Origin, Trajectories, and Correlations of Altruism and Mobility

60 Pages·2005·0.67 MB·English

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Interim Report IR-05-020
Adaptive Evolution of Social Traits:
Origin, Trajectories, and Correlations of Altruism and Mobility

Jean-François Le Galliard ([email protected])
Régis Ferrière ([email protected])
Ulf Dieckmann ([email protected])

Approved by
Leen Hordijk
Director, IIASA
March 2005

Interim Reports on work of the International Institute for Applied Systems Analysis receive only
limited review. Views or opinions expressed herein do not necessarily represent those of the
Institute, its National Member Organizations, or other organizations supporting the work.
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No.1 Metz JAJ, Geritz SAH, MeszénaG, JacobsFJA, van No.11 GeritzSAH,MetzJAJ,KisdiÉ,MeszénaG: TheDy-
HeerwaardenJS: AdaptiveDynamics: AGeometricalStudy namicsof AdaptationandEvolutionaryBranching. IIASA
oftheConsequencesofNearlyFaithfulReproduction. IIASA WorkingPaperWP-96-077(1996). PhysicalReviewLetters
WorkingPaperWP-95-099(1995). vanStrienSJ,Verduyn 78:2024-2027(1997).
LunelSM(eds):StochasticandSpatialStructuresofDynami-
No.12 Geritz SAH, Kisdi É, MeszénaG, Metz JAJ: Evo-
calSystems,ProceedingsoftheRoyalDutchAcademyofSci-
lutionary Singular Strategies and the Adaptive Growth and
ence (KNAW Verhandelingen), North Holland, Amsterdam,
BranchingoftheEvolutionaryTree. IIASA WorkingPaper
pp.183-231(1996).
WP-96-114(1996). EvolutionaryEcology12:35-57(1998).
No.2 DieckmannU,LawR: TheDynamicalTheoryofCo-
No.13 Heino M, Metz JAJ,Kaitala V: Evolutionof Mixed
evolution:ADerivationfromStochasticEcologicalProcesses.
Maturation Strategies in Semelparous Life-Histories: The
IIASAWorkingPaperWP-96-001(1996). JournalofMathe-
Crucial Role of Dimensionality of Feedback Environment.
maticalBiology34:579-612(1996).
IIASA Working Paper WP-96-126 (1996). Philosophi-
No.3 Dieckmann U, Marrow P, Law R: EvolutionaryCy- cal Transactions of the Royal Society of London Series B
clingofPredator-PreyInteractions:PopulationDynamicsand 352:1647-1655(1997).
theRedQueen. IIASAPreprint(1995). JournalofTheoreti-
No.14 Dieckmann U: Can Adaptive Dynamics Invade?
calBiology176:91-102(1995).
IIASA WorkingPaperWP-96-152(1996). TrendsinEcol-
No.4 Marrow P, Dieckmann U, Law R: EvolutionaryDy- ogyandEvolution12:128-131(1997).
namicsofPredator-PreySystems:AnEcologicalPerspective.
No.15 MeszénaG,CzibulaI,GeritzSAH: AdaptiveDynam-
IIASAWorkingPaperWP-96-002(1996). JournalofMathe-
icsina2-PatchEnvironment: ASimpleModelforAllopatric
maticalBiology34:556-578(1996).
andParapatricSpeciation. IIASAInterimReportIR-97-001
No.5 LawR,MarrowP,DieckmannU: OnEvolutionunder (1997). JournalofBiologicalSystems5:265-284(1997).
AsymmetricCompetition. IIASAWorkingPaperWP-96-003
No.16 Heino M, Metz JAJ, Kaitala V: The Enigma of
(1996). EvolutionaryEcology11:485-501(1997).
Frequency-DependentSelection. IIASA Interim ReportIR-
No.6 MetzJAJ,MyliusSD,DiekmannO: WhenDoesEvo- 97-061(1997). TrendsinEcologyandEvolution13:367-370
lution Optimize? Onthe RelationBetweenTypesof Density (1998).
DependenceandEvolutionarilyStableLife HistoryParame-
No.17 Heino M: Management of Evolving Fish Stocks.
ters. IIASAWorkingPaperWP-96-004(1996).
