Table Of ContentOUPCORRECTEDPROOF – FINAL,25/5/2020,SPi
TheoreticalEcology
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OUPCORRECTEDPROOF – FINAL,25/5/2020,SPi
Theoretical Ecology
Concepts and Applications
EDITED BY
Kevin S. McCann and Gabriel Gellner
DepartmentofIntegrativeBiology,UniversityofGuelph,Canada
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Foreword
Robert D.Holt
The word “theory” in its broadest sense has to reflecting maturation of our discipline. In the cur-
dowithaframeworkofconceptualconstructsthat rent volume there is much more emphasis than in
help explain or interpret phenomena in a particu- 1976onthekeyroletheinternalstructuresofpopu-
lar domain (Pickett et al. 2007). The discipline of lationsplayintheirdynamics,andonstochasticity,
ecology has in some of its branches robust theory lags, and transient dynamics—all crucial ingredi-
thatisnotparticularlymathematical(Scheinerand entsinbringingtobeartheoryonimportantapplied
Willig 2011). More narrowly construed, however, questions.Thereismuchmoreemphasisinthecur-
“theoreticalecology”pertainstotheuseofmathe- rentvolumeonaspectsofecologicalcomplexitythat
matical and computational models to explore eco- canberenderedasnetworks,oninteractionsamong
logicalquestions,rangingfromindividualbehavior, species,onparasitesinflictinghosts,andonflowsof
throughtheintertwinedcomplexitiesofcommuni- individualsacrossspace.Acentralthemeinmany
ties, to even global-scale phenomena. This is the chaptersiselucidatingtherelationshipbetweennet-
senseinwhichRobertMayusedthetermincrafting workstructure,andthestabilityandrobustnessof
theseriesofadvancedtextstitled“TheoreticalEcol- ecologicalsystems—anabidingthemeinMay’sown
ogy,”aninfluentiallineageofvolumesnowstretch- researchcontributions.
ingbacknearlyhalfacentury.Ahistorianofecology In1976,onechapterwasonschistosomiasis;now,
could glean much insight into the development of emphasis is placed on the community context of
ecologyatitsconceptualcorebycarefullyperusing infectiousdisease.Appliedissues(e.g.,schistosomi-
thecontentsofthesevolumes,startingwiththefirst asisasacasestudy)in1976weresomewhatloosely
edition in 1976, and now manifest in the current tiedtoecologicaltheory,butinthepresentvolume,
fourthedition.Mayhimselfhasplayedacentralrole issues such as the role of diversity in infectious
inthematurationoftheoreticalecology,andevenin disease dynamics, abiotic drivers of interspecific
thisvolumenearlyallthechaptersciteoneormore interactions, and the quest for early warning sig-
ofhismanypapersandbooks. nalsofimpendingradicalchange,squarelybuildon
As noted by the editors, comparing the table other areas of ecological theory. The 1976 volume
of contents of the timely and authoritative book nodded towards evolution, for instance, with dis-
you have before you, with the 1976 first edition, cussions of bionomic strategies and sociobiology;
is quite revealing. They both start with a focus thecurrentvolumeincludesthemessuchasadap-
onsinglespeciesdynamicsandinteractionsamong tive dynamics as tools for examining trait evo-
a small number of species, then build to multi- lution in a community context. One could well
speciesensemblesbeforeendingwithapplications, imagine that the fusion of ecology and evolution
andareall writtenbyleadingauthorities. Butitis will become a central theme in yet future editions
instructive to ponder the differences in emphases, of this renowned text (see Pásztor et al. 2016 for a
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vi FOREWORD
perceptive synthesis, grounding ecological theory biodiversity on our planet mandates a clear and
onDarwinianprinciples). rigorous understanding of the factors that govern
In the final chapter in this volume, the editors their persistence and spatial arrangements and
reflect on likely “Areas of Current and Future temporalfluxes,inthefirstplace.
