GameTheory

KennethPrestwich

kprestwi@holycross.edu

DepartmentofBiology
CollegeoftheHolyCross
Worcester,MAUSA01610

WithAdditionalNotesandTypesettingby

KevinMitchell

mitchell@hws.edu

DepartmentofMathematicsandComputerScience

HobartandWilliamSmithColleges

Geneva,NYUSA14456

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HawkStrategy
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Gametheorymodeling:Hawks(dashedline)andDoves(solidline).
GameTheory

KennethPrestwich

kprestwi@holycross.edu

DepartmentofBiology
CollegeoftheHolyCross
Worcester,MAUSA01610

WithAdditionalNotesandTypesettingby

KevinMitchell

mitchell@hws.edu

DepartmentofMathematicsandComputerScience

HobartandWilliamSmithColleges

Geneva,NYUSA14456

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HawkStrategy
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Gametheorymodeling:Hawks(dashedline)andDoves(solidline).
1999byKennethPrestwich.Allrightsreserved.Donotreproducewithoutpermissionofthe
author.
Thisandotherrelatedmaterialsareavailableon-lineat:

http://science.holycross.edu/departments/biology/kprestwi/behavior/ESS/ESSindexf
rmset.html
Contents

1ModelingBehavior:GameandOptimalityTheory
1
1.1AdaptationversusNeutralModelsofEvolution.....................1
1.2MathematicalModelsandRigorousScience.......................2
1.3OptimalityModels.....................................2
1.4ABriefIntroductiontoGameTheory..........................6
1.5ABriefSketchoftheHistoryofGameTheory.....................7
2GameTheoryandEvolutionaryStableStrategies
9
2.1Introduction:BehavioralStrategiesandGames.....................9
2.2FormalAnalysisofaPairwiseGame...........................11
2.3Stasis:EvolutionaryStableStrategies..........................15
2.4PureESSsinTwoStrategyGames............................16
2.5MixedESSsforaTwoStrategyGame..........................20
2.6CanDierentStrategiesCo-ExistWithoutanESS?..................22
3HawksandDoves
23
3.1Introduction.........................................23
3.2EvolutionaryStableStrategies:AnExample.......................24
3.3Preliminary(Qualitative)Exploration..........................25
3.4FormalAnalysisoftheHawk-DoveESS..........................26
3.5TheFitnessofEachStrategyandMixedESSs.....................29
4TheHawksandDovesSimulation
32
4.1WhattheSimulationDoes.................................32
4.2DierencesBetweentheAppletandApplication....................38
4.3QuestionstoAddressandThingstoTry.........................38
4.4Appendix:TheConceptofFitness............................39
ictandOwnership:TheBourgeoisStrategy
5Con
41
5.1Introduction.........................................41
5.2De
nitionoftheBourgeoisStrategy...........................42
5.3PayosfortheBourgeois,Dove,andHawkGame....................42
5.4IsBourgeoisaPureESS?.................................43
CONTENTS
ii
6AThreeStrategySimulation
44
6.1AbouttheSimulation...................................44
6.2DierencesBetweentheAppletandApplication....................49
6.3QuestionstoAddressandThingstoTry.........................49
7SupplementaryMaterialfortheHv.DandHv.Dv.BGames
51
7.1MoreontheHawkandDoveGame............................51
7.2MoreontheHawk,Dove,andBourgeoisGame.....................54
8WarsofAttrition:FixedCostStrategies
58
8.1Introduction.........................................58
8.2WaitingGamesandtheirCurrencies...........................59
8.3IsthereanESSforaWarofAttrition?..........................60
8.4CanaFixedWaitingTimeStrategybeaPureESS?..................61
8.5AMixedESSSolutiontotheWaitingGame.......................65
9AMixedESSSolutiontotheWarofAttrition
67
9.1TheBasicsofaMixedESSintheWarofAttrition...................68
9.2SupportingStrategyProbabilitiesatEquilibrium....................69
9.3AnIntroductiontoIntegration..............................70
9.4TheMathematicsoftheMixedEquilibrium.......................78
9.5ADescriptionoftheMixedEquilibrium.........................82
9.6ProvingthatvarisEvolutionarilyStable........................85
10References
93
11Glossary
95
12Appendix:DiscussionandSelectedAnswers
100
12.1AnswersforChapter2...................................100
12.2AnswersforChapter4...................................102
12.3AnswersforChapter5...................................105
12.4AnswersforChapter6...................................106
12.5AnswersforChapter7...................................107
12.6AnswersforChapter9...................................109
Chapter1

ModelingBehavior:Gameand
OptimalityTheory

Synopsis:Thischapterpresentsageneraloverviewoftheuseofmodelsinevolutionary
biology.Thedi
ydiscussed.Theerencesbetweenadaptationalandneutralmodelsarebrie
bulkofthematerialdealswithanoverviewoftwotypesofadaptationalistmodels|optimality
andgametheory(especiallyoptimalitytheory)andendswithacomparisonbetweenthem.
Laterchapterswillpresentamoredetailedexplanationofgametheory.

1.1AdaptationversusNeutralModelsofEvolution

AdaptationalModels:FollowingthesuccessofDarwinandWallace'stheoryofnaturalselecti
on,
mostmodernbiologistsbelievethatmostoftheaspectsofthemorphology,behaviorandphysio
logy
ofanorganismrepresentadaptations.Thatis,theseaspectsofthephenotypeexistinapopula
tionbecauseintherecentpasttheyallowedtheirpossessorstoreproducemoresuccessfullyt
han
individualswithalternativetraits.

NeutralistModels:However,onemustkeepinmindthatadaptationisoftenonlyanassumption.

Inthe1930sSewellWright[1931]developedthemaintheoreticalunderpinningsofanalternat
ive
meansofaccountingforevolutionbasedongeneticdrift,aprocesswhichworkswellinsmall
populationsespeciallywhencompetingtraitsconferlittlerelativesurvivaladvantageove
reachother
(thesetraitsaresaidtobeadaptivelyneutral).

ExtendingWright'swork,othershaveshownthattheparticulartraitsfoundinapopulation
canbetheresultofhistoricalaccidents.Forinstance,ErnstMayr(certainlyanadaptationa
list.
pointedoutthatapopulation'scharacteristicscouldhavemuchtodowiththegeneticmakeupo
f
asmallnumberofprogenitorsorfounders[Mayr,1954].Morerecently,StephenJayGouldhas
writtenextensivelyabouttheroleofhistoryandaccident(contingency)indeterminingthep
resent
phenotypesofmembersofapopulationandforthatmattertheactualrangeoforganismsthatexi
st
atagiventime[Gould,1990].

ItisfairtosaythatmuchofthevalueoftheworkofGouldandothershasbeentoforcebiologists

toacknowledgethatallaspectsofthephenotypeneednotrepresentspeci
cadaptations|thatthe
phenotypeisinpartaccident,notanidealdesign,andinmanycasesanumberofcompetingversi
ons
ofaphenotypemightalldoequallywell,especiallygiventhenatureofenvironmentalchange.
Thus,
oneshouldgainevidencethataparticularphenotypicfeaturerepresentsanadaptation.Ones
hould
Chapter1.ModelingBehavior:GameandOptimalityTheory
2

notsimplymakeupcircular\just-so"storiesthatpurporttoshowadaptationbyassumingitan
d
thenspinningoutanexplanationbasedonthe(untested)assumptionofadaptation.

1.2MathematicalModelsandRigorousScience

Mathematicalmodelsareabstractionsthataninvestigatorhopeswill,tovaryingdegreesofp
recision,
havepredictivepower.Modelsrepresentthescientist'sbest(mostinformed)guessasto:

theidentityandfunctionofimportantvariables,
thewaysthesevariablesinteractwitheachother.
Mathematicalmodelsareusefulbecauseoftheveryfactthattheytakeascientist'sideasand
produceamorecomplexabstraction(sinceamodelconsistsofpartsandtheirinteraction).Co
mplex,
multi-elementmodelsoftenyieldnewinsightsandnovelpredictions.Mathematicalmodelsha
ve
theadvantagethattheyyieldquantitativepredictions.Quantitativepredictionsareoften
less
ambiguousthanothertypesofpredictions.Sincetestsofhypothesesareattemptstoshowthe
predictionsareincorrect(testsattempttofalsifythemodel)quantitativepredictionsare
usually
easiertotest|didthemodelbehaveexactlyaspredictedornot?Ifnothowdierentwasitfrom
prediction?Howcouldthemodelbemodi
edtomakeitmoreconsistentwiththeresultsandthen
re-tested.

Inabilitytofalsifythemodeldoesnotvalidateit.Inabilitytofalsifymeansnothingmoreth
an
that.Notshowingthatamodeliswrongmeansonlytentativeacceptance,notproofofitstruth.
Amodelthathasnotbeenfalsi
edisnothingmorethanausefulworkinghypothesis.Forexample,
atellingobservationwasmadebyDr.DavidNormanaboutrestorationmountsofdinosaurs(thes
e
mountsare,ofcourse,nothingmorethanhypotheses)whenhewrylyobserved,\We'vegotitrigh
t|
fornow."Muchoftheformalismoftestinganddescribingthescienti
cprocesscanbetracedtothe
workoftheEnglishphilosopherSirKarlPopper[1972].(ForareferencetoErnstMayr'sextend
ed
andfascinatingtreatmentofbiologicalmethodology,see[Mayr,1982]).

Mostcommonly,modelsaremodi
edastheresultof(i)experimentalfalsi
cationofoneor
moreoftheircomponentsor(ii)independentre
nementsinourunderstandingofthevariables
andinteractionsthatalreadymakeupamodelorthatshouldbeaddedtothemodel.Youhave
probablynoticedthatthesameprocessisnormallyfollowedthroughoutthescienti
cprocess;the
maindierenceifanyisthathypothesesintheformofmathematicalmodelsareoftenmoreconcret
e
andquantitativelypredictivethanareothertypesofhypotheses.However,keepinmindthatw
orking
withexperimentallysupportedquantitativemodelsisfraughtwiththesamedangersaswithle
ss
quantitativemodels|likeanyhypothesismodelsshouldalwaysbeviewedwithskepticismando
nly
trustedtotheextentthattheyhavebeenstronglytested.

Oneothernoteaboutmathematicalmodelsinevolutionarybiology:theycanspringfromeither
anadaptationalistoraneutralistviewpoint.Twoimportanttypesofmathematicaladaptatio
nbasedmodelsareoptimalityandgametheorymodels
.Thenexttwosectionscomparethese
twoapproaches.Sincebothareadaptationmodels,bothwilllookforbehavioralcharacterist
icsthat
maximizeanindividual'sreproductivesuccessorsomerelatedvariable.

1.3OptimalityModels

Oftenabehavioristisinterestedinpredictingthebestway(intermsofits
tnessconsequences.
foraparticularanimaltobehaveirrespectiveofwhatotherindividualsaredoing.Toillustr
ate,
Chapter1.ModelingBehavior:GameandOptimalityTheory

supposewearetryingtounderstandhowloudlyananimalshouldmakeanadvertisementcall(one
designedtoattractamate).Thus,wearelookingatageneralbehavior(producinganadvertise
ment
call)andwearetryingunderstandtheselectiveforcesthatdeterminethebestwaytoperforma
particularpartofthebehavior|inthiscaseitsloudness.Inthisparticularexample(andina
ll
optimalitymodels)westartfromtheassumptionthattheloudnessofothercallershasnothing
todo
withpredictingtheloudnessofagivenindividual.Sowemightimagineasituationwhereanani
mal
callswithoutothersnearby(aswouldbethecaseinmanyspeciesofcrickets).

Optimalitytheoryisadaptational:thusitspringsfromthetheoryofnaturalselectionwhich
predictsthatananimalshouldbehavesoastomaximizeits
tness.Allbehaviorscanbeviewed
ashavingboth
tnessbene
tsand
tnesscosts.Sinceoptimalitymodelsarequantitative,a
rst
stepwillbetoestablishtherelationshipsbetweenthevariableofinterest(thedecisionvar
iable.
andtheassociatedcostsandbene
ts.Bene
ts(B)andcosts(C)arekeptstrictlyseparate(just
asispracticedinbusinessbookkeeping)andthustwoseparaterelationships(bene
tversusthe
decisionvariableandcostversusthedecisionvariable)areimplicitlypartoftheprocessof
setting
upanoptimalitymodel.Noticethatsinceweassumethatthebehavior(decisionvariable)has
consequenceson
tness,thebehavioristheindependentvariableforeachoftheserelationshipswith
bene
torcostbeingthedependentvariables.Now,sincethe
tnessconsequenceofbehavioris

NetChangeinFitness=Bene
t..Cost,
(1.1.

thesolutiontoanoptimalitymodelisto
ndthepointwhereB..Cismaximized.

Inprinciplethisformulationiseasytounderstand.However,inpractice,itcanbemorecompl
ex.
Let'sreturntoourexampleofcallloudnesstoillustratetheprocessofconstructingandsolv
inga
simpleoptimalitymodel.

Inacousticcommunication,producingaloudcallisenergeticallyexpensive[Prestwich,199
4].
However,loudercallstendtoattractmoremates,(forexample,see[ForrestandGreen,1991])
.We
wanttoconstructanoptimalitymodelinordertotrytopredicthowloudlytocallwiththe\goal
"
ofmaximizinglifetime
tness.

