THEORETICAL ECOLOGY Third Edition PRINCIPLES AND APPLICATIONS ROBERT MAY & ANGELA MCLEAN OXFORD
OXFORDUNIVERSITYPRESSGreat Clarendon Street,Oxford OX2 6DPOxford UniversityPress is a department of the University of Oxford.It furthers the University's objective of excellence in research, scholarship,and education by publishing worldwide inOxford New YorkAuckland CapeTown Darees SalaamHongKongKarachiKuala Lumpur Madrid Melbourne Mexico City NairobiNew Delhi Shanghai Taipei TorontoWith offices inArgentina Austria Brazil Chile Czech Republic France GreeceGuatemala Hungary Italy Japan Poland Portugal SingaporeSouth Korea Switzerland Thailand Turkey Ukraine VietnamOxford is a registered trade mark of Oxford University Pressin the UK and in certain other countriesPublished in the United Statesby Oxford University Press Inc., New YorkC R. M. May and A. R. McLean 2007The moral rights of the authors have been assertedDatabaseright Oxford University Press (maker)First published 2007All rights reserved. No part of this publication may be reproduced,stored in a retrieval system, or transmitted, in any form or by any means,without the prior permission in writing of Oxford University Press,or as expressly permitted by law, or under terms agreed with the appropriatereprographics rights organization. Enquiries concerning reproductionoutside the scope of the above should be sent to the Rights Department,Oxford University Press, at the address aboveYou must not circulate this book in any other binding or coverand you must impose the same condition on any acquirerBritish Library Cataloguing in Publication DataData availableLibrary of Congress Cataloging in Publication DataData availableTypeset by Newgen Imaging Systems (P) Ltd, Chennai, IndiaPrinted in Great Britainon acid-free paper byAntony Rowe Ltd., Chippenham, WiltshireISBN978-0-19-920998-9 (Hbk)978-019-920999-6 (Pbk)10987654321
1 Great Clarendon Street, Oxford OX2 6DP Oxford University Press is a department of the University of Oxford. It furthers the University’s objective of excellence in research, scholarship, and education by publishing worldwide in Oxford New York Auckland Cape Town Dare es Salaam Hong Kong Karachi Kuala Lumpur Madrid Melbourne Mexico City Nairobi New Delhi Shanghai Taipei Toronto With offices in Argentina Austria Brazil Chile Czech Republic France Greece Guatemala Hungary Italy Japan Poland Portugal Singapore South Korea Switzerland Thailand Turkey Ukraine Vietnam Oxford is a registered trade mark of Oxford University Press in the UK and in certain other countries Published in the United States by Oxford University Press Inc., New York # R. M. May and A. R. McLean 2007 The moral rights of the authors have been asserted Database right Oxford University Press (maker) First published 2007 All rights reserved. No part of this publication may be reproduced, stored in a retrieval system, or transmitted, in any form or by any means, without the prior permission in writing of Oxford University Press, or as expressly permitted by law, or under terms agreed with the appropriate reprographics rights organization. Enquiries concerning reproduction outside the scope of the above should be sent to the Rights Department, Oxford University Press, at the address above You must not circulate this book in any other binding or cover and you must impose the same condition on any acquirer British Library Cataloguing in Publication Data Data available Library of Congress Cataloging in Publication Data Data available Typeset by Newgen Imaging Systems (P) Ltd., Chennai, India Printed in Great Britain on acid-free paper by Antony Rowe Ltd., Chippenham, Wiltshire ISBN 978–0–19–920998–9 (Hbk) 978–0–19–920999–6 (Pbk) 10 9 8 7 6 5 4 3 2 1
ContentsviiAcknowledgementsixContributors11 IntroductionAngela R.McLean and Robert M.May72 How populations cohere: five rules for cooperationMartinA.NouwakandKarlSigmund173 Single-species dynamicsTim Coulson and H. Charles J. Godfray354 Metapopulations and their spatial dynamicsSean Nee465Predator-prey interactionsMichaelB.Bonsall andMichaelP.Hassell 62 6PlantpopulationdynamicsMichael J.Crazwley847Interspecificcompetitionandmultispecies coexistenceDavid Tilman988 Diversity and stability in ecological communitiesAnthony R. Ives1119Communities:patternsRobert M.May,Michael J.Crawley,and George Sugihara13210 Dynamics of infectious diseaseBryan Grenfell and Matthew Keeling14811 FisheriesJohn R.Beddington and Geoffrey P.Kirkzwood15812 A doubly Green Revolution:ecology and food productionGordon Conway17213Conservation biology: unsolved problems and their policy implicationsAndyDobson,Will R.Turner,and David S.Wilcove>