IIASAInterim ReportIR-97-062(1997). CanadianJournal
No.7 Ferrière R, Gatto M: Lyapunov Exponents and the ofFisheriesandAquaticSciences55:1971-1982(1998).
Mathematics of Invasion in Oscillatory or Chaotic Popula-
No.18 HeinoM: EvolutionofMixedReproductiveStrategies
tions. TheoreticalPopulationBiology48:126-171(1995).
inSimpleLife-HistoryModels. IIASAInterimReportIR-97-
No.8 Ferrière R, Fox GA: ChaosandEvolution. IIASA 063(1997).
Preprint (1996). Trendsin EcologyandEvolution10:480-
No.19 GeritzSAH,vanderMeijdenE,MetzJAJ: Evolution-
485(1995).
aryDynamicsofSeedSizeandSeedlingCompetitiveAbility.
No.9 FerrièreR,MichodRE: TheEvolutionofCooperation IIASAInterimReportIR-97-071(1997). TheoreticalPopu-
inSpatiallyHeterogeneousPopulations. IIASAWorkingPa- lationBiology55:324-343(1999).
per WP-96-029(1996). TheAmericanNaturalist147:692-
No.20 GalisF,MetzJAJ: WhyAreThereSoManyCichlid
717(1996).
Species? OntheInterplayof SpeciationandAdaptiveRadi-
No.10 vanDoorenTJM,MetzJAJ: DelayedMaturationin ation. IIASA InterimReportIR-97-072(1997). Trendsin
TemporallyStructuredPopulationswithNon-EquilibriumDy- EcologyandEvolution13:1-2(1998).
namics. IIASAWorkingPaperWP-96-070(1996). Journal
ofEvolutionaryBiology11:41-62(1998).
No.21 Boerlijst MC, NowakMA, Sigmund K: Equal Pay No.37 Gyllenberg M, Metz JAJ: On Fitness in Structured
for all Prisoners/TheLogicof Contrition. IIASA Interim Metapopulations. IIASAInterim ReportIR-99-037(1999).
Report IR-97-073(1997). AmericanMathematicalSociety JournalofMathematicalBiology43:545-560(2001).
Monthly104:303-307(1997). JournalofTheoreticalBiology
No.38 MeszénaG,MetzJAJ: SpeciesDiversityandPopula-
185:281-293(1997).
tionRegulation: TheImportanceofEnvironmentalFeedback
No.22 LawR,DieckmannU: SymbiosisWithoutMutualism Dimensionality. IIASAInterimReportIR-99-045(1999).
andtheMergerofLineagesinEvolution. IIASAInterimRe-
No.39 Kisdi É, Geritz SAH: Evolutionary Branching and
portIR-97-074(1997). ProceedingsoftheRoyalSocietyof
SympatricSpeciationinDiploidPopulations. IIASAInterim
LondonSeriesB265:1245-1253(1998).
ReportIR-99-048(1999).
No.23 KlinkhamerPGL,deJongTJ,MetzJAJ: SexandSize
No.40 Ylikarjula J, Heino M, DieckmannU: Ecologyand
inCosexualPlants. IIASAInterimReportIR-97-078(1997).
AdaptationofStuntedGrowthinFish. IIASAInterimReport
TrendsinEcologyandEvolution12:260-265(1997).
IR-99-050(1999). EvolutionaryEcology13:433-453(1999).
No.24 FontanaW, SchusterP: ShapingSpace: ThePossi-
No.41 NowakMA,SigmundK: GamesonGrids. IIASA
bleandtheAttainableinRNAGenotype-PhenotypeMapping.
Interim Report IR-99-038 (1999). Dieckmann U, Law R,
IIASAInterimReportIR-98-004(1998). JournalofTheoret-
Metz JAJ (eds): The Geometry of Ecological Interactions:
icalBiology194:491-515(1998).
SimplifyingSpatialComplexity,CambridgeUniversityPress,
No.25 Kisdi É, Geritz SAH: Adaptive Dynamics in Allele Cambridge,UK,pp.135-150(2000).
Space: Evolution of Genetic Polymorphismby Small Muta-
No.42 Ferrière R, Michod RE: Wave Patterns in Spatial
tionsinaHeterogeneousEnvironment. IIASAInterimReport
Games and the Evolution of Cooperation. IIASA Interim
IR-98-038(1998). Evolution53:993-1008(1999).