Let'sstartwiththebene
tsofcallingmoreloudlyforacertainperiodoftime.Intheorythere
areanumberofpossiblerelationshipsbetweenloudnessandbene
t.Onewouldbelinear|get
ectionwould
louderandproportionatelymoremateswillcome(graphI,Figure1.1).Alittlere
suggestthatthiscannotgoonforever|atsomepointincreasingloudnessbringsinsomanymate
s
thatthefocalanimalcan'thandleallofthemandsothereisnofurtherincreasein
tnesswith
loudness(GraphII,below).Alternately,onemightassumethattherateatwhichmatingsincre
ase
dropobefore
nallyreachingamaximum(graphIII,below)|i.e.,becauseofotherthingsthe
callerhastoattendto,asthenumberofmatingopportunitiesincrease,thepercentagethatar
e
actuallyconsummatedbecomesless:

Thereareseveralthingstonoteabouttheseplots:

Eachrepresentsadistincthypothesisastotherelationshipbetweenbene
tandloudness;

Beyondtheirgeneralshapes,morespeci
cadjustmentscouldbemade.Forinstanceinan
environmentsuchasonethatisheavilyvegetated,soundwouldbeheavilyattenuatedandthe
slopesofallofthegraphswouldprobablydecrease.

Noticethatthe
tnessmeasureusedhere(numberofmatings)isratherstraightforwardand
probablyeasilyrelatedtorelativereproductivesuccess.Thatisnotalwaysthecaseaswewil
l
seebelow.

CostsCurves:Itislogicaltoassumethatloudcallingwillhaveconsequenceson
tnesssince
itcostsconsiderableamountsofenergyandthereforecouldweakenthecaller.Thisleadsdire
ctlyto
Chapter1.ModelingBehavior:GameandOptimalityTheory

Figure1.1:Threepossiblerelationsbetweenloudnessofcallandbene
t.

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Loudness

expressingcostasenergylosswhichcanbemeasureddirectlyandrelativelyeasily).Butabso
lute
energytermssuchascaloriesorjoulesmaynotbethemostrelevantwaytomeasuretheenergy
portionofthecostofcalling.Forinstance,ifthecostofgettinglouderisviewedrelativeto
energy
stores,lowcostcallsmighthavelittleimpactonreservesandtherefore,uptoapoint,increa
sing
loudnessmighthavelittleaccompanyingincreaseincost.However,whentheenergydemandsof
makingloudercallsincreasesbeyondacertainpoint,theymaysigni
cantlyaectananimalsenergy
reservesandthereforeforceittoeithercallforashortertimeperiod(perhapstherebylower
ingits
tness)oreatless(alsoloweringits
tness).Noticethatonceagain,theexactpositionandshapeof
thecurvewoulddependonmanyfactors|forinstancesizeoffoodreservesandtheeasewithwhic
h
theyarereplacedrelativetotheincrementalcostsofloudercalls.

Moreover,anotherimportantcostofcallingisincreasedchanceoffallingvictimtopredatio
n
(seeMikeRyan's[1985]workonbatspreyingoncallingfrogs).Whatunitsdoyouusetomeasure
thiscost?Thecostcouldbeexpressedasthechanceofbeingkilled,orinjuredorsomethingels
e,
orbetteryet,thenumberoffuturematingsorfutureospringlostasresultthechanceofinjuryo
f
deathassociatedwithacertainloudnessofcall.

Let'sassumethatwebelievetherearejusttwoimportantcoststocalling|energyandrisk.We
needtocombinethemintoonecostrelationship.Botharealreadyrelatedtothesamedecision
variable(loudness)butthesetwotypesofcostsareusuallyexpressedinunitsthataredierent
from
eachother(forexample,joulesforenergyandchanceofdeathforpredationrisk).

Ifwewanttocombineallcostsintoonecurveweneedtoputthemintoacommonunitof
measurement(acommoncurrency).Thiscurrencymustalsobethesameoneusedinthebene
ts
function.Thiscanbeverydicultbutletssaythatwehavefoundawaytoexpressbothenergy
andpredationriskaslostfuturematings.Andlet'ssaythatourbestunderstandingisthatwea
k
callsdonotattractpredatorsatallandconfernoadditionalfeedingdemands.However,event
ually
apointisreachedwherepredationstartstoincreaseandeventuallyalsothatsigni
cantincreases
infeedingmustoccur.Figure1.2showsagraphicalexampleofourcostshypothesis.

Nowlet'scompleteandsolveourmodel.Let'ssaythatwedecidethatbene
tsmodelIIIisthe
bestonetouse.Ifwenowexpressbene
tsandcostsofcallloudnessinacurrencyofmatings,then
wecansuperimposethebene
tandcostsplotonthesameaxisandthensolvefortheloudnessthat
givesthegreatestincreasein
tnessasmeasuredbythegreatestnumberofmatings(Figure1.3).

Noticethatthemodelpredictsthatthebestloudnesstocallisnottheloudest(whichiswhat
Chapter1.ModelingBehavior:GameandOptimalityTheory

Figure1.2:ApossiblerelationbetweenLoudnessofcallandCost(inlostmatings).

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Cost
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Loudness

Figure1.3:GraphofthedierencesbetweenBene
tsandCostsversusloudness.

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Loudness
Chapter1.ModelingBehavior:GameandOptimalityTheory
6

youwouldpredictifonlybene
tsofloudnesshadbeenconsidered).Noticealsothatinthiscase,
themodeldoesnotkeepcoststoanabsoluteminimum.
WecanusetheBandCcurves(above)tomakeanothergraphinFigure1.4,thistimeof
B..Cversusloudnesstoillustratetheoptimumanotherway:

Figure1.4:GraphofB..Cversusloudness.

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Lifetime
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Matings

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Loudness

Thistypeofdepictionclearlyshowsthatthegreatestlifetime
tnessinourhypotheticalsituation
isachievedbycallingwithsometypeofintermediateloudness.

Finally,realizethatthemodelisnothingmorethananintegratedsetofhypothesesthattoget
her
attempttopredictthebestloudnesstocall,irrespectiveofwhatothersaredoing.Aswesawab
ove,
thispredictionwillneedtobetested(forinstancebyattemptingtodeterminethelifetimema
ting
curveforanimalsofdierentloudness).Inpractice,thisessentialstepisoftenverydicult(th
ink
howharditwouldbetodeterminethecurvegivenaboveforsometypeofanimal)andmuchofthe
artandhardworkofbehavioralscienceliesinthesetestsofthemodels.

1.4ABriefIntroductiontoGameTheory

OK,wejustwentthroughalongintroductiontoanimportanttopicinbehavior,physiology,and
ecology:optimalitytheory.Butyouthoughtthischapterwasaboutgametheory.Itis.Thestu.
onoptimality(above)wasimportantbecauseyouneedtoknowsomethingaboutoptimalitytheor
y
tounderstandgametheoryandespeciallytounderstandwhichtechniquetousewhentryingto
understandanimalbehavior.

So,whatisgametheory?Inoptimalitytheoryitisassumedthatwecanpredictthebest
behaviorforaparticular(focal)animalirrespectiveofwhatothersaredoing.However,thef
requencywithwhichothersareperformingaparticularbehaviorisoftenhighlyrelevanttoth
e
tness
consequencesofactingacertainway.Thus,thecrucialaspectofgametheoryisthatselection
amongalternativebehaviorsdependstoalargedegreeonwhatothersaredoing.Abehaviorthat
worksverywellwhenrareinapopulation(forinstance,sometypeofdeception)maynotbenearl
y
asadvantageoustoitsactorswhenitbecomescommon.Thusexplorationsofgametheoryinvolve
studiesofaformoffrequency-dependentselection.

Here'sabriefexample.Let'sreturntocallingbehavior.Let'sassumethatfemalesare
attractedtocallingmalesandtraveltothemtomate.Butlet'salsoassumethatarrivingfemal
es
arenotinfallibleinspottingthemalethatactuallywasmakingthecalltowhichtheywereattr
acted.
Or,femalesmaybeinterceptedbymalesastheyapproachandsometimestheinterceptingmales
werenottheonesthatattractedthefemaleinthe
rstplace.

What'sthebestthingforamaletodo?Ifnooneiscalling,itisprobablybesttocalland
becomeconspicuoustofemalesinterestedinmating.However,iftherearemanycallers,itmig
htbe
Chapter1.ModelingBehavior:GameandOptimalityTheory
7

besttokeepquiet(avoidingenergyandpredationriskcosts)andtrytointerceptfemalesasth
ey
approachthecallingmale.Suchamaleistermedasatellite.

Asastrategysatellitecouldwellbeasormore
tthancalling,dependingonitsfrequencyrelative
tocallers.Thus,eventhoughasatellitemight(perday)havefeweropportunitiesformating(
lower
bene
t),thefactthathiscostsarelessmeansthathemighthaveasmanyifnotmorelifetime
opportunitiesformating.However,asshouldbeobvious,hisrelativesuccessdoesnotsimply
reduce
tocostsandbene
tsasinoptimalitytheory{instead,his
tnessdependsverymuchonthefrequency
ofcallers(versussatellites)inthepopulation.Iffewotherscall,satelliteisprobablyno
tagood
strategyandrelativelyspeaking,callingisanexcellentstrategy(ortakingittoanextreme
,ifno
oneiscalling,satellite(simplesilence)makesverylittlesensesincefemalesareattracte
dtocalling
males).Ontheotherhand,asmoreandmorecallersarepresent,satelliteworksbetterandat
somefrequenciesmight,overalifetimepayobetterthancalling.Thus,frequencydependence
distinguishesthisexamplefromsimpleoptimality.

Note:Don'tgettheideathatIamarguingherethatsatelliteisgenerallyamoreorless
t
strategythanthealternative,call.Dependingonconditions,atanymomentintimeallofthes
eare
possible.Laterwewillconsiderthemostinterestingoutcomeofevolutionarygames,anevolu
tionary
stablestrategy(ESS).WewillseethatonetypeofESS(mixedESS)predictsthatbothstrategie
s
wouldcoexistatconstantrelativeproportionstoeachotherandatthesefrequenciesthe
tnessof
individualsofeachstrategyareequal.

Summary

Noticethesimilaritiesanddierencesbetweenoptimalityandgametheory.Bothdealwiththe
bene
tsandcostsofabehavior.Insituationswhereoptimalitytheoryisapplicable,thatisalltha
t
isneeded.However,asintheexampleabove,therearemanycaseswherethe
tnessassociated
withaparticularbehaviordependsonwhatothersaredoing,thatis,
tnessisfrequency-dependent.
Thus,whatisbestwilldependonwhateveryoneelseinthepopulationisdoing(andthereforeon
thelikelihoodofcertaintypesofinteractionswithcertaintypesof
tnessconsequences).

Younowhaveabasicideaabouttheusesofmathematicalmodelinginstudiesofbehavioral
evolution.Youshouldalsobefamiliarwiththebasicsofgameandoptimalitymodels.Youmayno
w
continueontoamoredetailedintroductiontogametheoryespeciallyinregardstoanimportan
t
concept,theevolutionarystablestrategy,oryoumaycontinuereadingthischapterandlearn
abit
aboutthehistoryofgametheory.

1.5ABriefSketchoftheHistoryofGameTheory

Gametheoryisequallyusefulinstudiesoflearnedandinnatebehavior.Infact,whenoriginal
ly
developedbyvonNeumannandMorgenstern[1953]duringthemid20thcentury,it'sprimary
purposewastounderstandthemostrationalwayforhumanstomakedecisionsbetweenalternati
ve
coursesofaction,inparticularastheyappliedtoeconomics.

However,anytechniquethatwillallowustostudythepayosofalearnedbehaviorcanbeused
equallywelltostudyinnatebehavioralstrategies.JohnMaynardSmith[1982]pointsoutthat
the
ideaofrationalinterestineconomicsissimplyreplacedbytheconceptof
tness.Hetracestheuse
ofgametheoryinbiology
rsttotheworkofLewontin[1961]andlaterSlobodkinandRapoport
[1974].Theseworkersappliedgametheorytosituationsof\speciesorgroupsurvival"(Didyo
uever
playtheboardgame\Extinction").ThepresentmainstreamofgametheorybeganwithHamilton
[1967]andthenafewyearslaterwithMaynardSmithandPrice[1972]andthenMaynardSmith
[1974].Intheinterveningyears,gametheoryhasbeenadaptedbyanumberofbiologiststoexam
ine
Chapter1.ModelingBehavior:GameandOptimalityTheory

evolutionaryproblems|hardlyanissueofAnimalBehaviourpasseswhenonedoesnotseeanarti
cle
usinggametheory.

Perhapsthebestthoroughintroductiontotheuseofgametheoryinbiologyisbyoneofthe
pioneersinapplyingittobiology|JohnMaynardSmith[1982].Theinterestedandmathematica
lly
inclinedstudentisurgedtoreadhisclassicintroductiontothe
eld.Thisunitisbasedlargelyon
thematerialinthe
rstfewchaptersofhisbook.
Chapter2

GameTheoryandEvolutionary
StableStrategies

Synopsis:Thischapterintroducesthecentralconceptoftheapplicationofgametheoryto
evolutionarybiology|theEvolutionaryStableStrategy.Youwilllearnthebasicterminolog
y
ofandtechniquesforsolvingevolutionarygameswithtwostrategies.Aftercompletionofthi
s
chapter,youwillmoveontosolvingaclassictwostrategygame(HawksandDoves)andthen
tosimulationsandmorecomplexgames.

