Community Ecology - Peter J. Morin - E-Book

Community Ecology E-Book

Peter J. Morin

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Beschreibung

All life on earth occurs in natural assemblages called communities. Community ecology is the study of patterns and processes involving these collections of two or more species. Communities are typically studied using a diversity of techniques, including observations of natural history, statistical descriptions of natural patterns, laboratory and field experiments, and mathematical modelling. Community patterns arise from a complex assortment of processes including competition, predation, mutualism, indirect effects, habitat selection, which result in the most complex biological entities on earth – including iconic systems such as rain forests and coral reefs.

This book introduces the reader to a balanced coverage of concepts and theories central to community ecology, using examples drawn from terrestrial, freshwater, and marine systems, and focusing on  animal, plant, and microbial species. The historical development of key concepts is described using descriptions of classic studies, while examples of exciting new developments in recent studies are used to point toward future advances in our understanding of community organization. Throughout, there is an emphasis on the crucial interplay between observations, experiments, and mathematical models.

This second updated edition is a valuable resource for advanced undergraduates, graduate students, and  established scientists  who seek a broad overview of community ecology. The book has developed from a course in community ecology that has been taught by the author since 1983.

Figures and tables can be downloaded for free from www.wiley.com/go/morin/communityecology

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Veröffentlichungsjahr: 2011

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Table of Contents

Cover

Title page

Copyright page

Preface to the Second Edition

Preface to the First Edition

Part 1: Communities: Basic Patterns and Elementary Processes

1 Communities

1.1 Overview

1.2 Communities

1.3 Communities and their members

1.4 Community properties

1.5 Interspecific interactions

1.6 Community patterns as the inspiration for theory: alternate hypotheses and their critical evaluation

1.7 Community patterns are a consequence of a hierarchy of interacting processes

1.8 Conclusions

2 Competition: Mechanisms, Models, and Niches

2.1 Overview

2.2 Interspecific competition

2.3 Mechanisms of interspecific competition

2.4 Descriptive models of competition

2.5 Mechanistic models of competition

2.6 Neighborhood models of competition among plants

2.7 Competition, niches, and resource partitioning

2.8 The many meanings of the niche

2.9 Other ways of thinking about the niche

2.10 Guild structure in niche space

2.11 Conclusions

3 Competition: Experiments, Observations, and Null Models

3.1 Overview

3.2 Experimental approaches to interspecific competition

3.3 Experimental studies of interspecific competition

3.4 Competition in marine communities

3.5 Competition in terrestrial communities

3.6 Competition in freshwater communities

3.7 An overview of patterns found in surveys of published experiments on interspecific competition

3.8 Null models and statistical/observational approaches to the study of interspecific competition

3.9 Conclusions

4 Predation and Communities: Empirical Patterns

4.1 Overview

4.2 Predation

4.3 Examples from biological control

4.4 Impacts of predators on different kinds of communities

4.5 Examples of predation in marine communities

4.6 Examples of predation in terrestrial communities

4.7 Examples of predation in freshwater communities

4.8 Inducible defenses

4.9 When is predation likely to regulate prey population size and community structure?

4.10 Overviews of general patterns based on reviews of experimental studies of predation

4.11 Trade-offs between competitive ability and resistance to predation

4.12 Conclusions

5 Models of Predation in Simple Communities

5.1 Overview

5.2 Simple predator–prey models

5.3 Models of predation on more than one prey

5.4 Models of intraguild predation

5.5 Models of infectious disease

5.6 Conclusions

6 Food Webs

6.1 Overview

6.2 Food-web attributes

6.3 Patterns in collections of food webs

6.4 Explanations for food-web patterns

6.5 Other approaches to modeling food-web patterns

6.6 Experimental tests of food-web theory

6.7 Omnivory, increasing trophic complexity, and stability

6.8 Interaction strength

6.9 Some final qualifications about empirical patterns

6.10 Conclusions

7 Mutualisms

7.1 Overview

7.2 Kinds of mutualisms

7.3 Direct and indirect mutualisms

7.4 Simple models of mutualistic interactions

7.5 Examples of obligate mutualisms

7.6 Energetic and nutritional mutualisms

7.7 Examples of facultative mutualisms and commensalisms

7.8 Theories about the conditions leading to positive interactions among species