Contents Acknowledgements vii Contributors ix 1 Introduction 1 Angela R. McLean and Robert M. May 2 How populations cohere: five rules for cooperation 7 Martin A. Nowak and Karl Sigmund 3 Single-species dynamics 17 Tim Coulson and H. Charles J. Godfray 4 Metapopulations and their spatial dynamics 35 Sean Nee 5 Predator–prey interactions 46 Michael B. Bonsall and Michael P. Hassell 6 Plant population dynamics 62 Michael J. Crawley 7 Interspecific competition and multispecies coexistence 84 David Tilman 8 Diversity and stability in ecological communities 98 Anthony R. Ives 9 Communities: patterns 111 Robert M. May, Michael J. Crawley, and George Sugihara 10 Dynamics of infectious disease 132 Bryan Grenfell and Matthew Keeling 11 Fisheries 148 John R. Beddington and Geoffrey P. Kirkwood 12 A doubly Green Revolution: ecology and food production 158 Gordon Conway 13 Conservation biology: unsolved problems and their policy implications 172 Andy Dobson, Will R. Turner, and David S. Wilcove v
viCONTENTS19014 Climate change and conservation biologyJeremy T. Kerr and Heather M. Kharouba20515Unanswered questions and why they matterRobert M.May216References249Index
14 Climate change and conservation biology 190 Jeremy T. Kerr and Heather M. Kharouba 15 Unanswered questions and why they matter 205 Robert M. May References 216 Index 249 vi CONTENTS
AcknowledgementsThe aims and scope of this book are set out in thebring the authors and others together for a 2-daybeginning of the first chapter (unimaginativelyconference, in which the sweep of material in thislabelled Introduction). So these prefacing com-book was exposed to discussion and constructivecriticism. It helped shape the book.ments are confined to acknowledging some of thehelp we and the other authors have received inOur thanks are also owed to enthusiastic andhelpful people at Oxford University Press, parti-putting this book together.The two of us are deeply indebted to the other 21cularlythe commissioningeditor,lan Sherman,theauthors who have contributed to the book, bothproduction editor, Christine Rode, and the copy-for the work they did and for their exemplaryeditor, Nik Prowse.R.M.M's assistant, Chrisadherence to a rather fast production scheduleBond, was her usual invaluable self, helping inIndividual authors have wished to thank bothevery facet of the enterprise with unflappablefunding agencies and helpful colleagues who gavecompetence.assistance of various kinds.This would have beenSadly,one of the authorsGeoff Kirkwood-an impressively long list, but we unkindly decidedunexpectedly died the week after the gatheringagainst including it.in Oxford.Everyoneremembershim with affec-We must, however, recognize the generosity oftion, and his shadow lies on the book. He will beMerton Collegeand theZoologyDepartment atmissed.the University of Oxford, and particularly theirrespective heads,Dame Jessica Rawson andA.R. McLean and R.M. MayProfessor Paul Harvey.They made it possibleto22September2006vi
Acknowledgements The aims and scope of this book are set out in the beginning of the first chapter (unimaginatively labelled Introduction). So these prefacing comments are confined to acknowledging some of the help we and the other authors have received in putting this book together. The two of us are deeply indebted to the other 21 authors who have contributed to the book, both for the work they did and for their exemplary adherence to a rather fast production schedule. Individual authors have wished to thank both funding agencies and helpful colleagues who gave assistance of various kinds. This would have been an impressively long list, but we unkindly decided against including it. We must, however, recognize the generosity of Merton College and the Zoology Department at the University of Oxford, and particularly their respective heads, Dame Jessica Rawson and Professor Paul Harvey. They made it possible to bring the authors and others together for a 2-day conference, in which the sweep of material in this book was exposed to discussion and constructive criticism. It helped shape the book. Our thanks are also owed to enthusiastic and helpful people at Oxford University Press, particularly the commissioning editor, Ian Sherman, the production editor, Christine Rode, and the copyeditor, Nik Prowse. R.M.M.’s assistant, Chris Bond, was her usual invaluable self, helping in every facet of the enterprise with unflappable competence. Sadly, one of the authors—Geoff Kirkwood— unexpectedly died the week after the gathering in Oxford. Everyone remembers him with affection, and his shadow lies on the book. He will be missed. A.R. McLean and R.M. May 22 September 2006 vii