2.1Introduction:BehavioralStrategiesandGames

Whycallitgametheory.

Intheprevioussection,ModelingBehavior:GameandOptimalityTheory,welearnedthatcompe
titionwasanimportantfeatureofgametheory
.1Thus,theanalogybetweenhumanbehaviorand
gametheoryisofcompetitors(players)seekingtowinsomethingthroughsomesortofcompetit
ion
(contestorthegameitself).Notethatingametheory,asinhumangames,theoutcomeofacontes
t
toaparticularplayerisshapedbyboththeactionsofthefocalplayerandher/hisopponent.An
d
bothhumanandevolutionarygamescanhavedierentstructures.Forinstance,theoutcomecan
bedeterminedasthesumofaseriesofone-on-oneencountersbetweentheplayersoritcaninvol
ve
acontestwhereeachplayerisworkingmoreorlessagainsteveryoneelseatonce.Andclearly,i
n
bothofthesecases,theoutcomewilldependonthebehavior(s)oftheplayers.

SomeDe
nitionsandCaveats

OK,soweunderstandwhythesearecalledgames,althoughfromthepointofviewoftheanimals,
theyaredeadlyserious.Let'sseehowweformallyanalyzeagame|howwemaketheoretical
calculationsofrelative
tnessthatarebasedonbene
ts,costsandfrequenciesofvarioustypesof
outcomes.

1Aboutcompetition:Obviously,competitionisalsoinvolvedatsomelevelinoptimalitymode
ls.Animalsthat
\discover"thebestwaytoperformaparticularbehaviorinsituationswherethepayosarefrequ
ency-independent
arestillcompetingwithothers,albeithighlyindirectly.Perhapsagoodwaytothinkaboutco
mpetition,inthesense
itisusedingametheory,isthatthecompetitionismoredirect.Itinvolvessomethingthatwem
ightanalogizeto
humancontests,althoughthosecontestscouldbetheeitherone-on-oneaairsweusuallythinko
fascompetitionor
oneagainsteveryoneelseplayingthe
eld.
Chapter2.GameTheoryandEvolutionaryStableStrategies
10

ContestsandGames:Let'sgetafewconventionsoutoftheway.First,weareonlygoingto
lookatonegeneralmodelofcompetition|whatMaynardSmithtermedpairwisecompetition.In
pairwisecompetition,eachcontestinvolvestwoindividualsplayerscompetingwitheachoth
erat
atime.Justasimportantly,pairwisemodelsviewthe
tnessconsequencesofaparticularcontestas
summingovertime(asmorecontestsoccur).Thistypeofmodelisquiteusefulinanimalbehavio
r
sincetherearemanysituationswherewecanseetwoindividualsinteractingoversomeresourc
e.
Furthermore,wecanseethattheconsequencesoftheseinteractionsseemtosumindetermining
the
tnessoftheplayers.2
Strategies:Theparticularbehaviororsuiteofbehaviorsthataplayerusesistermedastrate
gy.
3Strategiescanbebehaviorsthatareonsomecontinuum(e.g.,howlongtowaitordisplay)or
theymayrepresentdistinctlydierentbehaviors(e.g.,display,
ght,or
ee).Sometimestheterms
purestrategyandmixedstrategyareused.(DonotconfusethesewiththetermsPureESSand
MixedESS|itcanbeeasytodo.)Apurestrategyisastrategythatisnotde
nedinterms
ofotherstrategiespresentingame.Examplesofpurestrategiesthatwewillconsiderlaterar
e
\Hawk"and\Dove"|theyrepresentverydierentwaysoftryingtoobtainresources|
ghtingand
displaying.Ontheotherhand,sometimesstrategiesaremixesofothers.Anexampleofamixed
strategyiswhenoneindividualplaysamixof\Hawk"and\Dove"withacertainprobability.As
wewillseelater,pureandmixedstrategiesarenotnecessarilyESSs|justkeepallofthisinth
e
backofyourmind,you'llberemindedaboutitlateron!

Onemorepointaboutplayers(contestants,whatever)andstrategists.Playersareindividua
ls
whouse(play)acertainpureormixedstrategy.Whenwelookatagame,onewayistoconsider
the
tnessconsequencesofcontestsonanindividualwhoplaysacertainstrategy.However,wewill
morecommonlylookatthegamefromapopulationviewpoint.Theplayersinagamebecomethe
strategiesthemselves(orthegenesthatencodethesestrategies).Thegamewillthenconside
rthe
overall
tnesseectsoneachstrategyafterallpossiblecontestsareplayedinproportiontotheir
likelihood.(Trustme,thiswillmakemoresenseinamoment).Noticethatthissortoftreatmen
t
owesmuchtopopulationgeneticswheregenes(alleles)arecommonlyviewedincompetitionwit
h
eachother.However,pleasenotethatthisinnowayimpliesanysortofgroupselection|allele
or
strategycompetitionmodelscanbeviewedassomethinganditscopies(regardlessofwhethero
r
notthesearegenesorlearnedbehaviors)competingwithsomeotherthing(anditscopies)wher
e
thewinnersmakethemostadditionalcopiesofthemselves.

AsexualModels

Finally,tomakethegamessimple,weassumethatreproductionisasexual,or,iflearned,theb
ehavior
issimplycopiedbytheospring.Thus,therearenocomplicationsrelatingtogeneticinteracti
ons
betweentheallelesproducingdierentstrategies,noneedtoworryaboutsmallpopulationeects
,
etc.MaynardSmithandothershaveproducedmanygameswheresexisafactor.Theinterested
readerisurgedtolookatChapter4inMaynardSmith[1982]foranintroductiontothesemodels.
Noticealsothatassumingasexualreproductionmakesiteasytoadaptthesemodelstotransmis
sion
bylearning(assumingthatnomodi
cationoccursintheprocess).

2Amorecommontypeofgameisprobablyplayingthe
eld[MaynardSmith,1982],butwewillnotconsiderthis
model.

3Notethatinevolutionarystudies,thewordstrategydoesnotimplyconsciouschoiceorplann
ingonthepartof
theactor.Weusethetermasashorthandtodescribewhatishappeningintermsofourownexperie
nce.Butweare
awarethatinmostcasestheanimalisbehavingbysomesortofinstinctual,heritablerulesfor
behavior.Thus,the
strategyisprobablynotplannedoutinthesensethathumanstrategyisplanned.
Chapter2.GameTheoryandEvolutionaryStableStrategies
11

2.2FormalAnalysisofaPairwiseGame

Part1:ContestsandPayos

Asstatedabove,whenwetalkaboutstrategiesinthecontextofpairwisecompetitioningameth
eory,
wewillbeinterestedintheoutcomesofmanycontests.Acontestoccurswhentwoindividuals
interactwithinthecontextofthegame.Thatis,theycompeteforsomesortofresourceusing
thebehavioralstrategiesunderconsiderationbythegametheorist(rememberthatthesegame
s
arearti
cialconstructstoallowustounderstandwhattheanimalsaredoing).Contestscanoccur
betweenindividualsthatusethesamebehavioralstrategy(e.g.,displayforalenghtoftimet
1versus
displayfortimet1)ortheymayoccurbetweenindividualswithdierentstrategies(e.g.,displ
ayfor
timet1versusdisplayfortimet2).

Let'smakethisabitmoreconcreteusingExample1aboutsatellitebehavior.Wehavetwo
dierentstrategies,callandsatellite.
4Thepotentialcontestsare:

CallerversusCaller,
SatelliteversusCaller,
SatelliteversusSatellite.
Whataretheevolutionarysigni
cancesofthesecontests?Putinthetermsofanevolutionary
game,wewouldliketoknowthe
tness(orsomestand-infor
tness)consequencesontheactorsof
eachtypeofcontest.Weusuallyrefertothese
tnessconsequencesaspayos.

Ingamesinvolvingnon-continuousbehavioralstrategiesweusuallystartwiththeconstruct
ion
ofapayomatrix.ThematrixinTable2.1listsallthepossiblecontestsandtheirassociated
payos.

Table2.1:Thegeneralpayomatrixforatwostrategygame.

Opponent
FocalStrageyCallSatellite
Call
E(C;C)E(C;S.
SatelliteE(S;C)E(S;S.

Thereisaformalismtoitsconstruction:

Typicallytheleftcolumn(labeledFocalStrategy)liststhestrategiesinthegame.

Thesestrategiesarealsolistedastheheadsofthecenterandleftcolumns(bothlabeled
Opponent).

Nownoticethattheremainingcells,identi
edbyrowandcolumn,representeverypossible
typeofcontestinthisparticulargame.I'llrefertothemcollectivelyasthecontestorpayo.
cells.Ifyou'velearnedanybasicgenetics,thispayomatrixshouldremindyouofaPunnet
square(sinceitisthesamething).

4Thesestrategiescouldbeviewedasbeingeitheronacontinuumofcallingorastotallydierent
behaviors,the
distinctionislargelysemanticinthiscase.
Chapter2.GameTheoryandEvolutionaryStableStrategies

Therowheadedbyaparticularstrategythenlistsallpossiblecontesttypesforthatstrategy
.
{Thus,theinmatrixabove,thethirdrowshowsthetwopossiblecontesttypesthatCall
strategistswouldexperience.
{ThebottomrowdoesthesameforSatellitestrategists.
{Thus,aCallercancompetewithanotherCaller(centercell,labeledE(C;C))oraCaller
cancompetewithaSatellite(rightmostcell,labeledE(C;S)).Asimilararrangementis
usedforaSatellite(bottomrow).

Thepayosarelistedineachofthecontestcells.Whennotgivenexplicitly,theyareoften
abbreviated.Forinstance,theabbreviationforthepayotothestrategyCallinacontest
withanotherCallerisE(C;C)where:

{Estandsforpayoorexpectation,and

{the
rststrategywithintheparenthesesreferstothestrategywhosepayoisbeing
calculatedand

{thesecondstrategyintheparenthesesreferstotheotherstrategyinthecontest.
Thus,E(C;C)isthepayotoaCallstrategist5whenengagedinacontestwithanotherCall
strategist;E(S;C)isthepayotoaSatellitestrategistwheninacontestwithacaller.

Noticethatthepayomatrixisabitmorecomplicatedthanwasthesimplelistofcontests.In
thecallerversussatellitegame,therewerethreegeneraltypesofcontestsbutthepayomatrix
lists
fourpayos.Whyisthis?Thereasonisthatthepayomatrixliststheconsequencestoastrategy
foreachpossibletypeofcontest.Itthusbecomesobviousthatthestrategy\call"experience
sa
certaintypeofpayowhenevertwocallerscompete,E(C;C),andalsowheneveracallercompetes
againstasatellite,E(C;S).Likewise,thestrategysatelliteexperiencesonetypeofpayowhe
n
pittedagainstcall,E(S;C),andanotherwhenpittedagainstanothersatellite,E(S;S)).

Part2:CalculatingPayostoaStrategyinaParticularContest

Now,letsseehowtocalculateeachpayo.6

Firstwewillneedacompletedescriptionofthestrategy|howdoesitbehaveinregardsto
otherknownstrategies.
Next,wewillneedtoconvertthisdescriptionintopayos.Todothis,wewillneedtofactor
in:

{thevalueoftheresource,
{chancesofwinningaresource
{lessanycostsinvolvedinwinning
{thecostsoflosingand
nally

{thechanceofaloss.
5Aboutpayostostrategiesandstrategists:Onecanjustaswellthinkaboutthisintermsofpayos
tothe
strategy(orthestrategy'sgene)|butnogroupselectionisimplied.

6Morefussynotesaboutpayos:Realizethatpayosaregivenasexactamountsinthisgamewheninfa
cttheyare
averages;likewisechancesofvictoryareaverages|inarealsituationsomeanimalswouldcle
arlybemorecompetent
incompetitionsthanothers.Butwewanttokeepthissimple.
Chapter2.GameTheoryandEvolutionaryStableStrategies

Thus,allpayocalculationswillhavethegeneralform:

Payo(toStrategy1,whenversusStrategy2)=(Bene
tfromwin)..(Costofloss)(2.1.

Sincethesearecontests,procuringabene
torpayingacostdependsonanumberoffactors.Thus,
wemustfactorinthechanceofwinningaresourceofsomevalueandthechanceofpayingacostin
losing.Soweexpand(2.1)to

Payo(toStrategy1,whenversusStrategy2)=(chanceofwin)(resourcevalue.
+(chanceofloss)(costofloss)(2.2.

Even(2.2)isprobablynotsucientsince(2.2)statesthatwinninghasnocosts.However,inmany
conteststhereisacostpaidbythewinner.Goodexamplesmightbeenergeticortimecostsof
displays;thesecanbeseenasloweringthevalueofwinning.Thus,(2.2)mightbeexpandedto:

Payo(to1,whenvs.2)=chanceofwin(resourcevalue..costofwin.
+(chanceofloss)(costofloss.
(2.3.

Wenowhaveagoodgeneralizedequation.Noticethatanyofthetermsin(2.3)canbemade
todropoutsimplybysettingthemtozero.Thus,inaparticulartypeofcontestifthestrategy
underconsiderationincursnocosttowinningandthereisnochanceoflosing(thechanceofwin
ning
becomes1.0),thentheentireequationforpayoreducestothevalueoftheresource.

