Theoretical Ecology This page intentionally left blank Theoretical Ecology Principles and Applications EDITED BY Robert M. May, Department of Zoology, University of Oxford, Oxford, UK AND Angela R. McLean, Department of Zoology, University of Oxford, Oxford, UK 1 1 GreatClarendonStreet,OxfordOX26DP OxfordUniversityPressisadepartmentoftheUniversityofOxford. ItfurtherstheUniversity’sobjectiveofexcellenceinresearch,scholarship, andeducationbypublishingworldwidein Oxford NewYork Auckland CapeTown DareesSalaam HongKong Karachi KualaLumpur Madrid Melbourne MexicoCity Nairobi NewDelhi Shanghai Taipei Toronto Withofficesin Argentina Austria Brazil Chile CzechRepublic France Greece Guatemala Hungary Italy Japan Poland Portugal Singapore SouthKorea Switzerland Thailand Turkey Ukraine Vietnam OxfordisaregisteredtrademarkofOxfordUniversityPress intheUKandincertainothercountries PublishedintheUnitedStates byOxfordUniversityPressInc.,NewYork #R.M.MayandA.R.McLean2007 Themoralrightsoftheauthorshavebeenasserted DatabaserightOxfordUniversityPress(maker) Firstpublished2007 Allrightsreserved.Nopartofthispublicationmaybereproduced, storedinaretrievalsystem,ortransmitted,inanyformorbyanymeans, withoutthepriorpermissioninwritingofOxfordUniversityPress, orasexpresslypermittedbylaw,orundertermsagreedwiththeappropriate reprographicsrightsorganization.Enquiriesconcerningreproduction outsidethescopeoftheaboveshouldbesenttotheRightsDepartment, OxfordUniversityPress,attheaddressabove Youmustnotcirculatethisbookinanyotherbindingorcover andyoumustimposethesameconditiononanyacquirer BritishLibraryCataloguinginPublicationData Dataavailable LibraryofCongressCataloginginPublicationData Dataavailable TypesetbyNewgenImagingSystems(P)Ltd.,Chennai,India PrintedinGreatBritain onacid-freepaperby AntonyRoweLtd.,Chippenham,Wiltshire ISBN978–0–19–920998–9(Hbk) 978–0–19–920999–6(Pbk) 10 9 8 7 6 5 4 3 2 1 Contents Acknowledgements vii Contributors ix 1 Introduction 1 AngelaR.McLeanandRobert M.May 2 Howpopulations cohere:fiverulesforcooperation 7 MartinA.Nowakand KarlSigmund 3 Single-species dynamics 17 TimCoulsonand H.CharlesJ.Godfray 4 Metapopulations andtheirspatial dynamics 35 SeanNee 5 Predator–prey interactions 46 MichaelB.BonsallandMichaelP.Hassell 6 Plantpopulation dynamics 62 MichaelJ.Crawley 7 Interspecific competition andmultispecies coexistence 84 DavidTilman 8 Diversity andstabilityinecological communities 98 AnthonyR.Ives 9 Communities: patterns 111 RobertM.May,MichaelJ.Crawley,andGeorgeSugihara 10 Dynamics ofinfectiousdisease 132 BryanGrenfellandMatthewKeeling 11 Fisheries 148 JohnR.BeddingtonandGeoffreyP.Kirkwood 12 AdoublyGreen Revolution:ecology andfood production 158 GordonConway 13 Conservationbiology:unsolvedproblemsandtheirpolicyimplications 172 AndyDobson,WillR. Turner,andDavidS.Wilcove v vi CONTENTS 14 Climatechangeandconservationbiology 190 JeremyT.KerrandHeather M.Kharouba 15 Unansweredquestionsandwhytheymatter 205 RobertM.May References 216 Index 249 Acknowledgements The aims andscope ofthis book aresetout inthe bring the authors and others together for a 2-day beginning of the first chapter (unimaginatively conference, in which the sweep of material in this labelled Introduction). So these prefacing com- book was exposed to discussion and constructive ments are confined to acknowledging some of the criticism. It helped shape the book. help we and the other authors have received in Our thanks are also owed to enthusiastic and putting this book together. helpful people at Oxford University Press, parti- The two of us are deeply indebted to the other 21 cularlythecommissioningeditor,IanSherman,the authors who have contributed to the book, both production editor, Christine Rode, and the copy- for the work they did and for their exemplary editor, Nik Prowse. R.M.M.’s assistant, Chris adherence to a rather fast production schedule. Bond, was her usual invaluable self, helping in Individual authors have wished