ReportIR-99-041(1999). DieckmannU,LawR,MetzJAJ
No.26 FontanaW,SchusterP: ContinuityinEvolution: On (eds): TheGeometryofEcologicalInteractions: Simplifying
theNatureofTransitions. IIASAInterim ReportIR-98-039 SpatialComplexity,CambridgeUniversityPress,Cambridge,
(1998). Science280:1451-1455(1998). UK,pp.318-332(2000).
No.27 NowakMA,SigmundK: EvolutionofIndirectReci- No.43 Kisdi É, JacobsFJA,Geritz SAH: RedQueenEvo-
procity by Image Scoring/ The Dynamics of Indirect Reci- lution by Cycles of EvolutionaryBranching and Extinction.
procity. IIASA Interim ReportIR-98-040(1998). Nature IIASA Interim Report IR-00-030(2000). Selection2:161-
393:573-577(1998). JournalofTheoreticalBiology194:561- 176(2001).
574(1998).
No.44 MeszénaG,KisdiÉ,DieckmannU,GeritzSAH,Metz
No.28 KisdiÉ: EvolutionaryBranchingUnderAsymmetric JAJ: EvolutionaryOptimisationModelsandMatrixGamesin
Competition. IIASAInterimReportIR-98-045(1998). Jour- theUnifiedPerspectiveofAdaptiveDynamics. IIASAInterim
nalofTheoreticalBiology197:149-162(1999). ReportIR-00-039(2000). Selection2:193-210(2001).
No.29 BergerU: BestResponseAdaptationforRoleGames. No.45 ParvinenK,DieckmannU,GyllenbergM,MetzJAJ:
IIASAInterimReportIR-98-086(1998). EvolutionofDispersalinMetapopulationswithLocalDensity
DependenceandDemographicStochasticity. IIASAInterim
No.30 van Dooren TJM: The Evolutionary Ecology of
ReportIR-00-035(2000). JournalofEvolutionaryBiology
Dominance-Recessivity. IIASA Interim Report IR-98-096
16:143-153(2003).
(1998). JournalofTheoreticalBiology198:519-532(1999).
No.46 Doebeli M, Dieckmann U: Evolutionary Branch-
No.31 DieckmannU,O’HaraB,WeisserW: TheEvolution-
ing and Sympatric Speciation Causedby Different Types of
aryEcologyofDispersal. IIASAInterimReportIR-98-108
EcologicalInteractions. IIASA Interim Report IR-00-040
(1998). TrendsinEcologyandEvolution14:88-90(1999).
(2000). TheAmericanNaturalist156:S77-S101(2000).
No.32 SigmundK: ComplexAdaptiveSystemsandtheEvo-
No.47 Heino M, HanskiI: Evolutionof Migration Rate in
lution of Reciprocation. IIASA Interim Report IR-98-100
aSpatiallyRealisticMetapopulationModel. IIASAInterim
(1998). Ecosystems1:444-448(1998).
ReportIR-00-044(2000). TheAmericanNaturalist157:495-
No.33 PoschM, Pichler A, Sigmund K: The Efficiencyof 511(2001).
Adapting AspirationLevels. IIASA Interim Report IR-98-
No.48 GyllenbergM,ParvinenK,DieckmannU: Evolution-
103(1998). ProceedingsoftheRoyalSocietyLondonSeries
arySuicideandEvolutionofDispersalinStructuredMetapop-
B266:1427-1435(1999).
ulations. IIASAInterimReportIR-00-056(2000). Journal
No.34 MathiasA,KisdiÉ: EvolutionaryBranchingandCo- ofMathematicalBiology45:79-105(2002).
existenceofGerminationStrategies. IIASA Interim Report
No.49 vanDoorenTJM: TheEvolutionaryDynamicsofDi-
IR-99-014(1999).
rect Phenotypic Overdominance: EmergencePossible, Loss
No.35 DieckmannU,DoebeliM: OntheOriginofSpecies Probable. IIASAInterimReportIR-00-048(2000). Evolu-
bySympatricSpeciation. IIASA Interim ReportIR-99-013 tion54:1899-1914(2000).
(1999). Nature400:354-357(1999).