Herearefewimportantconsiderationsaboutcalculationsofpayos

Ingamesimulationsitiscommontoassignbene
tsandcostsusingsometypeofrelative
butarbitraryscaleofvalue.Thus,thebene
tofobtainingtheresourcemightbeassigneda
valueof1.0andthecostofatypicaldisplaymightbeassignedavalueof..0:1.Whatthese
assignmentsreallysayisthattheresourceisworthabout10timesthecostofatypicaldisplay
.

Selectionoftheserelativevaluesisextremelyimportant|asyouwilldemonstratetoyoursel
f
whenyouusethesimulations,dierentvaluesofbene
tsandcostscanresultinverydierent
outcomes.

Asintheoptimalityexample,thesearbitraryunitsmustbeacommoncurrency.
Aboutsigns:
{Bene
tsareusuallygivenpositivevalues.

{Costsmaybeassignedeitherpositiveornegativevaluesdependingontheformofthe
payoequation.
Thus,whencostsaregivennegativevaluestheyareaddedtothebene
ts(aswas
showninthegeneralequationforcalculationofpayosatthestartofthissection).
Whenpositivevaluesareusedforcosts,theyareusuallysubtractedfrombene
ts.

Obviouslytheeectisthesame;theonlythingthatmattersisthatthegametheoristis
consistent.

Finally,sincegamemodelsareabstractions,itisquitecommontoignorecertaincostsor
bene
tsorsometimesrollthemtogether.Forinstance,inthegameswewillrunassimulations,
youwillseethatsomecostsareignoredorareexpressedimplicitly.
Chapter2.GameTheoryandEvolutionaryStableStrategies

Part3:CalculationoftheFitnessofEachStrategy

Neitherthepayosforacontest,e.g.,thevalueofE(C;C),northesimplesumofalltypesof
payostoastrategy,e.g.,thepayorowfortheC-strategy:E(C;C)+E(C;S),fromthematrix
inTable2.1willgivethe
tnessofastrategy.Recallthatingames,
tnessalsodependsonthe
frequencyofotherbehaviors.

Amoment'sre
ectionwillwillrevealthatthefrequencyofeachtypeofinteractionisavitalpart
ofany
tnesscalculation|ifsatellitesareveryrarethenthe
tnessconsequencesofinteractingwith
asatellitearerelativelysmallascomparedtoiftheyweremorecommon.So,if
tnessisdenoted
byW,thentheoverall
tnessconsequencestoaparticularstrategistinaparticulartypeofcontest,
forexampleacallerversuscallercontest,aregivenas:

ChangeinW(Strategy1)=E(toStrategy1,versusStrategy2)frequency(encounter)(2.4.

Sinceourexamplegameonlyconsiderstwostrategies(callandsatellite),ifwedenotethefre
quencyofcallerascandthethefrequencyofsatelliteass
,then

FrequencyofCaller=c:
(2.5.

Ifthereareonlytwofrequencies,theymustsumto1,so

FrequencyofSatellite=s=1..c:
(2.6.

Thus,inthisgamethe
tnessconsequencestocall,W(C),arethesumofthepayosforeach
typeofinteractiontimesthefrequencyofthatinteraction:FitnessofCallStrategy=Fitnes
s
ChangeDuetoInteractionwithotherCallers+FitnessChangeDuetoInteractionwithSatellit
es,
orsymbolicallyas

W(C)=E(C;C)c+E(C;S)s:
(2.7.

Moreusefully,ifwesubstitute1..cfors:

W(C)=E(C;C)c+E(C;S)(1..c):
(2.8.

Asimilarcalculationcanbemadeforthe
tnessofsatellite:

W(S)=E(S;C)c+E(S;S)(1..c):
(2.9.

Youhavenowlearnedthebasicformalismstosettinguponeespeciallyusefultypeofgametheor
y
simulation.Inthenextsection,wewilllookatoneofthemostimportantoutcomesinevolution
ary
gametheory|theEvolutionaryStableStrategy(ESS).

Problem

1.Noticethattheselastthreeequations(2.7){(2.9)quantify
tnessassomesortofnumber.One
mightthinkthatthelarger(morepositive)thenumbergivenbyeitherequation,themore
successfulthestrategyinthisevolutionarycompetition.Butisthatreallycorrect?Here's
your
question:Isitcorrecttotalkaboutthe
tnessofeitherstrategyinisolation,thatis,ifonlyone
strategyispresentinthepopulation,dothe\
tnessvalues"calculatedbytheequationsabove
haveanyrealmeaning.
Chapter2.GameTheoryandEvolutionaryStableStrategies
15

2.3Stasis:EvolutionaryStableStrategies

Oneofthemostimportantconsequencesofgametheoryisthatitcanbeusedtopredictsituation
s
where:

onebehaviorismore
tthanallknownalternatives

oralternately,aspeci
cmixofbehaviorswherenoonebehaviorismore
tthananyother.

Inbothcases,theresultisevolutionarystasiswithrespecttothebehaviorsbeingconsidere
d;there
isnochangeinrelativefrequencyofstrategiesovertime.Thesesituationsaretermedevolut
ionary
stablestrategiesorESSs.TherearetwotypesofESS:

1.PureESS:whereonestrategytotallyout-competesallothers.Thatmeansthatregardless
ofitsfrequency,itisalwaysmore
tthananyknownalternative.Astrategythatisapure
ESSisimmunetoinvasionbyotherknownstrategies.Thus,anyalternativethatappearsby
mutationorimmigrationwillnotbeabletoincreaseandwilleventuallygoextinct.

2.MixedESS:wheretwostrategiespermanentlycoexist.Foragivensetofpayos,therewill
beonesetoffrequencieswherethismixisstable.AmixedESScanbeachievedifindividuals
either

playonestrategyallofthetimeinapopulationwherethetwostrategiesareatthe
equilibrialfrequencies.Forexample,60%oftheindividualsalwayscalland40%always
actassatellites.Or
allindividualsplayamixedstrategywhereeachofthebehaviorsinthemixisperformed
attheequilibrialfrequency.Forexample,allindividualscall60%ofthetimeandactas
satellites40%ofthetime.
IneithermixedESScase,attheESS,allindividualshavethesame
tnessregardlessoftheir
strategy.7Atanyotherfrequenciestherewillbe
tnessdierences.Forexample,ifanew
individualentersthepopulation(e.g.,anewcallerorsomeonewhocalls70%andsatellites
30%ofthetime),the
tnessofallindividualswiththisbehaviorarelowerthanthealternative.

Fortheexampleofanextracaller,allofthecallerswouldhavelowered
tnessrelativeto
satellite.

Fortheexampleofthemixedstrategy(70%caller:30%satellite)invader,itwillhavea
lower
tnessthaneveryanimalthatadoptsthemixedESSmixedstrategyof60:40.

Asnotedabove,atamixedESS,thepurestrategiesexistatthefrequencieswheretheir
tnesses
areequal.Itisveryimportanttorealizethatequal
tnessesdonotimplyequalfrequencyinthe
population!Aswewillsoonsee,thefrequencieswilldependonthepayomatrix.Wewillseea
numberofexamplesofmixedESSslaterandyouwillbeabletosimulatethem|butyouwillseldom
ndonewherethetwofrequenciesareequal.
7About
tness:Rememberthatallofthesediscussionsonlyrelatetothetraitsunderconsiderationin
thegame.
Notalltheindividualshavethesameoverall
tnessbutthesedierencesarenotcorrelatedwiththestrategybeing
usedandsoatthisequilibriumthereisnodierencein
tnessofeitherofthetwostrategiesandthereforenochange
ineitheroftheirfrequencies.
Chapter2.GameTheoryandEvolutionaryStableStrategies
16

OtherPointsAboutESSs

Intwostrategygames,therewillalwaysbeapureormixedESS:

ItshouldbeobviouswhysomesortofESSmustoccurwhenthereareonlytwostrategies:The
pureESScaseisselfexplanatory.ThepureESSstrategywilltotallydominatethepopulation
exceptforoccasionalmigrants,mutants,orindividualstemporarilytryingtogainany
tness
theycan.
However,ifthereisnopureESSwhymusttherebeamixedESS?Recallthat
tnessis
frequency-dependent.Thus,aslongasthetwostrategiesdonothaveidenticalpayos8then
atsomefrequenciesonewillbemore
twhileatotherstheotherwillbemore
t.(Again,if
oneisalwaysmore
t,itisapureESS).Sinceoneisnotalwaysmore
tthantheotherthen
theremustbeapointwherethetwohaveequal
tnesses.Thefrequencywherethathappens
isthemixedESS.

Ingameswherethreeormorestrategiesplay,itispossibletohaveasituationwherethereis
noESS.We'lldiscussthisatamoreadvancedpartofthisunit.
ACoupleofNotes

Iftheprecedingparagraphsconfuseyou,couragemesamis,we'lllookattheseconceptsinmore
detailbelowandthenyou'llhaveachancetoinvestigatethemwiththeaidofmodels.

NotethatwhenwesayanESS(whethermixedorpure)cannotbeinvadedwemeanthatitcannot
beinvadedbyanyotherknownstrategy.ESSsarealwaysde
nedagainstotherknownalternative
behavioralstrategies.AnESSisalwayspotentiallyvulnerabletoanynewstrategythatmight
come
along.Wewillseeexamplesofthiswhenwegofromatwotoathreestrategygame.

Problems

2.WhatisthemeaningofE(B;A).

3.AssumethattwoalternativestrategiesmakeupamixedESSatfrequenciesof0.8forstrategy
Aand0.2forstrategyB.Furthermore,assumethatallindividualspracticebothAandB.
Describeeachindividual'sbehavior.

4.ExplainthedierencesbetweenapurestrategyandapureESS.Betweenamixedstrategy
andamixedESS.

2.4PureESSsinTwoStrategyGames

Intwostrategygamesitisarelativelysimplemattertodetermineifoneofthestrategiesisap
ure
ESS,providedcertainveryreasonableassumptionsaremet.Inthissection,wewillreviewthe
procedureformakingthisdeterminationandthelogicbehindthisprocedure.

RecallthatapureESSisastrategythatisunbeatablebyotherknownstrategies.Thismeans
that:

8Iguessitshouldbementionedthatifbothstrategiessomehowalwayshaveexactlythesame
tness,thena
situationhasbeenproducedwherechangeispossiblethroughmutationormigration(orinnon-
geneticmodels,
learning).NoticethatthiswouldnotbeanESSsincethechangeswouldnotcause
tnessestobeunequaland
therebyfavorashiftbacktothepreviousfrequencies.
Chapter2.GameTheoryandEvolutionaryStableStrategies

apureESSisimmunetoinvasionbyanyotherknownstrategy.
and,astrategythatisapureESSisalsocapableofinvadinganddisplacingotherknown
strategies.
Ifwecanshowthateitherofthestatementsaboveistrue,thenwehaveshownthatthestrategy
isapureESS(eitheroneis
ne,theyareessentiallyequivalentasfarasthemathematicsofthe
gameisconcerned).

Wecanusethepayomatrixandasimplifyingassumptiontomakethedetermination.Let's
getawayfromthecaller/satellitemodelandinsteadde
netwoabstractstrategies:AandB.The
purposeofthisswitchissimplytogetyoumoreusedtomanipulatingandusingthepayomatrix.

Asalways,thematrixliststherelativepayostoeachstrategyforeachtypeofencounter.In
thisexamplewewillassignavaluetoeachpayo.SeeTable2.2.

Table2.2:AssigningpayostotheAversusBstrategies.

Opponent'sStrategy
FocalStrageyA
B
AE(A;A)=0E(A;B)=1
BE(B;A)=..0:5E(B;B)=0:5

Let'sassumethat:

ThepopulationinitiallyconsistsentirelyofindividualswhousestrategyA.
AverysmallnumberofstrategyBinvadersappear.Inthesimplestcase,thiscouldbea
singleindividualinvadingalargegroupofAstrategists.Thisinvasioncouldbetheresultof
mutation,learning,orimmigration.
Meetingsthatleadtocontestsbetweendierentindividualsoccuratrandom.Thus,thereis
notendencyforindividualstocollectbystrategyincertainplacesandincreasecertaintype
s
ofinteractionsoverwhatwouldhappenbychanceencounter.

Whattypesofinteractionsoccurandhowfrequentarethey?Themostcommoncontestswill
involveAstrategists.Whyisthisthecase?TheansweristhatnearlyeveryoneisanAstrategis
t
ictswithanalternativestrategyaredirectlyrelatedtothefrequencyof
andthatmeetingsandcon
thatstrategy.Thus:

AversusAcon
ictswillbethemostcommonforAstrategistsandwillthereforelargely
determinethe
tnessofAstrategists.FromthepointofviewofA,theseoccuratthefrequency
ofA.

BversusAcon
ictsarethemostcommonforBstrategistsandthereforewilllargelydetermine
the
tnessofBstrategists.InthemostextremecasewherethereisasingleBinvader,B
versusAwillbetheonlytypeofencounterthatmatterswithrespecttoits
tness.

