result from non-equilibrium dynamics 294
12.7 Stability and complexity 298
12.8 Productivity–diversity curves 301
12.9 Effects of diversity on the variability of processes 314
12.10 Effects of diversity on invasibility 316
12.11 Conclusions 318
13 Succession 319
13.1 Overview 319
13.2 Succession 319
13.3 A brief history of succession 321
13.4 Quantitative models of ecological succession 325
13.5 Case studies of succession in different kinds of habitats 331
13.6 Effects of plant succession on animal assemblages 336
13.7 Succession in microbial assemblages 337
13.8 Conclusions 338
14 Applied Community Ecology 340
14.1 Overview 340
14.2 Anthropogenic changes and applied community ecology 340
14.3 Epidemiology of animal borne diseases 341
14.4 Restoration of community composition and function 342
14.5 Biological control of invasive species 343
14.6 Biomanipulation of water quality 344
14.7 Management of multispecies fi sheries 344
14.8 Optimal design of nature preserves 345
14.9 Predicting and managing responses to global environmental change 345
14.10 Maximization of yield in mixed species agricultural and biofuel systems 347
14.11 Assembly of viable communities in novel environments 347
14.12 Conclusions 348
Appendix: Stability Analysis 349
References 353
Index 384
COMPANION WEBSITE
This book has a companion website:
www.wiley.com/go/morin/communityecology
with Figures and Tables from the book for downloading
Preface to the Second Edition
The second edition of Community Ecology represents an effort to update information that has
been published since the fi rst edition appeared in 1999, as well as to fi ll in some gaps present in
the fi rst edition. As before, the limits of space demand that the book cannot be encyclopedic. The
examples used to illustrate key concepts are the ones that I use in my own graduate course in
community ecology, and I realize that many other fi ne examples of important research in these
areas could have been used instead, but have necessarily gone uncited by me. For that, I apologize
to the many fi ne ecologists whose work I was unable to include here.
The overall organization of the book remains largely unchanged, while I have made an effort
to update the references used in most of the chapters. Some areas of community ecology have
advanced importantly since the fi rst edition appeared, and readers will notice those changes are
particularly refl ected by new content in the chapters on food webs (Chapter 6 ) and the causes
and consequences of diversity (Chapter 1 2 ). The second edition also appears at a time when some
prominent ecologists have questioned whether ecological communities are in fact real entities
whose properties can be understood through studies of local interactions among organisms.
Obviously, having written this book, I do not share this concern, and I hope that the book will
emphasize the many aspects of community ecology that emerge from interactions among organ-
isms in different environments.
A number of colleagues at other universities who have used the fi rst edition in their teaching
have made many helpful comments and suggestions that I have tried to incorporate in the second
edition. For that I am grateful to Laurel Fox, Bob Kooi, Robert Marquis, Wilfred R ö ling, Marcel
van der Heijden, and Herman Verhoef. Thanks also go to the students in my graduate course,
Community Dynamics, who have made comments and suggestions over the years.
Finally, Marsha Morin gets special praise for putting up with me, and running interference for
me, while this project took place. As with the fi rst edition, I could not have completed it without
her love, help, support, and understanding.
Peter Morin
New Brunswick, NJ
2011
ix
Preface to the First Edition
This book is based on the lectures that I have given in a Community Ecology course offered at
Rutgers University over the last 15 years. The audience is typically fi rst year graduate students
who come to the course with a diversity of backgrounds in biology, ecology, and mathematics. I
have tried to produce a book that will be useful both to upper level undergraduates and to gradu-
ate students. The course is structured around lectures on the topics covered here, and those
lectures are supplemented with readings and discussions of original research papers; some are
classic studies, and others are more recent. Throughout that course, the guiding theme is that
progress in community ecology comes from the interplay between theory and experiments.
I fi nd that the examples and case studies highlighted here are particularly useful for making
important points about key issues and concepts in community ecology. I have tried to maintain
a balance between describing the classic studies that every student should know about, and
emphasizing recent work that has the potential to change the way that we think about communi-
ties. Limits imposed by space, time, and economy mean that the coverage of important studies
could not even begin to be encyclopedic. I apologize to the many excellent hard - working ecolo-
gists whose work I was unable to include. I also encourage readers to suggest their favorite
examples or topics that would make this book more useful.
Early drafts of most of these chapters were written while I was a visiting scientist at the Centre
for Population Biology, Imperial College at Silwood Park, Ascot, UK. Professor John Lawton was
an ideal host during those stays, and he deserves special thanks for making those visits possible.
The CPB is a stimulating place to work and write while free from the distractions of one ’ s home
university.
During the prolonged period during which this book took form, several of my graduate stu-
dents, current and past, took the time to read most of the chapters and make careful comments
on them. For that I thank Sharon Lawler, Jill McGrady- Steed, Mark Laska, Christina Kaunzinger,
Jeremy Fox, Yoko Kato, Marlene Cole, and Timon McPhearson. Other colleagues at other universi-
ties including Norma Fowler, Mark McPeek, Tom Miller, and Jim Clark commented on various
drafts of different chapters. Any errors or omissions remain my responsibility.
Simon Rallison of Blackwell originally encouraged me to begin writing this book. Along the
way the process was facilitated by the able editorial efforts of Jane Humphreys, Nancy Hill -
Whilton, and Irene Herlihy. Jennifer Rosenblum and Jill Connor provided frequent editorial
feedback and the necessary prodding to keep the project going. They have been patient beyond
all reason.
Finally, Marsha Morin deserves special praise for putting up with my many moods while
this project slowly took form. I could not have completed it without her support and
understanding.
P. J. M.

x
Part 1
Communities: Basic Patterns
and Elementary Processes