No.50 NowakMA,PageKM,SigmundK: FairnessVersus
No.36 MetzJAJ,GyllenbergM: HowShouldWeDefineFit- ReasonintheUltimatum Game. IIASAInterim ReportIR-
nessinStructuredMetapopulationModels?IncludinganAp- 00-57(2000). Science289:1773-1775(2000).
plicationtotheCalculationofEvolutionarilyStableDispersal
No.51 de Feo O, Ferrière R: Bifurcation Analysis of Pop-
Strategies. IIASA Interim ReportIR-99-019(1999). Pro-
ulation Invasion: On-Off Intermittency and Basin Riddling.
ceedingsof the RoyalSocietyof LondonSeries B268:499-
IIASAInterimReportIR-00-074(2000). InternationalJour-
508(2001).
nalofBifurcationandChaos10:443-452(2000).
No.52 HeinoM, Laaka-LindbergS: Clonal Dynamicsand No.66 Dercole F, Rinaldi S: Evolution of Cannibalistic
Evolution of Dormancy in the Leafy Hepatic Lophozia Sil- Traits: ScenariosDerivedfromAdaptiveDynamics. IIASA
vicola. IIASA Interim Report IR-01-018 (2001). Oikos InterimReportIR-02-054(2002). TheoreticalPopulationBi-
94:525-532(2001). ology62:365-374(2002).
No.53 SigmundK,HauertC,NowakMA: RewardandPun- No.67 Bürger R, Gimelfarb A: Fluctuating Environments
ishment in Minigames. IIASA Interim Report IR-01-031 andtheRoleofMutationinMaintainingQuantitativeGenetic
(2001). Proceedingsof the National Academyof Sciences Variation. IIASAInterimReportIR-02-058(2002). Geneti-
oftheUSA98:10757-10762(2001). calResearch80:31-46(2002).
No.54 HauertC,DeMonteS,SigmundK,HofbauerJ: Os- No.68 BürgerR: OnaGeneticModelofIntraspecificCom-
cillations in OptionalPublic GoodGames. IIASA Interim petitionandStabilizingSelection. IIASAInterimReportIR-
ReportIR-01-036(2001). 02-062(2002). Amer.Natur.160:661-682(2002).
No.55 FerrièreR,LeGalliardJ: InvasionFitnessandAdap- No.69 DoebeliM,DieckmannU: SpeciationAlongEnviron-
tiveDynamicsinSpatialPopulationModels. IIASAInterim mentalGradients. IIASAInterimReportIR-02-079(2002).
ReportIR-01-043(2001). ClobertJ,DhondtA,DanchinE, Nature421:259-264(2003).
NicholsJ(eds):Dispersal,OxfordUniversityPress,pp.57-79
No.70 DercoleF,IrissonJ,RinaldiS: BifurcationAnalysisof
(2001).
aPrey-PredatorCoevolutionModel. IIASAInterimReport
No.56 deMazancourtC,LoreauM,DieckmannU: Canthe IR-02-078(2002). SIAM JournalonAppliedMathematics
EvolutionofPlantDefenseLeadtoPlant-HerbivoreMutual- 63:1378-1391(2003).
ism. IIASAInterimReportIR-01-053(2001). TheAmerican
No.71 LeGalliardJ,FerrièreR,DieckmannU: TheAdaptive
Naturalist158:109-123(2001).
DynamicsofAltruisminSpatiallyHeterogeneousPopulations.
No.57 ClaessenD,DieckmannU: OntogeneticNicheShifts IIASAInterimReportIR-03-006(2003). Evolution57:1-17
and EvolutionaryBranching in Size-StructuredPopulations. (2003).
IIASAInterimReportIR-01-056(2001). EvolutionaryEcol-
No.72 Taborsky B, Dieckmann U, Heino M: Unex-
ogyResearch4:189-217(2002).
pectedDiscontinuities in Life-History Evolution underSize-
No.58 Brandt H: Correlation Analysis of Fitness Land- Dependent Mortality. IIASA Interim Report IR-03-004
scapes. IIASAInterimReportIR-01-058(2001). (2003). ProceedingsoftheRoyalSocietyofLondonSeriesB
270:713-721(2003).