7.9 Integrating positive interactions into ecological networks

7.10 Conclusions: Consequences of mutualism and commensalism for community development

8 Indirect Effects

8.1 Overview

8.2 Types of indirect effects

8.3 Apparent competition

8.4 Indirect mutualism and indirect commensalism

8.5 Trophic cascades, tri-trophic interactions, and bottom-up effects

8.6 Interaction modifications: Higher-order interactions, non-additive effects, and trait-mediated indirect effects

8.7 Indirect effects can complicate the interpretation of manipulative community studies

8.8 Conclusions: Factors contributing to the importance of indirect effects

Part 2: Factors Influencing Interactions Among Species

9 Temporal Patterns: Seasonal Dynamics, Priority Effects, and Assembly Rules

9.1 Overview

9.2 The importance of history

9.3 Interactions among temporally segregated species

9.4 Consequences of phenological variation: case studies of priority effects

9.5 Assembly rules

9.6 Examples of assembly rules derived from theory

9.7 Conclusions

10 Habitat Selection

10.1 Overview

10.2 Features of habitat selection

10.3 Correlations between organisms and habitat characteristics

10.4 Cues and consequences

10.5 A graphical theory of habitat selection

10.6 Conclusions

11 Spatial Dynamics

11.1 Overview

11.2 Spatial dynamics in open systems

11.3 Metapopulations and metacommunities

11.4 Interspecific interactions in patchy, subdivided habitats

11.5 Competition in spatially complex habitats

11.6 Predator–prey interactions in spatially complex habitats

11.7 Habitat fragmentation and dispersal corridors affect diversity and movement among patches

11.8 Recruitment -limited interactions – “supply-side ecology”

11.9 Large-scale spatial patterns: island biogeography and macroecology

11.10 Conclusions

Part 3: Large-Scale, Integrative Community Phenomena

12 Causes and Consequences of Diversity

12.1 Overview

12.2 Equilibrium and non-equilibrium communities

12.3 Experimental studies of community stability and alternate stable states

12.4 Examples of stable community patterns

12.5 Equilibrium explanations for diversity

12.6 Situations where diversity may result from non-equilibrium dynamics

12.7 Stability and complexity

12.8 Productivity–diversity curves

12.9 Effects of diversity on the variability of processes

12.10 Effects of diversity on invasibility

12.11 Conclusions

13 Succession

13.1 Overview

13.2 Succession

13.3 A brief history of succession

13.4 Quantitative models of ecological succession

13.5 Case studies of succession in different kinds of habitats

13.6 Effects of plant succession on animal assemblages

13.7 Succession in microbial assemblages

13.8 Conclusions

14 Applied Community Ecology

14.1 Overview

14.2 Anthropogenic changes and applied community ecology

14.3 Epidemiology of animal borne diseases

14.4 Restoration of community composition and function

14.5 Biological control of invasive species

14.6 Biomanipulation of water quality

14.7 Management of multispecies fisheries

14.8 Optimal design of nature preserves

14.9 Predicting and managing responses to global environmental change

14.10 Maximization of yield in mixed species agricultural and biofuel systems

14.11 Assembly of viable communities in novel environments

14.12 CONCLUSIONS

Appendix: Stability Analysis

References

Index

This edition first 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

Preface to the Second Edition

The second edition of Community Ecology represents an effort to update information that has been published since the first edition appeared in 1999, as well as to fill in some gaps present in the first 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 fine 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 fine 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 first edition appeared, and readers will notice those changes are particularly reflected by new content in the chapters on food webs (Chapter 6) and the causes and consequences of diversity (Chapter 12). 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 organisms in different environments.

A number of colleagues at other universities who have used the first 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 Röling, 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 first edition, I could not have completed it without her love, help, support, and understanding.

Peter Morin

New Brunswick, NJ

2011

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 first 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 graduate 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 find 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 communities. 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 ecologists 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 students, 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 universities 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.