AnyBversusAcon
ict!Putanotherway,sucha
ictcanalsobeviewedasanAversusBcon
ictinvolvespayostobothstrategies,E(B;A)tostrategyBandE(A;B)tostrategyA.Notice
con
however,thatfromthepointofviewofA,interactionswithBareextremelyrareascomparedto
Chapter2.GameTheoryandEvolutionaryStableStrategies

thosewithA.Thus,wewillassumethatwecanignorethe
tnesscontributionofAversusB
interactionstotheoverall
tnessofstrategyA.Youwillhaveachancetolookatthisassumption
inmoredetaillateron.

ANoteofWarning

Rememberthatweareattemptingtocalculatestrategy
tnesses.Thus,weareinterestedinthe
frequencyofcertaintypesofinteractionsfromthepointofviewofthestrategy.Sinceweare
consideringpairwisecontests,thefrequenciesofeachcontestfromthepointofviewofoneco
ntestant
(strategist)willbeequaltothefrequencyoftheopponentinthecontest.

Sometimesstudentswhoarefamiliarwithbasicprobabilityandpopulationbiologyassumetha
t
thefrequencyofaparticularpayoequalsaterminabinomialexpansionofthestrategyfrequenc
ies.
Forexample,ifaisthefrequencyofstrategyAandbisthefrequencyofstrategyB,then(a+b)2
isexpandedtopredictthat:

AversusAcon
ictswouldoccuratthefrequencya2
,

ictsat2ab,and
AversusBcon

BversusBcon
ictsatb2
.

Thissortofformulationistrueifonewantstoestimatetherateofoccurrenceoftheseinterac
tionsin
thewholepopulation.Butitisnotcorrectwhenweareonlyinterestedinthefrequencyofinter
actions
fromthepointofviewofaparticularstrategy!

Now,ifweconsultthepayomatrix,wecanseehowthisinvasionturnsout.Inourexample:

ThepayotostrategyAwhenitisanAversusAcontestis0.(Ourconventionisthat
thismeansthattheinteractionhasno
tnessconsequences|itneitherincreasesnordecrease
reproduction).

WhataboutstrategyB?All(ifasingleinvader)ofthecontestsitfacesareBversusA.From
thematrix,weseethatthepayotostrategyBincontestsversusstrategyAis..0:5.That
is,Blosessome
tnessasaresultofthisinteraction.

ItshouldnowberatherobviousthatBcannotinvadeA.
Fromthesituationwejustconsidered,wecanconstructageneralruletodeterminewhetheror
notatwostrategygamecontainsapureESS:IfE(A;A)>E(B;A)(themostcommonencounter
foreachstrategy),thenAisstableversusB(itisapureESSversusB).

Youmaybewonderingwhatwouldhappenifthe
tnessconsequencesofthemostcommontypes
ofinteractionsareequal,i.e.,E(A;A)=E(B;A).Doesthatmeanthatneitherisstableagainst
the
other?Notnecessarily.Inthisonecase,thereisanadditionaltestthatmustbeperformedbef
ore
concludingwhetherornotthereisapureESS.

IfthereismorethanoneBinvader,theremayalsosomerareinteractions,withpayoE(B;B).
Also,inthisparticularsituation,thepayoE(A;B)startstomatter,eventhoughitisstillext
remely
rare.

Whynowbutnotbefore?Inthepreviousexample,theAversusBinteractionwasveryrare
incomparisonwiththecommonAversusAcontests.Thus,anyeectsontheoverall
tnessofA
duetointeractionswithBweresosmallastoprobablynotmatter.However,inthecaseweare
ictconfersnorelative
nowconsideringE(A;A)=E(B;A).Thus,thecommonAversusAcon
advantageordisadvantage.(ThesamelogicappliestothemostcommoncontestBexperiences,B
Chapter2.GameTheoryandEvolutionaryStableStrategies

versusA).SotheremaininginteractionswilldecidewhetherornotthereisapureESS.Thus,if
E(A;B)>E(B;B),thenAmuststillhaveanadvantageoverBandthereforeitwillbestable!

Toreviewthis,considerthefollowingscenario.ApopulationofAstrategistsisinvadedbya
smallnumberofBstrategists.Inthemostcommontypesofcontestsforeachstrategythepayos
E(A;A)andE(B;A)areequal.Thus,neitherstrategyiscompetitivelyaidedorhinderedbythes
e
contests.However,intherarecontests,AisdoingbetterthanBsinceE(A;B)>E(B;B)andso
Awilleventuallyout-competeB.

SummaryofRulesforFindingaPureESSinaTwoStrategyGame

Assumption:Onestrategyisveryrarecomparedtotheother.Inthisexample,letAbethe
commonstrategy.ItisimmunefrominvasionbyBif

Rule1eitherE(A;A)>E(B;A.

Rule2orE(A;A)=E(B;A)andE(A;B)>E(B;B).

AnothergoodwaytothinkaboutRule1isthinkofitasthe\equilibriumproperty,"aterm
usedbyRiechertandHammerstein[1983]toindicatethatthebeststrategytofacestrategyAis

alsostrategyA.TheequivalencebetweenthislaststatementandRule1shouldbequiteevident
sinceRule1saysthatthepayotoAversusAisgreaterthanwhatBwouldgetagainstA.

RiechertandHammersteinrefertoRule2asthe\stabilityproperty."Thereasoninghereis
thatifBdoesjustaswellAagainstasdoesAitself,thenAwillonlybestableifitdoesbetter
againstBthanBdoesagainstitself.

YoumaybeuneasyaboutRule1.MathematicallyyoucanvaguelyimaginecaseswhereE(A;A.
isgreaterthanE(B;A)yetAisnotstableagainstB!Thesesituationsrequiremorethanone
Bstrategyinvadersothatallthepayosmightmatter.Sincemorethanoneinvaderisnotan
unreasonablescenario,youbecomesuspiciousthatgametheoristsareeitherintellectually
shallow
oraretryingtosweepthingsunderarug.

Problems

5.Aquickreviewonnotation:WhatdoesE(B;A)mean.

6.Thefollowingsimpleproblemsillustratetheassumptionswemadeaboutthefrequencyofvar
ious
contestsinourpopulationmainlycomposedofAstrategists.Assumethatthefrequencyof
strategyAis0.9999.Calculate:

a)thefrequencyofstrategyB.
b)thefrequencyofAversusAinteractions.
c)thefrequencyofBversusBinteractions.
d)thefrequencyofAversusBinteractionswherethepayoistoA,i.e.,E(A;B).
e)thefrequencyofBversusAinteractionswherethepayoistoB,i.e.,E(B;A).

f)WilltheproportionofthetotalnumberofpayostoAwhenversusBbeanydierentthan
theproportionofthetotalnumberofpayostoBwhenversusA.

7.WritetheexpressionfordeterminingwhetherornotstrategyBisapureESSagainstA.
Chapter2.GameTheoryandEvolutionaryStableStrategies
20

8.UsingtheexpressionforBversusAthatyoujustwroteandthematrixinTable2.2,explain
whetherornotBisstableagainstinvasionbyA.

9.AssumethatE(A;A)=E(B;A).WhatifE(B;B)>E(A;B)?DoesthatmeanthatBisnow
stableagainstA.

10.IfastrategyisnotapureESS,doesthatmeanthattheopposingstrategyisapureESS.

11.WhenusingMaynardSmith'sshortcutmethodto
ndapureESS,whatisthehypothesized
situationwithrespecttothefrequenciesofeachstrategy.

12.Inathreeormorestrategygame,willfailureto
ndanypureESSstrategymeanthatthe
remainingstrategiesformamixedESS.

2.5MixedESSsforaTwoStrategyGame

WhatifneitherstrategyisapureESS?Ifthereareonlytwostrategies,thentheremustbeamixe
d
ESS.ThereasonwhyamixedESSisrequiredinthissituationiseasytounderstand.Imaginea
situationwhereneitherAnorBisapureESS:

Apopulationthatiscomposedentirelyofmembersofonestrategy,Aforexample,isinvaded
bytheother.ThefactthatBcaninvadeAmeansthatatleastatlowfrequencies,Bismore
tthanA.
Butlet'sturnthisaround.SinceBisalsonotapureESS,apopulationofBstrategistscould
beinvadedbyA.Thus,atlowfrequenciesAismore
tthanB!
Whatdoesthismean?Itmeansthataseitherstrategyincreasesfrombeingrare,itsrelative
tnessmustbegintodecreaseandeventually,bythetimeitisverycommon,itmustbeless
thanthe
tnessoftheotherstrategy.
Thus,theremustbesomeintermediatepointwherebothhavethesame
tness.Thefrequencies
atthispointarethemixedESS.

Noticethatifeitherstrategyincreasesaboveitsequilibriumfrequency,itbecomesrelativ
ely
less
t.Thus,wehaveatrueequilibriumsincethestrategyfrequencywilltendtoreturnto
itsoriginalvalue!
So,let'sseehowto
ndamixedESSmathematically.Wewillusethissolutioninthesimulation
asanaidinvisualizingmixedandpureESSs.Lettherebetwostrategies,AandBatrespective
frequenciesaandb.Wehavealreadyseentheexpressionsforcalculatingthe
tnessofeachstrategy.
ForA:

W(A)=E(A;A)a+E(A;B)b
(2.10.

andforB

W(B)=E(B;A)a+E(B;B)b
(2.11.
Chapter2.GameTheoryandEvolutionaryStableStrategies

Now,toachieveandequilibrium,the
tnessesmustbeequal.Ifnot,onestrategywillbeincreasing
relativetotheother.Thus:

W(A)=W(B.
E(A;A)a+E(A;B)b=E(B;A)a+E(B;B)b:
(2.12.

Substituting1..aforbandrearranging,wecansolveforthefrequencyofstrategyAwhereits
tnessisequaltoB,i.e.,forthefrequencyofAatthemixedESS:
aE(B;B)..E(A;B.

.
:
(2.13.

1..aE(A;A)..E(B;A.

Wecannowuseato
ndbatthemixedESS.
Thissolutioncanbevisualizedgraphically.Ifwewriteboth(2.10)and(2.11)intheform
y=mx+bwhereyis
tnessandxisthefrequencyofstrategyA,then:

W(A)=[E(A;A)..E(A;B)]a+E(A;B.
(2.14.

and

W(B)=[E(B;A)..E(B;B)]a+E(B;B):
(2.15.

Ifweplotsthese,wegettwostraightlinesthatintersectatsomefrequencyofstrategyAthat
dependsonthevaluesinthepayomatrix.Figure2.1showsanexamplefromtheHawksand
Dovesgamethatwewilllookatnext.Pleasenotethat
tnessisbeingexpressedin\payounits"
andbeawarethattheslopes,intercepts,etc.,willbedierentwithdierentpayomatrices.

Figure2.1:Gametheorymodeling:Hawks(dashedline)andDoves(solidline).

50
50
.
.............
.
.............
.
.............
.
.............
.
.............
.
.............
.
.............
.
.............
.
..........
....
.
.............
.
................................................................................
................................................................................
................................................................................
................................................................................
.............................................
.....
..............
..
.............
.

.
................................................................................
................................................................................
......................................
........
.
.............
.
.............
Payo.
..
.............. .
................................................................................
.................
.
0
.
.
.............
.
.............
.
.............
ESS0.5
.
.............
.
.............
.
.............1.0
.
.............
.
.............
.
.........
Frequencyof
.
.............
.
.............
HawkStrategy

..50

Noticethattheintersectionisastableevolutionaryequilibriumsincetheadditionofindiv
iduals
ofeitherstrategylowerstheaverage
tnessofallmembersofthatstrategy.

Thus,anincreaseinHawksresultsinmovingtotherightalongtheHawk(dashed)linetoa
lower
tnessthanthatofDove;thatis,awayfromequilibrium.

Likewise,additionofDoves(movementtotheleftalongtheDove(solid)linesincetheir
frequencyis1..hlowerstheir
tnessrelativetoHawk.

Inbothcases,thelowered
tnessresultsinreductionoftheirnumbersbacktotheequilibrium
point.
Chapter2.GameTheoryandEvolutionaryStableStrategies
22

2.6CanDierentStrategiesCo-ExistWithoutanESS.

Atthispointyoumightexpectthatwhenevertwostrategiespersistinapopulationthattheywo
uld
alwaysformamixedESS.Thisisnottrue.TheyareonlyamixedESSiftheir
tnessesareequal.
Youmightaskhowbothstrategiescouldpossiblypersistinthesamepopulationiftheir
tnesses
werenotequal.Whatfollowsisanon-exhaustivelistofrealisticalternativestoamixedESS.

Disequilibrium:Inadisequilibrium,onestrategyismore
tbuttherehasnotyetbeensucient
timetoreachequilibrium.Disequilibriumcouldbemaintainedbyanumberofprocesses,forin
stance
thearrivalofmigrantswhoexhibittheless
tstrategy(althoughpresumablyitisquite
tinthe
populationtheyoriginatedfrom).Geneticdriftcouldalsocontributetodisequilibriumasc
ould
geneticlinkages.

ChangingEnvironments:Anextensionoftheideaabove.Here,anequilibriumisnotreached
becausetheenvironmentchangesandfavorsoneandthentheotherstrategy.Therearemany
examplesinevolutionofthissortofcyclicalvariationandthedisequilibriumthatresultsa
ndithas
beenahotareaofresearchinpopulationgenetics.