No.59 DieckmannU: AdaptiveDynamicsofPathogen-Host
Interacations. IIASA Interim Report IR-02-007 (2002). No.73 Gardmark A, Dieckmann U, Lundberg P: Life-
Dieckmann U, Metz JAJ, Sabelis MW, Sigmund K (eds): HistoryEvolutioninHarvestedPopulations:TheRoleofNat-
AdaptiveDynamicsofInfectiousDiseases:InPursuitofViru- ural Predation. IIASA Interim Report IR-03-008 (2003).
lenceManagement,CambridgeUniversityPress,Cambridge, EvolutionaryEcologyResearch5:239-257(2003).
UK,pp.39-59(2002).
No.74 MizeraF,MeszénaG: SpatialNichePacking,Char-
No.60 Nowak MA, Sigmund K: Super- and Coinfection: acter Displacement and Adaptive Speciation Along an En-
TheTwoExtremes. IIASAInterimReportIR-02-008(2002). vironmental Gradient. IIASA Interim Report IR-03-062
Dieckmann U, Metz JAJ, Sabelis MW, Sigmund K (eds): (2003). EvolutionaryEcologyResearch5:363-382(2003).
AdaptiveDynamicsofInfectiousDiseases:InPursuitofViru-
No.75 DercoleF: RemarksonBranching-ExtinctionEvolu-
lenceManagement,CambridgeUniversityPress,Cambridge,
tionary Cycles. IIASA Interim Report IR-03-075 (2003).
UK,pp.124-137(2002).
JournalofMathematicalBiology47:569-580(2003).
No.61 Sabelis MW, Metz JAJ: Perspectivesfor Virulence
No.76 HofbauerJ,SigmundK: EvolutionaryGameDynam-
Management:RelatingTheorytoExperiment. IIASAInterim
ics. IIASAInterimReportIR-03-078(2003). Bulletinofthe
ReportIR-02-009(2002). DieckmannU,MetzJAJ,Sabelis
AmericanMathematicalSociety40:479-519(2003).
MW,SigmundK(eds):AdaptiveDynamicsofInfectiousDis-
eases: InPursuitofVirulenceManagement,CambridgeUni- No.77 Ernande B, Dieckmann U, Heino M: Adaptive
versityPress,Cambridge,UK,pp.379-398(2002). ChangesinHarvestedPopulations: PlasticityandEvolution
of AgeandSize at Maturation. IIASA Interim Report IR-
No.62 Cheptou P, Dieckmann U: The Evolution of Self-
03-058(2003). ProceedingsoftheRoyalSocietyofLondon
Fertilization inDensity-RegulatedPopulations. IIASA In-
SeriesB-BiologicalSciences,271:415-423(2004).
terim Report IR-02-024 (2002). Proceedingsof the Royal
SocietyofLondonSeriesB269:1177-1186(2002). No.78 HanskiI,HeinoM: Metapopulation-LevelAdaptation
of InsectHost Plant PreferenceandExtinction-Colonization
No.63 BürgerR: AdditiveGeneticVariationUnderIntraspe-
Dynamics in Heterogeneous Landscapes. IIASA Interim
cific Competition and Stabilizing Selection: A Two-Locus
Report IR-03-028 (2003). Theoretical Population Biology
Study. IIASA Interim ReportIR-02-013(2002). Theoret-
63:309-338(2003).
icalPopulationBiology61:197-213(2002).
No.79 vanDoornG,DieckmannU,WeissingFJ: Sympatric
No.64 HauertC,DeMonteS,HofbauerJ,SigmundK: Vol-
Speciation by Sexual Selection: A Critical Re-Evaluation.
unteeringasRedQueenMechanismforCo-operationinPub-
IIASA Interim Report IR-04-003 (2004). American Natu-
licGoodsGames. IIASAInterimReportIR-02-041(2002).
ralist163:709-725(2004).
Science296:1129-1132(2002).
No.80 EgasM, DieckmannU, SabelisMW: EvolutionRe-
No.65 DercoleF,FerrièreR,RinaldiS: EcologicalBistabil-
stricts the Coexistence of Specialists and Generalists - the
ityandEvolutionaryReversalsunderAsymmetricalCompeti-
RoleofTrade-offStructure. IIASAInterimReportIR-04-004
tion. IIASA Interim Report IR-02-053(2002). Evolution
(2004).
56:1081-1090(2002).
