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. Blackwell Publishing was acquired by John Wiley & Sons in February 2007. Blackwell’s publishing program has been merged with Wiley’s global Scientifi c, Technical and Medical business to form Wiley-Blackwell. 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Designations used by companies to distinguish their products are often claimed as trademarks. All brand names and product names used in this book are trade names, service marks, trademarks or registered trademarks of their respective owners. The publisher is not associated with any product or vendor mentioned in this book. This publication is designed to provide accurate and authoritative information in regard to the subject matter covered. It is sold on the understanding that the publisher is not engaged in rendering professional services. If profes- sional advice or other expert assistance is required, the services of a competent professional should be sought. 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.