Growth,” specifically noting seasonal drivers The ideas summarized in this splendid volume,
of dynamics, ecosystem issues, coupled social- carryingon the grand tradition Robert Mayestab-
ecological systems, and grappling with the many lishedstartingin1976—fromcoexistencetheory,to
dimensions of change in the world. In closing, I network analyses, to embracing stochasticity and
wanttoreturntoRobertMay’sownclosingchapter lags, to grounding theory on an underpinning of
in the third 2007 edition of Theoretical Ecology (co- keyindividualtraitsandsubtleissuesofpopulation
edited with Angela McLean), titled “Unanswered structure—areessentialingredientsneededforsuch
questions and why they matter,” which mulls on understanding.Allactiveecologists,fromstudents
the consequences of the current largest driver of toseasonedpractitioners,wouldbenefitfromread-
change on the planet—ourselves. May remarks on inganddigestingthechaptersinthisvolume,andI
thestartlingsingularityofourpointintime,when applaudtheeditorsandauthorsonthefruitsoftheir
overafewgenerationsourspecieshasburgeonedin labors.
numbers,withcorrespondingimpactsontheplanet.
May cites an estimate that will reach 9 billion by
2050;themodalprojectionoftoday’sdemographic References
models worrisomely is even higher, at 9.8 billion.
Backin1800,wellwithinthelifespanofmanyforest Pásztor, L., Z. Botta-Dukát, G. Magyar, T. Czárán, and
trees still standing, there were only 1 billion or so G. Meszéna. 2016. Theory-based Ecology: A Darwinian
Approach.Oxford,UK:OxfordUniversityPress.
ofus.Maythenremarksthatbysomeestimates,we
Pickett,S.T.A.,J.Kolasa,andC.G.Jones.2007.Ecological
have already exceeded the “ecological footprint”
Understanding: The Nature of Theory and the Theory of
ofsustainability,andendswithreflectionson“the
Nature. 2nd edition. Burlington NY: Academic Press,
aspiration of optimizing the preservation of our
Elsevier.
evolutionaryheritage.”Theethicalgoalofpassing Scheiner,S.M.andM.R.Willig.2011.TheTheoryofEcology.
on to future generations some of the richness of ChicagoIL:UniversityofChicagoPress.
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Detailed table of contents
Listofcontributors xiii
1 Introduction 1
GabrielGellner,KevinS.McCann,andEmilyJ.Champagne
1.1 Thisbookanditspredecessors 1
1.2 Thisbook:Themesanddirections 2
2 Speciescoexistence 5
PeterChesson
2.1 Introduction 5
2.2 Models 6
2.3 Overallinteractionandaveragefitnessdifferences 11
2.4 Competitionforresources 12
2.5 Roleofnaturalenemies 15
2.6 Roleofenvironmentalvariation 18
2.7 Discussion 21
Acknowledgments 24
References 24
3 Thesynergisticeffectsofinteractionstrengthandlagsonecologicalstability 28
GabrielGellner,KevinS.McCann,andChristopherGreyson-Gaito
3.1 Introduction 28
3.2 Populationmodels:Theinteractiveroleofgrowthandlags 31
3.3 Consumer-Resourcemodels:TheinteractiveroleofISandlags 34
3.4 Lagexcitationandlaginterference 36
3.5 Asynchronyasaformoflaginterference? 36
3.6 Summary 38
References 38
4 Non-equilibriumdynamicsandstochasticprocesses 40
KarenC.Abbott
4.1 Introductiontostochasticityandtransients 40
4.2 Challenge1:Stabilityinstochasticecologicalsystems 42
4.2.1 Whyisthisachallengefornon-equilibriumsystems? 42
4.2.2 Awayforward 44
4.2.3 LessonfromChallenge1:Non-equilibriumdynamicsstrengthen
ecologicalunderstanding 46
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viii DETAILED TABLE OF CONTENTS
4.3 Challenge2:Predictingregimeshifts 46
4.3.1 Whyisthisachallengefornon-equilibriumsystems? 46
4.3.2 Awayforward 46