No.81 ErnandeB, DieckmannU: TheEvolutionof Pheno- No.88 CadetCR,MetzJAJ,KlinkhamerPGL: Sizeandthe
typicPlasticityinSpatiallyStructuredEnvironments:Implica- Not-So-SingleSex: disentanglingtheeffectsofsizeonsexal-
tionsofIntraspecificCompetition,PlasticityCosts,andEnvi- location. IIASAInterimReportIR-04-084(2004). Ameri-
ronmentalCharacteristics. IIASAInterimReportIR-04-006 canNaturalist,164:779-792(2004).
(2004). Journal of Evolutionary Biology 17 (3): 613-628
No.89 Rueffler C, van Dooren TJM, Metz JAJ: Adaptive
(2004).
WalksonChangingLandscapes:Levins’ApproachExtended.
No.82 Cressman R, Hofbauer J: Measure Dynamics on a IIASAInterimReportIR-04-083(2004). TheoreticalPopu-
One-DimensionalContinuousTraitSpace:TheoreticalFoun- lationBiology,65:165-178(2004).
dationsforAdaptiveDynamics. IIASA Interim Report IR-
No.90 deMazancourtC,LoreauM,DieckmannU: Under-
04-016(2004).
standingMutualismWhenThereisAdaptationtothePartner.
No.83 Cressman R: Dynamic Stability of the Replicator IIASAInterimReportIR-05-016(2005).
Equation with ContinuousStrategySpace. IIASA Interim
No.91 DieckmannU,DoebeliM: PluralisminEvolutionary
ReportIR-04-017(2004).
Theory. IIASAInterimReportIR-05-017(2005).
No.84 RavignéV,OlivieriI,DieckmannU: Implicationsof
HabitatChoiceforProtectedPolymorphisms. IIASAInterim No.92 DoebeliM,DieckmannU,MetzJAJ,TautzD: What
We Have Also Learned. IIASA Interim Report IR-05-018
ReportIR-04-005(2004). EvolutionaryEcologyResearch6:
(2005).
125-145(2004).
No.85 NowakMA,SigmundK: EvolutionaryDynamicsof No.93 Egas M, Sabelis MW, Dieckmann U: Evolution of
BiologicalGames. IIASAInterimReportIR-04-013(2004). Specialization and Ecological Character Displacement of
Science303:793-799(2004). HerbivoresAlong a Gradient of Plant Quality. IIASA In-
terimReportIR-05-019(2005).
No.86 VukicsA,AsbóthJ,MeszénaG: SpeciationinMul-
tidimensional Evolutionary Space. IIASA Interim Report No.94 Le Galliard J, Ferrière R, Dieckmann U: Adaptive
IR-04-028(2004). PhysicalReviewE684(2003). Evolution of Social Traits: Origin, Trajectories, and Corre-
lationsofAltruismandMobility. IIASAInterimReportIR-
No.87 deMazancourtC, DieckmannU: Trade-off Geome- 05-020(2005).
triesandFrequency-dependentSelection. IIASAInterimRe-
portIR-04-039(2004).
IssuesoftheIIASAStudiesinAdaptiveDynamicsseriescanbeobtainedatwww.iiasa.ac.at/Research/ADN/Series.htmlorby
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Contents
Abstract.............................................................................................................................3
Introduction......................................................................................................................4
Model Description............................................................................................................6
Population dynamics on social networks.....................................................................7
Evolutionary dynamics on social networks.................................................................8
Spatial invasion fitness..............................................................................................10
Selective Pressures on Mobility and Altruism...............................................................13
Separate Evolution of Mobility and Altruism................................................................17
Altruism.....................................................................................................................17
Mobility.....................................................................................................................18
Joint Evolution of Altruism and Mobility......................................................................20
Origin of altruism.......................................................................................................20
Evolutionary trajectories of social traits....................................................................21
Evolutionarily induced correlations between altruism and mobility.........................22
Discussion.......................................................................................................................24
Joint evolution versus single-trait evolution..............................................................24
Correlative patterns of social traits............................................................................26
The habitat saturation hypothesis..............................................................................29
Concluding remarks...................................................................................................32
Acknowledgements........................................................................................................33
Appendix A....................................................................................................................34
Appendix B.....................................................................................................................36
Literature Cited...............................................................................................................39
Table 1............................................................................................................................45
Figure Legends...............................................................................................................46
Figures............................................................................................................................49
1 ARTICLE
2 ADAPTIVE EVOLUTION OF SOCIAL TRAITS: ORIGIN, TRAJECTORIES, AND
3 CORRELATIONS OF ALTRUISM AND MOBILITY
4
5 Jean-François Le Galliard
6 Fonctionnement et évolution des systèmes écologiques, CNRS UMR 7625
7 Ecole Normale Supérieure, 75005 Paris, France
8 Present address: Center for Ecological and Evolutionary Synthesis, University of Oslo
9 P.O. Box 1050, Blindern, Oslo NO-0316, Norway
10 E-mail: [email protected]
11
12 Régis Ferrière
13 1) Fonctionnement et évolution des systèmes écologiques, CNRS UMR 7625
14 Ecole Normale Supérieure, 75005 Paris, France
15 E-mail: [email protected]
16 2) Department of Ecology and Evolutionary Biology
17 University of Arizona, Tucson AZ 85721, USA
18
19 Ulf Dieckmann
20 Adaptive Dynamics Network, International Institute for Applied Systems Analysis
21 Schlossplatz 1, A-2361 Laxenburg, Austria
22 E-mail: [email protected]
1
1 Keywords
2 Evolutionary dynamics, spatial invasion fitness, altruism, mobility, habitat saturation, kin selec-
3 tion.