Part 1: Communities: Basic Patterns and Elementary Processes

1

Communities

“Ecology is the science of communities. A study of the relations of a single species to the environment conceived without reference to communities and, in the end, unrelated to the natural phenomena of its habitat and community associations is not properly included in the field of ecology.”

Victor Shelford (1913)

1.1 Overview

This chapter briefly describes how ecological communities are defined and classified, and introduces some of the properties and interactions that community ecologists study. The major interspecific interactions, or elementary processes, among pairs of species include competition, predation, and mutualism. Complex indirect interactions can arise among chains of three or more interacting species. Important community properties include the number of species present, measures of diversity, which reflect both the number and relative abundances of species, and statistical distributions that describe how different species differ in abundance.

Observations of natural patterns and explorations of mathematical models have inspired generalizations about the underlying causes of community organization. One pattern important in the historical development of community ecology concerns an apparent limit to the similarity of coexisting species. The case of limiting similarity provides a cautionary example of the way in which community patterns are initially recognized, explained in terms of causal mechanisms, and eventually evaluated. Community patterns are the consequence of a hierarchy of interacting processes that interact in complex ways to mold the diversity of life on Earth.

1.2 Communities

Our best estimates suggest that somewhere between 1.5 million and 30 million different species of organisms live on Earth today (Erwin 1982; May 1990). The small fraction of this enormous global collection of species that can be found at any particular place is an ecological community. One important goal of community ecology is to understand the origin, maintenance, and consequences of biological diversity within local communities. Different processes, operating on very different time scales, can influence the number and identity of species in communities. Long-term evolutionary processes operating over time scales spanning millions of years can produce different numbers of species in different locations. Short-term ecological interactions can either exclude or facilitate species over shorter time scales ranging from a few hours to many years. This book provides an overview of community patterns and the processes that create them.

Like many fields of modern biology, community ecology began as a descriptive science. Early community ecology was preoccupied with identifying and listing the species found in particular localities (Clements 1916; Elton 1966). These surveys revealed some of the basic community patterns that continue to fascinate ecologists. In many temperate zone communities, a few species are much more common than others. The dominant species often play an important role in schemes used to identify and categorize different communities. But why should some species be much more common than others? Communities also change over time, often in ways that are quite repeatable. But what processes drive temporal patterns of community change, and why are those patterns so regular within a given area? Different communities can also contain very different numbers of species. A hectare of temperate forest in New Jersey in northeastern North America might hold up to 30 tree species (Robichaud and Buell 1973), while a similar sized plot of rainforest in Panama can yield over 200 tree species (Hubbell and Foster 1983). More than 10 different ideas have been proposed to explain the striking latitudinal gradient in biodiversity that contributes to the differences between temperate and tropical communities (Pianka 1988)! While there are many reasonable competing explanations for the commonness and rarity of species, and for latitudinal differences in biodiversity, the exact causes of these very basic patterns remain speculative. Related questions address the consequences of biodiversity for community processes. Do communities with many species function differently from those with fewer species? How do similar species manage to coexist in diverse communities?

The central questions in community ecology are disarmingly simple. Our ability to answer these questions says something important about our understanding of the sources of biological diversity and the processes that maintain biodiversity in an increasingly stressed and fragmented natural ecosystem. Answering these questions allows us to wisely manage the human-dominated artificial communities that include the major agricultural systems that we depend on for food and biologically produced materials, and to restore the natural communities that we have damaged either through habitat destruction or overexploitation.

Ecologists use a variety of approaches to explore the sources of community patterns. Modern community ecology has progressed far beyond basic description of patterns, and often experiments can identify which processes create particular patterns (Hairston 1989). However, some patterns and their underlying processes are experimentally intractable, owing to the fact that the organisms driving those processes are so large, long-lived, or wide-ranging that experimental manipulations are impossible. Consequently, community ecologists must rely on information from many sources, including mathematical models, statistical comparisons, and experiments to understand what maintains patterns in the diversity of life. The interplay among description, experiments, and mathematical models is a hallmark of modern community ecology.

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