CopingorMakingtheBestoftheSituation:Let'ssaythatacertainstrategyisapureESS.
Andlet'ssaythatanimalshaveachoiceaboutwhetherornottoexhibitthisstrategy.Onemight
thinkthatananimalwouldalwaysexhibitthepureESSstrategy.Butthisisnotnecessarilyso.
Whatifthestrategyisverycostlyandnotlikelytosucceedforanindividualwhoisinacertain
condition?Itmaybethatthealternativebehaviors(tothepureESS)availabletoanindividua
lare
ofdemonstrablylower
tness(e.g.,theyallyieldlowerchancesofmatinginthepresent).

Let'sillustratethisbyreturningtothecaller-satellitesituation.Sofarwehavetreatedt
hesetwo
strategiesasiftheywerepartofamixedESS.Infact,somestudiesofanuranshavesupportedth
e
notionthatthe
tnessesofcallerandsatelliteareequal.However,whatiftheywerenotequal?If
westartwiththelogicalassumptionthatcallismore
tthansatellite,thenwewouldexpecttosee
satellitedisappear.Orwouldwe?Wehavealreadyconsideredthatcallingiscostly.Whatif,o
na
particularnight,amaledidnothavesucientenergytocallandhaveagoodchanceofattracting
amate?Aslongastherewassomechancethatsatellitesgainmates,eveniffewerthancallers,i
t
wouldpaya\weak"maletosatelliteorengageinsomeotherlessenergeticallyexpensivewayto
obtainamate(e.g.,searching).Satellitebehaviorwouldnotdisappearbutwouldremainatal
ow
frequencyandwouldtendtobepracticedbyindividualswhentheirenergyreserveswerelow.Wh
en
theywereinbettercondition,theycouldcall.Buteventhoughtbothstrategiespersist,they
are
notamixedESS.

Anotherwaytolookatthissortofcopingbehaviorisasanoptimizationproblem|willengaging
inlower
tnessbehavioratcertaintimeincreasemylifetime
tnessbygivingmesomegainsnow
andperhapsgreatergainslater.

Testingtheseideas

Inallofthesecases,adecisionaboutwhetherornotthebehaviorswereanESSwouldrequiredat
a
onrelative
tnessandtheirpersistenceoveranumberofgenerations.Theimportantthingtorealize
isthatsimply
ndingalternativestrategiesinapopulationdoesnotprovethatamixedESSexists
anymorethan
ndingasinglestrategyprovesapureESSexists.

OK,youarenowfamiliarwiththebasicsofpairwisegames.Wewillnowmoveontoaseriesof
simplegamesthatwillhelpyoutounderstandhowgameswork,theirimplicationsforbehavior,
and
perhapsalsohelpyouseehowtoapplyabstractgamestothebehaviorsofrealorganisms.
Chapter3

HawksandDoves

Synopsis:Hereyouwillhaveachancetoapplywhatyouhavelearnedaboutgamesandtheir
solutiontoaclassictwostrategygame|HawksandDoves.Youwillbeintroducedtotothese
strategieswhichhaveutilityinunderstandinghow
ghtinganddisplaystrategiescouldco-exist
inapopulation.Afterthisintroduction,youwillbeguidedthroughtheconstructionofapayo
.
matrixwhichyouwillusetodeterminewhetherornotHawkorDovearepureESSs.Youwill
alsobeintroducedtoagraphicaldepictionofevolutionarygames.Thischaptermarkstheend
ofyour\basictraining"ingametheoryandisthegatewaytousingthesimulationsthatwill
beprovided.

3.1Introduction

Inthelastsection,welearnedthebasicsofsettingupandsolvingatwostrategygame.However
,
wedidnotactuallyconstructandsolveagame.Inthissection,wewillconstructaclassicbutv
ery
simplegameknownasHawksandDoves.Thesetwosimpli
edbehavioralstrategiesemployvery
dierentmeanstoobtainresources|
ghtinginHawksanddisplayinDove.
1Thesedierencesin
behaviorhavemarkedconsequencesonthechanceofwinningandofpayingcertaintypesofcosts
.
Thisleadstoverydierentpayos.

UsetheHawkandDoveexampleinthischaptertosolidifyyourunderstandingofbasicgame
theory.Yourfundamentalgoalshouldbetofeelthoroughlycomfortablewiththebasicconcept
sof
evolutionarygametheoryandwithsolutionstotwostrategygames.Inaddition,asyoustudyth
e
material:

ThinkhardaboutspecieswherestrategieslikeDove,Hawk,oramixmightoccur.Trytomove
beyondthegametoapplicationtorealanimals.
Oneofthemostimportantthingsyoushoulddoistothinkabouthowtherelativevaluesof
thefactorsthatdeterminethepayosultimatelyaecttheequilibrialpointoftheHawksand
Dovesgame.Animportantpartofyouruseoftheaccompanyingsimulationswillbetestout
yourideasabouthowsettingdierentbene
tsandcostswillchangetheequilibrialpointofa
twostrategygame.
1NotethattheHawkandDoverefertotwodierentbehavioralstrategiesadoptedbymembersofthe
samespecies.
Wearenotpittingtwodierentspeciesagainsteachother.
Chapter3.HawksandDoves
24

ThinkaboutthelimitationsofsimplemodelslikeHawksandDoves,yetatthesametimebe
suretothinkabouttheinsightsithasgivenyou.
3.2EvolutionaryStableStrategies:AnExample

Aswithanygamemodel,ourcentralquestioniswhetherornotDoveandHawkcancoexistandif
so,atwhatfrequency.Hereisadescriptionofthetwoalternativebehaviors:
Hawk:veryaggressive,always
ghtsforsomeresource.

FightsbetweenHawksarebrutalaairswiththeloserbeingtheonewho
rstsustainsinjury.
Thewinnertakessolepossessionoftheresource.

AlthoughHawksthatloseacontestareinjured,themathematicsofthegamerequiresthat
theynotdieandinfactarefullymendedbeforetheirnextexpectedcontest.2
Forsimplicity,wewillassumethatallHawksareequalin
ghtingability,thatis,eachHawk
ict.AnotherwayofsayingthisisthatHawk
hasa50%chanceofwinningaHawk{Hawkcon
versusHawkcontestsaresymmetrical.

Dove:never
ghtsforaresource|itdisplaysinanycon
ictandifitisattackeditimmediately
withdrawsbeforeitgetsinjured.

Thus,inanycon
ictsituation,DovewillalwayslosetheresourcetoaHawk,butitnever
getshurt(neversustainsadecreasein
tness)whenconfrontingaHawkandthereforethe
interactionsareneutralwithrespecttotheDove's
tness.3

AcorollarytothisruleisthatDovesdonotdisplayforverylongagainstHawks.Afterstarting
theirdisplays,theyimmediatelyrecognizethattheiropponentisaHawkandtheywithdraw
withoutpayingameaningfuldisplaycost.
Ontheotherhand,ifaDovemeetsaDovetherewillbeaperiodofdisplayingwithsome
cost(time,energyfordisplay)butnoinjury.WeassumethatallDovesareequallygoodat
displayingandandtheyadaptastrategyofwaitingforarandomtimeperiod(seeChapter
5,WarsofAttrition)thereforewhentwoDovesfaceo,eachhasa50%chanceofwinning.
NoticethatbothDoveswillpayessentiallythesamedisplaycostinanycontest.Thewinneris
theindividualwillingtopaymore.However,notethatthewinnerquitsdisplayingessentiall
y
atthesametimeastheloserwithdraws(seeWarsofAttrition).
2Whycan'tHawksdieorgetpermanentlyknockedoutofaction?Whymusttheybemiraculouslyres
toredto
health?Thereasonisverysimple.Ifthiswerenotthecase,theninanypopulationcontainingm
orethanoneHawk,
HawkversusHawkcontestswouldcausethefrequencyofHawktodecrease.ThemoreHawks,themor
eHawkversusHawk
contestsandthefasterfreq(Hawk)willdecrease!Noticethattheequationswelearnedearlie
rfor
ndingthe
tnessof
thestrategyallimpliedaconstantfrequencyofthestrategy.Thus,thebadthingsthathappen
inHawkversusHawk
contestsshouldbeseenaschanging(inthiscaselowering)thegeneral
tnessofHawkindividualsinthepopulation
withoutchangingtheirfrequencies.

3Thereareacoupleofthingstonoticehere.First,noDovesgetkilled.Toreiteratethemateri
alaboutfreqof
Hawkandinjury,noticethatifinjuredHawksdiddropout,thefreq.ofDovewouldincrease.Als
onoticethedierence
inthepayo(accordingtothedescriptionsofHandDthatyouhavejustreadorinthesameexamplep
ayomatrix
thatweconsideredwithHawk)|negativepayostendtomeanlowered
tnessasaresultofthecontestbutnotdeath
andpayosof0(thepayotoDoveversusHawkinthisexample)meannoeecton
tness|theDovegoesonasbefore.
Chapter3.HawksandDoves
25

Noticethatwehaveassumedthereareasymmetrieswithinastrategy|allHawksareequally
goodat
ghtingandallDovesareequallygoodatdisplays.Ananimalthatwinsonecontestisjustas
likelytowinortolosethenext.Thus,inanycontestbetweenmembersofthesamestrategy,eith
er
contestanthasanequalchanceofwinning|thereisnocorrelationwithpastsuccess,conditio
n,
whatever.Thisisclearlynotaveryreasonableassumption,butwe'rejuststartingoutsolet'
skeep
thingssimple.

Therearetwootherimportantassumptions.

Assumethattheattackinganimal(theonethateitherstarts
rsttophysicallyattackorto
display)hasnoknowledgeofthestrategythatitsopponentwillplay.

Assumethattheseinteractionsincreaseordecreasetheanimal's
tnessfromsomebaseline
tness.Inotherwords,theseinteractionssimplymodifyananimal's
tnessupordown|
winningacontestdoescreate
tness.Thisassumptionisassociatedwithourconventionthat
injuriesanddisplaycostswillbeassignednegativescores|losinganimalsdonothavenegati
ve
tnesses.4
3.3Preliminary(Qualitative)Exploration

Let'sstartbymakinganqualitativeanalysisofthegame.Thenwe'llusegametheorytomake
amuchmorequantitativeprediction(aswasdiscussedintheintroductorymaterialdealingwi
th
games).Let'sstartwiththefollowingquestion:Areeitherofthetwostrategiesbythemselve
s
impervioustoinvasion?Thatis,doeseitherrepresentapureESS.

TomostpeopleitimmediatelyappearsthatDoveisnotapureESS.Imagineapopulation
entirelyofDoves.Itisprobablyaveryniceplacetoliveandeveryoneisdoingreasonablywell
withoutinjurieswhenitcomestocon
ictsoverresources|theworstthingthathappenstoyouis
thatyouwastetimeandenergydisplaying.ButthatisOKbecauseontheaverageyouwin50%of
theencountersandthereforeontheaverageyouwillcomeoutaheadprovidedthedisplaycostsa
re
notlargecomparedtotheresourcevalue.

Now,imaginewhathappensifaHawkappearsbymutationorimmigration.TheHawkwilldo
extremelywellrelativetoanyDove|winningeveryencounterand,initiallyatleast,sueringn
o
injuries.Thus,itsfrequencywillincreaseattheexpenseofDove.Thus,DoveisnotapureESS.
If
DoveisnotanESS,whataboutHawk.

Let'sdotheanalysisagain,thistimestartingwithapopulationmadeentirelyofHawks.This
wouldbeanastyplace,anasphaltjungle,thatyouwouldn'tliketoliveinwithlotsofinjuriou
s
ghts.Althoughthese
ghtsdon'tkillyou,theytendtolowereveryone's
tness.Yet,justlikewith
theDovepopulation,noHawkisdoingbetterthananyotherandtheresourcesaregettingdivide
d
equally.

CouldaDovepossiblyinvadethisroughplace?Itmightnotseemsosincetheyalwayslose
ghts
withHawks.Yetthinkaboutit:

Dovesdon'tgethurt.Thebestwaytothinkaboutthisisthattheydonotpayhighcostsin
tnessforlosingacontest.WhiletheyneverbeataHawktheydon'tgethurtbecausethey
eethemomenttheyrealizetheyareinacontestwithaHawk.Theresultisthat,unlike
Hawk-Hawkcontests,aDove's
tnessisnotloweredbyacon
ictwithaHawk(i.e.,itisthesame
4Theimportantideahereisthattheanimalmustbeabletoreproduceevenifitlosesalloftheco
ntestsitengages
in.Ifnot,youmightaswellcounttheanimalasdeadwiththesameconsequencesasoutlinedinth
ediscussionof
injuries.Again,theimportantconsequenceofthegameiscontestsmayalterthe
tnessofindividualsbutnotkill(or
essentiallykill)theindividual.
Chapter3.HawksandDoves
26

asifnocontestatallhadoccurred|rememberthatweassumeacertainbaselineof
tness
independentoftheoutcomeofcontests|contestsonlyincreaseordecreasethisamount).

Dovesarewinners50%ofthetimeagainstotherDovesandtheyprobablyloselittleinsuch
contests.

Thus,ifamutantappearsintheformofaDoveoronewandersinfromelsewhere,itwilldoquite
wellrelativetoHawkandincreaseinfrequency.Thus,HawkisalsonotapureESS.