to thank both every facet of the enterprise with unflappable fundingagenciesandhelpfulcolleagueswhogave competence. assistance ofvariouskinds.This would havebeen Sadly, one of the authors—Geoff Kirkwood— animpressivelylonglist,butweunkindlydecided unexpectedly died the week after the gathering against including it. in Oxford. Everyone remembers him with affec- We must, however, recognize the generosity of tion, and his shadow lies on the book. He will be Merton College and the Zoology Department at missed. the University of Oxford, and particularly their respective heads, Dame Jessica Rawson and A.R. McLean and R.M. May Professor Paul Harvey. They made it possible to 22 September 2006 vii This page intentionally left blank Contributors JohnR.Beddington,DivisionofBiology,Facultyof HeatherKharouba,CanadianFacilityforEcoinformatics NaturalSciences,RSMBuilding,ImperialCollege Research(CFER),DepartmentofBiology,Universityof London,SW72BP,UK.E-mail:j.beddington@ Ottawa,Box450,StationA,Ottawa,ON,K1N6N5, imperial.ac.uk Canada.E-mail:[email protected] MichaelB.Bonsall,DepartmentofZoology,Tinbergen GeoffreyP.Kirkwood,DivisionofBiology,Facultyof Building,UniversityofOxford,OxfordOX13PS,UK. NaturalSciences,RSMBuilding,ImperialCollege E-mail:[email protected] London,SW72BP,UK GordonConway,CentreforEnvironmentalPolicy,4th RobertM.May,DepartmentofZoology,Tinbergen Floor,RSMBuilding,ImperialCollege,South Building,UniversityofOxford,OxfordOX13PS,UK. Kensington,LondonSW72AZ,UK.E-mail: E-mail:[email protected] [email protected] AngelaR.McLean,DepartmentofZoology,Tinbergen TimCoulson,NERCCentreforPopulationBiologyand Building,UniversityofOxford,OxfordOX13PS,UK. DivisionofBiology,ImperialCollegeLondon,Silwood E-mail:[email protected] ParkCampus,Ascot,BerkshireSL57PY,UK.E-mail: SeanNee,InstituteofEvolutionaryBiology,Schoolof [email protected] BiologicalSciences,UniversityofEdinburgh,West MichaelJ.Crawley,DepartmentofBiologicalSciences, MainsRoad,EdinburghEH93JT,UK.E-mail: ImperialCollegeLondon,SilwoodPark,Ascot,Berk- [email protected] shireSL57PY,UK.E-mail:[email protected] MartinA.Nowak,TheProgramforEvolutionary AndyDobson,EcologyandEvolutionaryBiology, Dynamics,FacultyofArtsandScience,OneBrattle PrincetonUniversity,Princeton,NJ08544,USA. Square,HarvardUniversity,Cambridge,MA02138, E-mail:[email protected] USA.E-mail:[email protected] H.CharlesJ.Godfray,DepartmentofZoology, KarlSigmund,FacultyforMathematics,Universityof TinbergenBuilding,UniversityofOxford,Oxford Vienna,Nordbergstrasse15,A-1090Vienna,Austria. OX13PS,UK.E-mail:[email protected] E-mail:[email protected] BryanGrenfell,BiologyDepartment,208Mueller GeorgeSugihara,ScrippsInstitutionofOceanography, Laboratory,PennsylvaniaStateUniversity,University UniversityofCalifornia,SanDiego,9500Gilman Park,PA16802,USA.E-mail:grenfell@ psu.edu Drive,LaJolla,CA920930202,USA.E-mail: MichaelP.Hassell,DepartmentofBiologicalSciences, [email protected] ImperialCollegeLondon,SilwoodPark,Ascot,Berk- DavidTilman,DepartmentofEcology,Evolutionand shireSL57PY,UK.E-mail:[email protected] Behavior,UniversityofMinnesota,St.Paul,MN55108, AnthonyR.Ives,DepartmentofZoology,Universityof USA.E-mail:[email protected] Wisconsin,Madison,WI53706,USA. WillR.Turner,CenterforAppliedBiodiversity E-mail:[email protected] Science,ConservationInternational,1919MSt.NW MatthewKeeling,DepartmentofBiologicalSciences Suite600,Washington,DC20036,USA.E-mail: andMathematicsInstitute,UniversityofWarwick, [email protected] GibbetHillRoad,CoventryCV47AL,UK.E-mail: DavidS.Wilcove,EcologyandEvolutionaryBiologyand [email protected] PrincetonEnvironmentalInstituteandWoodrow JeremyT.Kerr,CanadianFacilityforEcoinformatics WilsonSchoolofPublicandInternationalAffairs, Research(CFER),DepartmentofBiology,Universityof PrincetonUniversity,Princeton,NJ08544,USA. Ottawa,Box450,StationA,Ottawa,ON,K1N6N5, E-mail:[email protected] Canada.E-mail:[email protected] ix
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