4.3.3 LessonfromChallenge2:Unstableequilibriacanrevealalotabout
non-equilibriumdynamics 49
4.4 Buildingontheselessonstoconfrontfuturechallenges 50
References 51
5 Theimpactofpopulationstructureonpopulationandcommunitydynamics 53
AndréM.deRoos
5.1 Introduction 53
5.2 StateconceptsinSPMs 54
5.3 Typesofstructuredpopulationmodels 55
5.4 Ecologicalconsequencesofchangingpopulationstructure 60
5.4.1 Juvenileandadult-drivenpopulationcycles 60
5.4.2 Biomassovercompensation 62
5.4.3 Communityconsequencesofbiomassovercompensation 64
5.5 Interfacingtheoryanddata 67
5.6 Ongeneralityandmodelspecificity 69
5.7 Outlook 70
References 71
6 Modelsforlargeecologicalcommunities—arandommatrixapproach 74
StefanoAllesinaandJacopoGrilli
6.1 Introduction 74
6.2 May’sstabilitycriterion 75
6.3 Randommatrices 78
6.4 Fundamentalresults 79
6.5 Structuredrandommatrices 83
6.6 Otherapplications 88
6.7 Openproblemsandconclusions 89
References 91
7 Astructuraltheoryofmutualisticnetworks 93
JordiBascompteandAntonioFerrera
7.1 Introduction 93
7.2 Apurelydynamicstabilityapproachtomutualisticnetworks 95
7.2.1 Earlymodels 95
7.2.2 Addingnon-linearfunctionalresponses 95
7.2.3 Addinginterspecificcompetitionwithinsets 97
7.3 Astructuralstabilityapproachtomutualisticnetworks 99
7.3.1 Preliminaryworkonthelimitstothenumberofcoexisting
speciesinpurelycompetitivesystems 99
7.3.2 Limitstothenumberofcoexistingspeciesinsystemswith
competitionplusmutualism 104
7.3.3 Robustnessofmutualisticnetworks 108
7.4 Concludingremarks 113
References 114
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DETAILED TABLE OF CONTENTS ix
8 Adata-drivenapproachtocomplexecologicalsystems 116
MichioKondoh,KazutakaKawatsu,YutakaOsada,andMasayukiUshio
8.1 Interspecificinteractionsandecologicaldynamics 116
8.1.1 Populationdynamicsandinterspecificinteractions 116
8.1.2 Communitydynamicsandinterspecificinteractions 117
8.2 Natureofpopulation-levelinterspecificinteractions 119
8.2.1 Diversityinbehavioralmechanisms 119
8.2.2 Scaledependencyofinterspecificinteractions 119
8.2.3 Dynamicnatureofinterspecificinteractions 120
8.3 Howtostudyinterspecificinteractionsinnature 121
8.3.1 Identifyingpopulation-levelinteractions 121
8.3.2 Identificationofinteractionsbasedonbehaviorbyindividuals 121
8.3.3 Manipulativefieldexperiments 122
8.4 Moderndata-drivenapproachtointerspecificinteractions 123
8.4.1 Estimatingpopulation-levelinteractionsfromtime-seriesdata 123
8.4.2 ConvergentcrossmappingandmultivariateS-map 124
8.4.3 ApplicationofEDMtointeractionnetworkstudies 125
8.5 Conclusionandfuturedirections 128
References 129
9 Trait-basedmodelsofcomplexecologicalnetworks 134
UlrichBrose
9.1 Modelingcomplexecologicalnetworks 134
9.2 Allometricpopulationmodels 136
9.3 Allometricmodelsofcomplexcommunities:TheYodzis
andInnesapproach 137
9.4 Morecomplexallometries 138
9.5 Modelingthetemperature-dependenceofnetworkdynamics 139
9.6 Outlook 139
References 141
10 Ecologicalnetworks:Fromstructuretodynamics 143
SoniaKéfi
10.1 Briefintroduction 143
10.2 Whatisanetwork? 144
10.3 Networksinecology 146
10.3.1 Interactionnetworks 146
10.3.2 Towardmulti-layerinteractionnetworks 147
10.3.3 Othertypesofecologicalnetworks 147
10.3.4 Broadquestionsaskedinecologicalnetworks 149
10.4 Quantifyingnetworksstructure 149
10.4.1 Localnetworkdescriptors 150
10.4.2 Quasi-localnetworkdescriptors(intermediatedescriptionlevel) 150
10.4.3 Globalnetworkdescriptors 150
10.4.4 Extensionstomultilayernetworks 152
10.5 Structuralpropertiesofecologicalnetworks 152
10.5.1 Foodwebs 152
10.5.2 Mutualisticwebs 154