4
5 Running head
6 Evolution of altruism and mobility.
7
8 List of elements to appear in the Expanded Online Edition
9 Online Appendix A
10 Online Appendix B
2
1 ABSTRACT
2 Social behavior involves “staying and helping”, two individual attributes that vary considerably
3 among organisms. Investigating the ultimate causes of such variation, this study integrates previ-
4 ously separate lines of research by analyzing the joint evolution of altruism and mobility. We
5 unfold the network of selective pressures and derive how these depend on physiological costs,
6 eco-evolutionary feedbacks, and a complex interaction between the evolving traits. Our analysis
7 highlights habitat saturation, both around individuals (local aggregation) and around unoccupied
8 space (local contention), as the key mediator of altruism and mobility evolution. Once altruism
9 and mobility are allowed to evolve jointly, three general insights emerge. First, the cost of mobil-
10 ity affects the origin of altruism, determining whether and how quickly selfishness is overcome.
11 Second, the cost of altruism determines which of two qualitatively different routes to sociality are
12 taken: an evolutionary reduction of mobility, resulting in higher habitat saturation, is either pre-
13 ceded or followed by the adaptive rise of altruism. Third, contrary to conventional expectations, a
14 positive correlation between evolutionarily stable levels of altruism and mobility can arise; this is
15 expected when comparing populations that evolved under different constraints on mobility or that
16 differ in other life history traits.
3
1 INTRODUCTION
2 Sociality is an essential characteristic of life. It involves specific individual behaviors that lead to
3 the emergence of collective properties, new levels of natural selection, and the adaptive com-
4 plexification of living systems (Michod 1999). One of the intriguing features of sociality is that it
5 causes a double cost to individuals. Sociality typically requires, first, some form of altruistic be-
6 havior through which individuals sacrifice their own fitness for the benefit of others (Hamilton
7 1964a, b), and second, some reduction in individual mobility, allowing for sustained interaction,
8 which exacerbates competition for local resources (Frank 1995; Perrin and Lehmann 2001). The
9 benefits associated with these costs must be substantial enough that the involved genes are not
10 eliminated by natural selection. Thus, one of the challenges facing evolutionary theory is to ex-
11 plain the role of adaptive evolution in molding individual altruism along with the underlying
12 population structure, to help us understand the wide diversity of social systems observed in the
13 wild (Choe and Crespi 1997; Crespi 2001).
14 The double cost of sociality reflects only some of the selective pressures acting on social
15 traits. Low individual mobility may increase genetic relatedness between interacting individuals,
16 thus promoting inbreeding as well as the evolution of helping behaviors through kin selection
17 (Hamilton 1964b). Yet, the enhancement of neighbors’ performance through altruistic interaction
18 may induce also habitat saturation and thus exacerbate local competition among kin (Grafen
19 1984; Queller 1992). Increased competition between relatives for local resources can in turn re-
20 duce or even totally negate the indirect genetic benefits of altruism (Taylor 1992; Wilson et al.
21 1992). The deleterious effects of kin competition resulting from low mobility have been demon-
22 strated in a recent comparative study of social traits in fig wasps. In these insects, strict philopatry
4