Competition Theory in Ecology PDF
Preview Competition Theory in Ecology
OxfordSeriesinEcologyandEvolution TheComparativeMethodinEvolutionaryBiology PaulH.HarveyandMarkD.Pagel TheCauseofMolecularEvolution JohnH.Gillespie DunnockBehaviourandSocialEvolution N.B.Davies NaturalSelection:Domains,Levels,andChallenges GeorgeC.Williams BehaviourandSocialEvolutionofWasps:TheCommunalAggregationHypothesis YosiakiItô LifeHistoryInvariants:SomeExplorationsofSymmetryinEvolutionaryEcology EricL.Charnov QuantitativeEcologyandtheBrownTrout J.M.Elliott SexualSelectionandtheBarnSwallow AndersPapeMøller EcologyandEvolutioninAnoxicWorlds TomFenchelandBlandJ.Finlay AnolisLizardsoftheCaribbean:Ecology,Evolution,andPlateTectonics JonathanRoughgarden FromIndividualBehaviourtoPopulationEcology WilliamJ.Sutherland EvolutionofSocialInsectColonies:SexAllocationandKinSelection RossH.CrozierandPekkaPamilo BiologicalInvasions:TheoryandPractice NanakoShigesadaandKohkichiKawasaki CooperationAmongAnimals:AnEvolutionaryPerspective LeeAlanDugatkin NaturalHybridizationandEvolution MichaelL.Arnold TheEvolutionofSiblingRivalry DouglasW.MockandGeoffreyA.Parker Asymmetry,DevelopmentalStability,andEvolution AndersPapeMøllerandJohnP.Swaddle MetapopulationEcology IlkkaHanski DynamicStateVariableModelsinEcology:MethodsandApplications ColinW.ClarkandMarcMangel TheOrigin,Expansion,andDemiseofPlantSpecies DonaldA.Levin TheSpatialandTemporalDynamicsofHost–ParasitoidInteractions MichaelP.Hassell TheEcologyofAdaptiveRadiation DolphSchluter ParasitesandtheBehaviorofAnimals JaniceMoore EvolutionaryEcologyofBirds PeterBennettandIanOwens TheRoleofChromosomalChangeinPlantEvolution DonaldA.Levin LivinginGroups JensKrauseandGraemeD.Ruxton StochasticPopulationDynamicsinEcologyandConservation RussellLande,SteinerEngen,andBernt-ErikSæther TheStructureandDynamicsofGeographicRanges KevinJ.Gaston AnimalSignals JohnMaynardSmithandDavidHarper EvolutionaryEcology:TheTrinidadianGuppy AnneE.Magurran InfectiousDiseasesinPrimates:Behavior,Ecology,andEvolution CharlesL.NunnandSoniaAltizer ComputationalMolecularEvolution ZihengYang TheEvolutionandEmergenceofRNAViruses EdwardC.Holmes Aboveground–BelowgroundLinkages:BioticInteractions,EcosystemProcesses,andGlobal Change RichardD.BardgettandDavidA.Wardle PrinciplesofSocialEvolution AndrewF.G.Bourke MaximumEntropyandEcology:ATheoryofAbundance,Distribution,and Energetics JohnHarte EcologicalSpeciation PatrikNosil EnergeticFoodWebs:AnAnalysisofRealandModelEcosystems JohnC.MooreandPeterC.deRuiter EvolutionaryBiomechanics:Selection,Phylogeny,andConstraint GrahamK.TaylorandAdrianL.R.Thomas QuantitativeEcologyandEvolutionaryBiology:IntegratingModelswithData OtsoOvaskainen,HenrikJohandeKnegt,andMariadelMarDelgado MitonuclearEcology GeoffreyE.Hill TheEvolutionaryBiologyofSpecies TimothyG.Barraclough GameTheoryinBiology:ConceptsandFrontiers JohnM.McNamaraandOlofLeimar AdaptationandtheBrain SusanD.Healy CompetitionTheoryinEcology PeterA.Abrams Competition Theory in Ecology PETER A. ABRAMS DepartmentofEcologyandEvolutionaryBiology,UniversityofToronto,Canada GreatClarendonStreet,Oxford,OX26DP, UnitedKingdom OxfordUniversityPressisadepartmentoftheUniversityofOxford. ItfurtherstheUniversity’sobjectiveofexcellenceinresearch,scholarship, andeducationbypublishingworldwide.Oxfordisaregisteredtrademarkof OxfordUniversityPressintheUKandincertainothercountries ©PeterA.Abrams2022 Themoralrightsoftheauthorhavebeenasserted Impression:1 Allrightsreserved.Nopartofthispublicationmaybereproduced,storedin aretrievalsystem,ortransmitted,inanyformorbyanymeans,withoutthe priorpermissioninwritingofOxfordUniversityPress,orasexpresslypermitted bylaw,bylicenceorundertermsagreedwiththeappropriatereprographics rightsorganization.Enquiriesconcerningreproductionoutsidethescopeofthe aboveshouldbesenttotheRightsDepartment,OxfordUniversityPress,atthe addressabove Youmustnotcirculatethisworkinanyotherform andyoumustimposethissameconditiononanyacquirer PublishedintheUnitedStatesofAmericabyOxfordUniversityPress 198MadisonAvenue,NewYork,NY10016,UnitedStatesofAmerica BritishLibraryCataloguinginPublicationData Dataavailable LibraryofCongressControlNumber:2022936370 ISBN978–0–19–289552–3(hbk) ISBN978–0–19–289553–0(pbk) DOI:10.1093/oso/9780192895523.001.0001 Printedandboundby CPIGroup(UK)Ltd,Croydon,CR04YY LinkstothirdpartywebsitesareprovidedbyOxfordingoodfaithand forinformationonly.Oxforddisclaimsanyresponsibilityforthematerials containedinanythirdpartywebsitereferencedinthiswork. Acknowledgements Thisbookarosefromaplannedreviewarticlethatgrewtoolong.CharleyKrebsand TomSchoenerencouragedmetoturnitintoabook.ItwasanideathatIhadresisted forquitesometime,sothebookwouldlikelynothavecomeintoexistencewithout theirinfluence.Tomalsousedsomepreliminarychapterdraftsinagraduateseminar course,whichfurnishedsomemuch-neededmotivationforspeedingupmywriting schedule.ChrisKlausmeier,BobHolt,andMarkMcPeekearlyonprovidedvaluable feedback,andMichaelCortezsentcommentsonthelongestchapter.Iamgrateful to Ian Sherman, Charlie Bath, and Giulia Lipparini at Oxford University Press for theirinputintothisproject.Iamalsoverymuchindebtedtomywife,JanetPelley, forherforbearanceasIspentaninordinateamountoftimewritingthiswork,and forprovidingfeedbackandcopyeditingonmanyofthechapters.Thebookwaswrit- tenduringthefirsttwoyearsoftheSARS-CoV2epidemic,whichlimitedaccessto libraries and people who could have made the writing process easier. Weekly dis- cussions with Tom Reimchen, Don Kramer, and Larry Dill have helped keep me informedofrecentdevelopmentsinecologyandevolutionduringthatperiod.The Natural Sciences and Engineering Research Council of Canada provided financial supportthroughaDiscoveryGrant. Contents 1 Introduction: competition theory past and present 1 1.1 Ashorthistory 1 1.2 Theneedforresourcesincompetitiontheory 3 1.3 TheLotka–Volterramodel 4 1.4 Thefirstrevivalofcompetitiontheory 7 1.5 Recentcompetitiontheory 8 1.6 Generalthemesofthisbook 9 1.7 Necessarybackgroundandalookahead 12 2 Defining and describing competition 13 2.1 Introduction 13 2.2 Historicaldefinitionsofcompetition 14 2.3 Whatshouldthedefinitionbe? 22 2.4 Implicationsofthedefinition 24 2.5 Competitionwithintheframeworkoffoodwebs 29 3 Measuring and describing competition: a consumer–resource framework 33 3.1 Themeasurementofcompetition(andotherinteractions) 33 3.2 Methodsofmeasuringanddescribingcompetition 36 3.3 Argumentsagainstresource-baseddefinitionsandmodels 40 3.4 MacArthur’sconnectionofLVtoconsumer–resourcemodels 42 3.5 Whatdocoexistenceandexclusionmean? 47 3.6 Whatdistinguishesasingleresourcefromothers? 