9781405156585_1_pre.qxd 11/5/07 14:38 Page i ESSENTIALS OF ECOLOGY 9781405156585_1_pre.qxd 11/5/07 14:38 Page ii 9781405156585_1_pre.qxd 11/5/07 14:39 Page iii 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 9781405156585_1_pre.qxd 11/5/07 14:39 Page iv © 2008 by Blackwell Publishing BLACKWELL PUBLISHING 350 Main Street, Malden, MA 02148-5020, USA 9600 Garsington Road, Oxford OX4 2DQ, UK 550 Swanston Street, Carlton, Victoria 3053, Australia The right of Colin R. Townsend, Michael Begon, and John L. Harper to be identified as the Authors of this Work has been asserted in accordance with the UK Copyright, Designs, and Patents Act 1988. All rights reserved. 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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. Set in 9.5/12pt ClassGarmond by Graphicraft Limited, Hong Kong Printed and bound in Singapore by C.O.S. Printers Pte Ltd The publisher’s policy is to use permanent paper from mills that operate a sustainable forestry policy, and which has been manufactured from pulp processed using acid-free and elementary chlorine-free practices. 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For further information on Blackwell Publishing, visit our website: www.blackwellpublishing.com 9781405156585_1_pre.qxd 11/5/07 14:39 Page v Short contents SHORT CONTENTS 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 v 9781405156585_1_pre.qxd 11/5/07 14:39 Page vi 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 9781405156585_1_pre.qxd 11/5/07 14:39 Page vii Contents vii 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 9781405156585_1_pre.qxd 11/5/07 14:39 Page viii viii Contents 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 9781405156585_1_pre.qxd 11/5/07 14:39 Page ix Contents ix 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