Townsend C.R., Begon M., Harper J.L. Essentials of Ecology
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ESSENTIALS
OF ECOLOGY
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ESSENTIALS
OF ECOLOGY
Third Edition
Colin R. Townsend
Department of Zoology, University of Otago, Dunedin, New Zealand
Michael Begon
Population Biology Research Group, School of Biological Sciences
The University of Liverpool, Liverpool, UK
John L. Harper
Professor Emeritus in the University of Wales
Visiting Professor in the University of Exeter, Exeter, UK
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© 2008 by Blackwell Publishing
BLACKWELL PUBLISHING
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the services of a competent professional should be sought.
First edition published 2000 by Blackwell Publishing
Second edition published 2003
Third edition published 2008
1 2008
Library of Congress Cataloging-in-Publication Data
Townsend, Colin R.
Essentials of ecology / Colin R. Townsend, Michael Begon, John L. Harper.—3rd ed.
p. cm.
Includes bibliographical references and index.
ISBN 978-1-4051-5658-5 (pbk. : alk. paper)
1. Ecology. I. Begon, Michael. II. Harper, John L. III. Title.
QH541.T66 2008
577—dc22 2007034694
A catalogue record for this title is available from the British Library.
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v
Full contents vi
Preface x
Acknowledgments xii
Part I Introduction 1
1 Ecology and how to do it 3
2 Ecology’s evolutionary backdrop 36
Part II Conditions and Resources 67
3 Physical conditions and the availability of resources 69
4 Conditions, resources and the world’s communities 110
Part III Individuals, Populations, Communities and Ecosystems 143
5 Birth, death and movement 145
6 Interspecific competition 182
7 Predation, grazing and disease 217
8 Evolutionary ecology 251
9 From populations to communities 281
10 Patterns in species richness 323
11 The flux of energy and matter through ecosystems 357
Part IV Applied Issues in Ecology 387
12 Sustainability 389
13 Habitat degradation 423
14 Conservation 455
References 483
Index 495
SHORT CONTENTS
Short contents
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CONTENTS
Preface x
Acknowledgments xii
Part I Introduction 1
1 Ecology and how to do it 3
1.1 Introduction 4
1.2 Scales, diversity and rigor 7
1.3 Ecology in practice 17
2 Ecology’s evolutionary backdrop 36
2.1 Introduction 37
2.2 Evolution by natural selection 37
2.3 Evolution within species 41
2.4 The ecology of speciation 51
2.5 Effects of climatic change on the evolution and distribution of species 58
2.6 Effects of continental drift on the ecology of evolution 60
2.7 Interpreting the results of evolution: convergents and parallels 63
Part II Conditions and Resources 67
3 Physical conditions and the availability
of resources
69
3.1 Introduction 70
3.2 Environmental conditions 71
3.3 Plant resources 84
3.4 Animals and their resources 95
3.5 Effects of intraspecific competition for resources 103
3.6 Conditions, resources and the ecological niche 106
vi
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4 Conditions, resources and the world’s
communities
110
4.1 Introduction 111
4.2 Geographic patterns at large and small scales 111
4.3 Temporal patterns in conditions and resources 117
4.4 Terrestrial biomes 119
4.5 Aquatic environments 130
Part III Individuals, Populations,
Communities and Ecosystems 143
5 Birth, death and movement 145
5.1 Introduction 146
5.2 Life cycles 151
5.3 Monitoring birth and death: life tables and fecundity schedules 156
5.4 Dispersal and migration 164
5.5 The impact of intraspecific competition on populations 169
5.6 Life history patterns 175
6 Interspecific competition 182
6.1 Introduction 183
6.2 Ecological effects of interspecific competition 183
6.3 Evolutionary effects of interspecific competition 197
6.4 Interspecific competition and community structure 200
6.5 How significant is interspecific competition in practice? 208
7 Predation, grazing and disease 217
7.1 Introduction 218
7.2 Prey fitness and abundance 220
7.3 The subtleties of predation 222
7.4 Predator behavior: foraging and transmission 228
7.5 Population dynamics of predation 233
7.6 Predation and community structure 246
8 Evolutionary ecology 251
8.1 Introduction 252
8.2 Molecular ecology: differentiation within and between species 253
8.3 Coevolutionary arms races 262
8.4 Mutualistic interactions 267
Contents
vii
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9 From populations to communities 281
9.1 Introduction 282
9.2 Multiple determinants of the dynamics of populations 283
9.3 Dispersal, patches and metapopulation dynamics 294