49 3.7 Functionalformsforthemodelcomponents 49 3.7.1 Resourcegrowth 52 3.7.2 Consumerfunctionalresponses 53 3.7.3 Consumernumericalresponses 56 3.8 Analysisofmodelsofcompetition 58 3.9 Summary 61 viii • Contents 4 Competition theory: its present state 63 4.1 Introduction 63 4.2 Questionsforassessingrecentinfluentialtheory 64 4.3 Choosingarticlestorepresentcurrentcompetitiontheory 66 4.4 Forgottenresultsin‘moderncompetitiontheory’ 70 4.5 WhytheLotka–VolterraandMacArthurmodelsareinsufficient 72 4.6 Reasonsforincludingresourcedynamics 74 4.7 Appendix:Problematicfeaturesinthefocalarticles 75 5 Understanding intraspecific and apparent competition 79 5.1 Introduction 79 5.2 Intraspecificcompetition 80 5.2.1 Thedefinitionandmechanismofintraspecific competition 81 5.2.2 Describing,measuring,andmodellingintraspecific competition 82 5.2.3 Modelsofdensitydependence 86 5.2.4 One-consumer–multi-resourcesystems 93 5.2.5 Amoremechanisticapproachtodensitydependence 101 5.3 Apparentcompetition 101 6 The negativity, constancy, and continuity of competitive effects 109 6.1 Introduction 109 6.2 Resourceextinctionandquasi-extinctioninMacArthur’smodel 111 6.3 Consequencesofnon-logisticresourcegrowth 122 6.4 Consequencesofnonlinearfunctionalresponses 125 6.4.1 Effectsofnonlinearfunctionalresponsesonconsumer competitioninsystemswithstableequilibria 126 6.4.2 InteractionsinunstablesystemswithtypeIIresponses 132 6.5 Interdependenceofcompetitiveeffectswithmoreconsumers 137 6.6 Otherneglectedaspectsofconsumer–resourcemodels 138 7 Resource use and the strength of interspecific competition 143 7.1 Theoryregardingthestrengthofcompetition 143 7.1.1 Theoryfromtheearly1970s 144 7.1.2 EarlyquestioningofMacArthur’slimitingsimilarity 147 Contents • ix 7.1.3 Recentandpotentialfuturetheoryonoverlapand competition 149 7.1.4 Continueduseofoutdatedsimilarity–competition relationships 153 7.2 Laboratorystudiesofcompetition 154 7.3 Fieldstudiesofcompetition 155 7.3.1 Ahistoricalreview 156 7.3.2 Anillustrativeexample:competitionbetweenhermit crabs 157 7.3.3 Currentstatusoffieldstudiesofcompetition 159 7.4 Doescompetitiveneutralityoccur? 160 7.5 Interspecificcompetitioninafoodwebcontext 162 7.6 Competitionbetweenspeciesintheoryandreality 169 8 Competition in seasonal environments: temporal overlap 171 8.1 Introduction 171 8.1.1 Abriefhistoryofworkonseasonalcompetition 172 8.1.2 Aspectsofseasonalvariationincompetitiontreatedhere 174 8.1.3 Whyarethedynamicsofseasonalsystemsimportant? 175 8.2 AmodellingframeworkandaseasonalMacArthursystem 176 8.2.1 Generalfeaturesofthemodels 176 8.2.2 ResourcelagsandmutualinvasibilityofMacArthur systems 178 8.2.3 Coexistenceina2-consumerMacArthursystem 184 8.2.4 Howrobustandrepresentativeistheexample? 188 8.2.5 Coexistenceofaseasonalandanaseasonalconsumer 189 8.2.6 Amorecompletedescriptionofseasonalinteractions 192 8.2.7 Seasonalityresourceconversionefficiency,b 195 8.2.8 A3-consumersystemwithvariationinc 197 8.2.9 A2-resourcesystemwithtemporalandnon-temporal partitioning 199 8.3 Competitioninother2-consumer–1-resourcemodels 200 8.3.1 Systemswithabioticresources 201 8.3.2 BioticresourceswithtypeIIfunctionalresponses 203 8.3.3 AbioticresourceswithtypeIIfunctionalresponses 204 8.4 Discussion 204 9 Relative nonlinearity and seasonality 209 9.1 Introduction 209 9.2 Inherentlyunstableconsumer–resourceinteractions 210 9.3 Differencesinnonlinearitywithseasonalresourcegrowth 213 9.4 Differencesinthenonlinearityofnumericalresponses 222
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