Noticethatinalloftheargumentsabove,wemadeimplicitassumptionsabouttherelative
valuesoftheresourceandthecostsofinjuryanddisplaythatareconsistentwiththebehavior
al
descriptions.Youprobablyrealizethatifwechangedsomeoftheseassumptionsofrelativeva
lue,
thegamemightturnoutdierently|perhapsHawkorDovecouldbecomeanESS.Moreover,evenif
westicktothequalitativevaluesandtoourconclusionthatthereisnopureESS,thetechnique
we
havejustusedwillnotallowustopredictthefrequenciesofDoveandHawkatthemixedESS.As
wasstatedearlier,thebestmodelsmakequantitativepredictionssincetheseareoftenmoste
asily
tested.

Thus,inthenextsectionwewillusetherulesandtechniqueswepreviouslylearnedtoquantita
tivelyanalyzetheHawksandDovesgame.

3.4FormalAnalysisoftheHawk-DoveESS

The
rststepofouranalysisistoset-upapayomatrix.Recallthatthematrixliststhepayosto
bothstrategiesinallpossiblecontests(seeTable3.1).

Table3.1:TheHawkversusDovepayomatrix.

Opponent
FocalStrageyHawk
Dove
Hawk
E(H;H)E(H;D.
Dove
E(D;H)E(D;D.

Weneedtomakeexplicithowwearriveateachpayo.Recallthatthegeneralformofan
equationusedtocalculatepayosis

Payo(toStrategy1,whenvs.Strategy2)=(chanceofwin)(resourcevalue)(3.1.
+(chanceofloss)(costofloss.

Wewillusethedescriptionsofthestrategiesgivenpreviouslytowritetheequationsforeach
payo.
But
rst,let'sassignsomebene
tsandcosts(wecoulddothislater,butlet'sdoitnowsothatwe
cancalculateeachpayoassoonaswewriteitsequation).

TherationaleforthevaluesinTable3.2isasfollows.For\gainresource,"itisself-explana
tory;
for\loseresource,"nothingisgained.For\injurytoself,"thecostofaninjuryislarge(ris
ky)ifit
isassumedthattherearelikelytobechancesinthefuturetogaintheresourceagain|injuryno
w
tendstoprecludegaininthefuture.Thismightappearquitedierenthoweverifthereisonlyone
chance|doesinjurymatter(ascomparedtowinning)ifyouareforcedintoacontesttohaveany
chanceatalltogainaresource?For\costofdisplay,"displaysgenerallyhavecostsalthough
how
Chapter3.HawksandDoves
27

Table3.2:ThepayovaluesforHawkandDove.

Action
Bene
torCost(arbitratyunits.

GainResource
+50
LoseResource
0
InjurytoSelf
..100
CosttoDisplaySelf
..10

hightheyarevaries|clearlytheyhavevariablecostsintermsofenergyandtimeandtheymay
alsoincreaseriskofbeingpreyedupon.Allofthesetypeofmeasurements,intheoryatleast,c
an
betranslatedinto
tnessterms.

Note:Alloftheseseparatepayosareinunitsof
tness(whatevertheyare!).Youwillsee
shortlythatthevaluesthatareassignedtoeachpayoiscrucialtooutcomeofthegame|thus
accurateestimatesarevitalinusefulnessofanyESSgameinunderstandingabehavior.

CalculationofthePayotoHawkwhenversusHawk

Relevantvariables(from(3.1)).
chanceofwinning(50%|i.e.thecontestsaresymmetrical.
resourcevalue(seeTable3.2.
chanceoflosing(50%.
costsoflosing|inthiscase,thecostisaninjurycost(seeTable3.2.

noticethatnocostsarepaidinwinning
Thus:
E(H;H)=(0:550)+0:5(..100)=25..50=..25:
(3.2.
Note:Thecostsoflosingareaddedinourmodelsincewegavethecostsanegativesigntoemphasi
ze
thattheyloweredthe
tnessoftheloser.

CalculationofthePayotoHawkwhenversusDove

chanceofwinning100%|i.e.,thecontestsareasymmetrical
resourcevalue(seeTable3.2.
nocoststotheHawksince(a)theyneverloseand(b)sincetheDoveimmediatelyretreatsonce

itrecognizestheHawk

Thus:
E(H;D)=1:050..0=+50:
(3.3.
Chapter3.HawksandDoves
28

CalculationofthePayotoDovewhenversusHawk
chanceofwinning0%|i.e.,thecontestsareasymmetricalandDovesalwayslose

nocoststotheDovesinceitimmediatelyretreatsonceitrecognizestheHawk
E(D;H)=050+1:00=0:
(3.4.

CalculationofthePayotoDovewhenversusDove

chanceofwinning(50%|DoveversusDovecontestsaresymmetrical

resourcevalue(seeTable3.2.

displaycostpaidinwinning|bothanimalswilldisplayessentiallythesameamountoftime.
Theonethatwinsistheonethatiswillingtodisplayforalongerperiodoftimethisparticular
time(seeWarsofAttrition).
chanceoflosing(50%.

costsoflosing|aswiththewinner,theloseralsopaysadisplaycostanditisthesameas
whatthewinnerpays(seeTable3.2.
noinjurycost|noviolenceplease,we'reDovesforheaven'ssake!
E(D;D)=0:5(50..10)..0:5(..10)=+15:
(3.5.

SoforthisparticularversionoftheHawkversusDovegame(de
nedbythesepayos),thepay-o.
matrixis:

Table3.3:AparticularHawkversusDovepayomatrix.

Opponent
FocalStrageyHawkDove
Hawk
..25+50
Dove
0+15

Problems

1.Inthelistofcostandbene
tsinTable3.2,itisassumedthatinjurycostsarelargecomparedto
thepayoforgainingtheresource.Giveasituationwherethisrelativeweighingmightaccurate
ly
ecttheforcesactingonananimal.
re

2.DoesitseemreasonablethatHawkspaynocostinwinning?Also,doesitseemreasonablethat
theloseronlypaysaninjurycost?Thinkaboutwhatanimalsdoandaboutsimpli
cationsof
models.
Chapter3.HawksandDoves
29

3.UsingthematrixinTable3.3,seeiftheHawkandDovegameabovemeetsthecriteriafora
pureESS.Hint:ReviewtherulesforapureESSandthenarbitrarilyde
neHasAandtestto
seeifHisapureESSwithpayoslistedabove(Dothisforbothstrategies|useHandthenD
asstrategyA.Shouldyougetthesameresultseachtime?.

3.5TheFitnessofEachStrategyandMixedESSs

Ifyoudidthelastproblemabove,youwillrealizethatneitherHawknorDovearepureESSsgiven
thepayoscalculatedfromtheequationsandvaluesforbene
tsandcostspresentedabove.(When
youusethesimulation,youwillseethatcertainbene
tsandcostscanbeusedtomakeeitherof
thestrategiespureESSs,althoughthesemightseemtoinvolveunreasonableassumptions).

Itisgoodtokeepinmindthefactthattherulesyouusedtodeterminethatneitherstrategy
wasapureESSrequiresomereasonableassumptions.IfwehavenopureESS,weknowthatin
atwostrategygametherewillbeamixedESSwhichisde
nedasthefrequencyofthestrategies
wherebothhaveequal
tness.Recallthatthe
tnessofastrategyisthesumofthepayostimes
thefrequencyoftheiroccurrence

W(strategy).
E(contest)frequency(contest):
(3.6.
contest
Thus,ifweassumethat
frequency(Hawk)=h;
(3.7.
then

frequency(Dove)=1..h:
(3.8.
Thus,the
tnessofHawk,W(H),is
W(H)=hE(H;H)+(1..h)E(H;D.
(3.9.
andthe
tnessofDove,W(D)is
W(D)=hE(D;H)+(1..h)E(D;D):
(3.10.
Noticethateachoftheequationsforstrategy
tnessyieldsastraightlinewhensolvedforaseriesof
frequencies.
NowsinceinamixedESSbothstrategiesmusthavethesame
tness,wecandeterminethe
equilibrialmixbysettingthe
tnessesofthetwostrategiesequaltoeachother,W(H)=W(D).
Forourgame,
hE(H;H)+(1..h)E(H;D)=hE(D;H)+(1..h)E(D;D)(3.11.
IfwenowsolveforthefrequencyofHawkatthisequilibrium,weobtain

hE(D;D)..E(H;D.

.
:
(3.12.

1..hE(H;H)..E(D;H.

Wecanunderstandthesolutionmoreclearlyifwegraph(3.8)and(3.9),asinFigure3.1where
thesolidlineisforDoveandthedashedlineisHawk.TheintersectionoftheHawkandDoveplots
representsthefrequencyofonestrategy(inthiscaseHawk)wherethe
tnessesofbothstrategiesare
equalintermsofpayounits.
Chapter3.HawksandDoves
30

Figure3.1:Gametheorymodeling:Hawks(dashedline)andDoves(solidline).

Payo.

50
50
.
.............
.
.............
.
.............
.
.............
.
.............
.
.............
.
.............
.
.............
.
..........
....
.
.............
.
................................................................................
................................................................................
......................................
.....
.
................................................................................
................................................................................
................................................................................
................................................................................
.............................................
....
...
.
........
.............
.
.
....
....
...
...
...
...
...
...
..
...
...
...
...
...
...
...
...
...
..
.
.............
.
.............
.
.............
Payo.
.
.............. .
................................................................................
.................
0
.
.
.............
.
.............
.
.............
ESS0.5
.
.............
.
.............
.
.............1.0
.
.............
.
.............
Frequencyof
.
.............
.
.............
.
.........
HawkStrategy

..50

RemarksAbouttheGraphicalResultstheHawkversusDoveGame

Figure3.1pointsupanumberofinterestingthings.

Notethatattheequilibrialpoint,theadditionofindividualsofeitherstrategylowersthe
relative
tnessofallmembersofthatstrategy.
{Thus,anincreaseinHawksresultsinmovingtotherightalongtheHawklinetoalower
tnessthanthatofDove|awayfromequilibrium.
{Likewise,additionofDoves(movementtotheleftalongtheDovelinesincetheirfrequency
is1..hlowerstheir
tnessrelativetoHawk.

{Inbothcases,thelowered
tnesswilleventuallyresultinareductionoftheirnumbers
andareturntotheequilibriumfrequency.

AlsonotethattheadditionofanyHawkoveralllowerseveryone'sabsolute
tness!|bothcurves
havenegativeslopes.

And,thegraphprovidesanotherwaytoseethatneitherHawknorDovearepureESSs:
{Hawkdoesverypoorlywheninhighfrequencycomparedtowhenitisrare.Thus,itis
easilyinvadedbywhatmightseemtobethemostimprobableofinvaders,thepaci
c
Dove.
{Ontheotherhand,DoveisnotstablesinceHawksdoextremelywellwhenenteringthe
population.
{Thus,althoughitisobviousthatanyHawkswouldalwaysbe\badforthespecies,"
theycannotbekeptoutoncetheyappear(somuchforgroupselectionbeingacommon
phenomenon).Forthisparticularsetofpayos,DoveisnotanESSanymorethanHawk
is.

Atthispointyouknowhowtosetupandsolveasimplegameandyouhaveabasicfamiliarity
withtheHawksandDovesgame.So,youarenowreadytoexploretheHawksandDovesgame
indetailusingPrestwich'ssimulationthatwillallowyoutoalterpayosbychangingbene
tsand
costs.Thesimulationwillprovideyouwithavisualrepresentationofthesolution,usingthe
same
techniquesyouhavejustlearned(exceptthecomputerwillnowdothecomputationalworkforyo
u).
Chapter3.HawksandDoves
31

And,you'llgettoseesomethingnew|you'llbeabletosetthefrequenciesofthetwostrategies
and
thenseehowapopulationwithagivenpayomatrixwillevolveovertime.

Problem

4.CalculatethemixedESSfrequenciesofHawkandDoveusingthepayomatrixinTable3.3.
Chapter4

TheHawksandDovesSimulation

Synopsis:ThischapterdescribeshowtousetheJavasimulationoftheHawksandDoves

game.Thisgamewasexplainedindetailpreviously;youwillnotgetmuchoutofthissimu

lationunlessyoualreadythoroughlyunderstandtheHawksandDovesgame.

Computersimulationsareusefulinthattheyallowyou:

toexplorealargenumberofsituationsinashorttimetherebyprovidingameanstoquickly
testyourunderstandingofthesystembeingmodeledand
visualizehowthesystemacts.
However,itcannotbeemphasizedtoostronglythatsimplyplayingwiththesimulationwithout
un-
derstandingwhatitdoesandwithoutrelatingtheinputsandoutputstosomebiologicallymean
ingful
situationislargely,ifnotentirely,afruitlessexercise.

4.1WhattheSimulationDoes

Thematerialbelowwillgiveyouanoverviewofthesimulation.Pleasetakethetimetoreaditso
thatyouhavesomebasicfamiliaritywithwhatitdoesandhowitworks.

WhenyoulaunchthesimulationapanelwillappearlikethatinFigure4.1.Atthebottomof
thepanelaretwobuttons:

Thebluebuttonwillbringupawindowthatgivesyougeneralinformationaboutthesimulation|
copyright,acknowledgments,etc.Closethiswindowusingthenormalcloseboxafteryouread
it.
Theredbuttonwilltakeyoutoanotherscreentosetupthegame.
Note:IfyouareusingtheApplet(web-based)version,this
rstscreenmustremainopenedthe
entiretimeyouusethesimulation.Closingitwillclosethesimulation.