9.4 Temporal patterns in community composition 299
9.5 Food webs 307
10 Patterns in species richness 323
10.1 Introduction 324
10.2 A simple model of species richness 326
10.3 Spatially varying factors that influence species richness 328
10.4 Temporally varying factors that influence species richness 337
10.5 Gradients of species richness 340
10.6 Patterns in taxon richness in the fossil record 349
10.7 Appraisal of patterns in species richness 352
11 The flux of energy and matter through ecosystems 357
11.1 Introduction 358
11.2 Primary productivity 360
11.3 The fate of primary productivity 364
11.4 The process of decomposition 369
11.5 The flux of matter through ecosystems 374
11.6 Global biogeochemical cycles 380
Part IV Applied Issues in Ecology 387
12 Sustainability 389
12.1 Introduction 390
12.2 The human population ‘problem’ 391
12.3 Harvesting living resources from the wild 399
12.4 The farming of monocultures 405
12.5 Pest control 412
12.6 Integrated farming systems 417
12.7 Forecasting agriculturally driven global environmental change 419
13 Habitat degradation 423
13.1 Introduction 424
13.2 Degradation via cultivation 428
13.3 Power generation and its diverse effects 435
13.4 Degradation in urban and industrial landscapes 442
13.5 Maintenance and restoration of ecosystem services 448
Contents
viii
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14 Conservation 455
14.1 Introduction 456
14.2 Threats to biodiversity 459
14.3 Conservation in practice 468
14.4 Conservation in a changing world 476
14.5 Finale 479
References 483
Index 495
Contents
ix
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B
y writing this book we hope to share with you some of our wonder at the
complexity of nature, but we must all also be aware that there is a darker
side: the fear that we are destroying our natural environments and the services
they provide. All of us need to be ecologically literate so that we can take part
in political debate and contribute to solving the ecological problems that we
carry with us into the new millennium. We hope our book will contribute to this
objective.
The genesis of this book can be found in the more comprehensive treatment of
ecology in our big book Ecology: from Individuals to Ecosystems (Begon, Townsend
& Harper, 4th edn, 2006). This is used as an advanced university text around
the world, but many of our colleagues have called for a more succinct treatment
of the essence of the subject. Thus, we were spurred into action to produce a
distinctively different book, written with clear objectives for a different audience
– those taking a semester-long beginners course in the essentials of ecology. We
hope that at least some readers will be excited enough to go on to sample the big
book and the rich literature of ecology that it can lead into.
In this third edition of Essentials of Ecology we have made the text, including
mathematical topics, even more accessible. Ecology is a vibrant subject and this
is reflected by our inclusion of literally hundreds of new studies. Some readers will
be engaged most by the fundamental principles of how ecological systems work.
Others will be impatient to focus on the ecological problems caused by human
activities. We place heavy emphasis on both fundamental and applied aspects of
ecology: there is no clear boundary between the two. However, we have chosen to
deal first in a systematic way with the fundamental side of the subject, and we have
done this for a particular reason. An understanding of the scope of the problems
facing us (the unsustainable use of ecological resources, pollution, extinctions
and the erosion of natural biodiversity) and the means to counter and solve these
problems depend absolutely on a proper grasp of ecological fundamentals.
The book is divided into four sections. In the introduction we deal with two
foundations for the subject that are often neglected in texts. Chapter 1 aims to
show not only what ecology is but also how ecologists do it – how ecological
understanding is achieved, what we understand (and, just as important, what we
do not yet understand) and how our understanding helps us predict and manage.
We then introduce ‘Ecology’s evolutionary backdrop’ and show that ecologists
need a full understanding of the evolutionary biologist’s discipline in order to
make sense of patterns and processes in nature (Chapter 2).
What makes an environment habitable for particular species is that they can
tolerate the physicochemical conditions there and find in it their essential resources.
PREFACE
Preface
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