Competition Theory in Ecology

337 Pages·2022·5.915 MB·English

by  Peter A. Abrams

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CompetitionTheoryinEcology
PeterA.Abrams
Competition Theory in
Ecology
PETER A. ABRAMS
DepartmentofEcologyandEvolutionaryBiology,UniversityofToronto,Canada
GreatClarendonStreet,Oxford,OX26DP,
UnitedKingdom
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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