TheSet-UpWindow(seeFigure4.2)willallowyoutoalterthepayosbychangingthe
bene
tsandcoststotheHawkandDovestrategies.Youwillnotbeabletochangethewaysthe
payostothestrategiesarecalculated,youcanonlychangetheparametersusedtomakethese
calculations.

Here'sabriefoutlineoftheset-upscreen:
Chapter4.TheHawksandDovesSimulation
33

Figure4.1:Whenyoulaunchthesimulationapanelwillappearlikethisone.

Thetoppanelgivesgeneralinformationaboutusingthesetupscreen|itismoredetailedthat
whatispresentedhere(sinceonlyanoverviewispresentedhere).
Themiddlepanelhasthreetext
eldsandabutton(seeFigure4.2).Whenyou
rstrun
thesimulationatanysession,thetext
eldswillcontaindefaultvalues(shown),whichare
thesamenumbersweusedwhenweconsideredtheHawksandDovesgame.However,youcan
changeanyofthesevaluesbysimplydraggingthemouseacrossthemandthentypinginthe
newvalueoryoucanpositiontheinsertionpointanduseyourdeletekey.

Note1:Imustemphasizethatthisistheonlywaythatyouwillbeabletoalterthepayos.
Note2:Also,pleasenotethatsimplytypinginanewvaluewillnotalterthepayomatrix.
Youmustpresstheredandyellowbuttonlabeled\CalculatePayoMatrix"torecalculate
withthenewvalues.Thisalsoappliestousingthedefaultvalues.

Thebottompanel(seeFigure4.2)hasanumberofimportantfeatures.

{The
rsttworowsarethegamepayomatrix.
Thebluebuttonswillbringupwindowsthatexplainhowagivenpayoiscalculated.
Adjacenttoeachbluebuttonistheassociatedpayo.Whenyou
rstopenthispage,
thepayoswillnotbegiven;youneedtopresstheredbutton(\CalculatePayo.
Matrix")inthemiddlepaneltogetadisplay.
{Thelastrowofthebottompanelhasthreebuttons(seeFigure4.2).

Ontheleftisagold\Reset"button|pressingitrestoresthebene
tsandcoststo
theiroriginalvalues.

Thebuttonlabeled\PlotFitnessGraph"makesaplotofthe
tnessofHandD
versusthefrequencyofHusingthecurrentpayovalues.Figure4.3showstheplot
forthedefaultvalues(yousawthisgraphwhenwe
rstlearnedaboutmixedESSs).
Chapter4.TheHawksandDovesSimulation
34

Figure4.2:TheSet-UpWindowconsistsofthreepanelsseparatedbyhorizontallines.

Thereareacoupleofthingstonoticeabouttheplot.First,
tnessonthey-axisis
expressedrelatively.Thus,italwayshasavaluebetweenzeroand1.0.
Secondly,beawarethattherearesomeslightroundingerrors.Forinstance,the
tnessoftheHawklinedoesnotstartexactlyat1.0onthey-axisnordoesitend
exactlyonthex-axisatafrequencyof1.0asitshould.Buttheerrorsarenotlarge.
Thirdly,belowthegraph,atextprint-outwilltellyouwhetherornotthereisapure
ormixedESSandifmixed,whattheequilibrialfrequencywillbe.Itusestherules
giveninprevioussectionstodeterminewhetherthereisapureormixedESSandif
amixedone,itusesthetechniquewelearnedearlierto
ndthevalueofthatmix.
Thelastyellowbutton,labeled\Evolve"willtakeyoutoanotherset-uppage.This
pageisusedtosetupasimulationthatshowsthechangeinstrategyfrequenciesand
relative
tnessovertimegiventhecurrentpayomatrixandaninitialfrequencyof
Hawk,whichyouset.SeeFigure4.4.

Noticethatthiswindowhasthreepanels:

Generalinstructionscanbeobtainedfromthebluebuttononthetoppanel.
Thebottompanelisforreference.Itcontainsadisplayofthepresentpayomatrixand
predictedstrategyequilibrialfrequencies.Notethatyouwillnotbeabletochangeanyof
thesevalues|theonlywaytoalterthemistogobacktothepreviouswindowandchangethe
valuesofthebene
tandcost.

Themiddlepaneliswheretheactionis!Hereyoucan:
Chapter4.TheHawksandDovesSimulation
35

Figure4.3:TheFitnessGraphwindowfortheHawksandDovesGame.
Chapter4.TheHawksandDovesSimulation
36

Figure4.4:TheSet-upforEvolutionSimulationwindow.
Chapter4.TheHawksandDovesSimulation
37

{Usethetop(yellow)text
eldtosettheinitialfrequencyoftheHawkstrategyandthereby
alsosetthefreq(Dove)sinceitequals1..freq(Hawk).Youshouldenteraninitialvalue
forthefrequencyoftheHawkstrategybytypinginavalueintheyellowtext
eldnextto
thetextabout\InitialFreq(Hawk)."Sincemutationandmigrationdonotoccur,besure
topickavaluegreaterthan0andlessthan1otherwiseyouareguaranteedstasis.

{Usethemiddlegoldpulldownmenutosetthenumberofgenerations(ithasadefault
valueof25generations).

{Whentheseareset,usetheredbuttontorunthesimulation.Whenyoupress\Run
Simulation"youwillgetagraphliketheoneinFigure4.5.

Figure4.5:TheChangesinFrequencyandFitnesswindow.

Noticethattherearetwographswithtwoplotsoneach:oneeachforthefrequencyofeach
strategyandonefortherelative
tnessforeachstrategy.Belowthegraphs,thereisatextmessage
thatwilltellyouthenumberofgenerationsrequiredtoreachanewequilibrium,ifatall.Plea
se
notethat,biologically-speaking,equilibriumwouldprobablyoccuratadierent,earliernum
berof
generations.TheprogramdoesnotassignequilibriumuntilthefrequencyofHremainsconstan
tto
38placesfortwosuccessivegenerations!

Ifyouwanttoreviewtheconceptsof
tnessandfrequencyandespeciallyifyouwantseean
exampleofhowtheevolvegraphiscalculated,seetheAppendixattheendofthischapter.
Chapter4.TheHawksandDovesSimulation
38

4.2DierencesBetweentheAppletandApplication

Thereareafewdierencesbetweenthestand-aloneapplicationandtheweb-basedapplet.Here
theyare.

Launching:
{Applet:Simplypresstheappropriatelink.
{Application:Firstdownloadtheapplicationandbesurethatitisunpacked.Yourweb
browsershoulddothisautomatically,butfollowtheinstructionsthatcanbefoundon
thedownloadwindow.Onceit'sunpacked,doubleclickonitandit'lllaunch(provided
youhaveaJavainterpreterinstalledinyourOS|ifyouusesomeversionofWindows32
thismaybeabitmorecomplicated;seenotesonthedownloadpage).
Quitting:
{Applet:Simplycloseallwindows,thiswillexityoufromthesimulation.
{Application:GototheFilemenuandselect\Quit"(Mac)or\Exit"(Windows).
4.3QuestionstoAddressandThingstoTry

Trytoanswerallofthequestionsbelow.Discussionmaterialisprovidedforsomeofthequesti
ons
intheAppendixattheendofthetext.Ifyouhavetroubleansweringotherquestions,askabout
theminclass.

1.PayosandpureversusmixedESSs.
a)TryalteringthepayosandseehowthisaectstheequilibriumofthemixedESS.
b)Findthevaluesofgain,loss,anddisplaythatproduceapureESSforbothHandD.You

shouldprobablylookatthemasratiosorinrelativeterms.
c)Whatgeneralizationscanyoumake.
d)IfyouproduceapureESS,aretherelativepayosthatyouareusingstillrealistic?Canyou

userealisticnumberstoproduceapureHawkESS?PureDove?Oraretotallyunbelievable
numbersrequired.

2.WhenthereisapureESS,wheredothe
tnesslinesforHandDconvergerelativetothe
frequencyofHawk.

3.Usingtheinsightsyougainedfromtryingtheprevioussetofquestions,alterallvaluesofc
osts
andbene
tsbutkeepthemconstantintermsofratioswithrespecttoeachother.
a)Iftheratiosremainthesame,doesitmakeanydierenceintheequilibrialfrequencies.
b)Shouldit|isratiowhatmattersorisitabsolutedierencethatmatters.

c)Wouldyouexpecttherelative
tnessesofthetwostrategiestobeequalordierent(andif
so,howdierent)whenamixedequilibriumisreached?HowaboutapureESS?Tryitand
see.
Chapter4.TheHawksandDovesSimulation
39

4.Whyisitthattherelative
tnessofHdoesnotchange(upordown)insimulationswhenH
increasesinfrequency?Recallthatwelearnedearlierthattheabsolute
tnessofbothHandD
decreaseasthefrequencyofHincreases.

a)Whatdoesthistellyouaboutmeasuresofrelative
tness.
b)Doyoustillthinkthatrelative
tnessisagoodmeasuretouseinevolutionarystudies.
c)Whyshouldthereshouldbecaseswherethe
tnessofastrategyincreasesasitsfrequency

decreases.

5.Imagineasituationwherelosinga
ghtcausessevereinjurybutthat
ghtingistheonlywayto
procureacriticalresource,withoutwhichreproductionisimpossible.
a)Inthissituation,whatisthe
tnessofanindividualplayingastrategythatdoesnot
ght
andthereforedoesnotobtaintheresourcebutlivesalongtime.

b)Comparedtotheindividualjustdiscussed,whatwouldbetheaveragerelativepayoto
anindividualplayinganalternativestrategythat
ghtsfortheresource.Assumethatmost
individualsofthisstrategydiein
ghtswithoutprocuringtheresource.However,someof
themaresuccessfulandleaveospring.

c)Ifdeathoccursina
ght,isitappropriatetousesequentialcontestgameslikeHawkand
Dove.

d)Therearemanycaseswheretherearehighlyescalatedcontestsleadingtoserious
ghtsthat
mightcausethedeathofoneofthecontestants.Forexample,maleelephantsealsengagein
suchcontestsoversectionsofabeachand\breedingrights"withthefemalesinthisarea.
Doesthatmeanthatmalesthatlosesuch
ghtsorthatdonotengagein
ghtinghaveno
tness?WhatdoesthistellyouaboutsimplegameslikeHawkandDove.

4.4Appendix:TheConceptofFitness

Whiletheunitofevolutionisthepopulation,nevertheless,selectionoccursbetweenindivi
duals
bearingalternativephenotypes(whichforourpurposeswillmeanindividualsusingdierentge
neticallybasedbehavioralstrategies
).Sincedierentphenotypesspringpartiallyfromalternative
alleles,selectioncanalsobeseenasacompetitionbetweenalternativealleles.Therearean
umber
ofwaystomeasurethesuccessofdierentindividualsoralleles.Allinvolvethe
tnessconcept.

Theterm
tnesscomesfromDarwinandWallace'sideathatanimalsthatsurvived(i.e.,were
most
t)weremostlikelytoleaveagreaternumberofospring.Whilethenotionofsurvival
primarilyhastodowithwhattheytermed\naturalselection,"wewillalsotoincludetheeectso
f
sexualselectiononanindividual'sreproductionwhenwediscuss
tness.1

Simplyput,
tnessisameasureofthenumberofcopiesofanindividual'sgenes,orifweare
consideringasinglegeneticlocus,thenumberofcopiesofanallele,thatareputintothenext
generation.

Actuallytogetaroundsomeoftheproblemsthatcanarisewhenanindividual'sospring(F1
generation)areinfertile(F2generation),themostformalanalysescountthenumberofgrand
children
(F2).Therearewell-knownexamplesofthissortofthing,termedhybridsterility|thinkofdo
nkey
andhorsecross.Ityieldsavigorous,valuableF1(mule)butnoF2|mulesaresterile.Neverthe
less,
incaseswherethereisnoreasontosuspecthybridsterility,itiscommonpracticesimplytoco
unt
thenumbersofospring.

1Forthepurposesofourpresentdiscussion,wewillleaveoutconceptsofindirectandinclusi
ve
tness.
Chapter4.TheHawksandDovesSimulation
40

FitnessisdenotedbyWwithsomeothernotationthatusuallyexplainswhose
tnessisbeing
considered(e.g.,W(Hawk)forthe
tnessofthestrategyHawk).Simplycountingthenumberof
ospringorgrandchildrengivesameasurethatcanbecalledabsolute
tness.

Evolutionisanumbersgameandsowhatreallymattersingenerationalcompetitionisnotthe
numberofospring(provideditismorethan0)buthowmanyanindividualproducesrelativeto
itscompetitors.Thesimplestwaytomakethiscomparisonistojustcomparehowmanycopiesof
eachalleleareproduced|forexample,strategyA(associatedwithalleleA)producesandaver
age
of1.7ospringwhilestrategyBproduces2.2.OnecaneasilyseethatstrategyBisdoingbetter.
Buthowmuchbetter.

Humanshaveaverygoodunderstandingofproportionalmeasurements|where1.7versus2.2
tellsussomething,sayingthatstrategyAisonly77%assuccessfulasstrategyBusuallytells
us
somethingthatweunderstandbetterwhenweareconsideringtheoutcomeofcompetition.Using
measuresofrelative
tnessdojustthissortofthing.Theyarbitrarilyde
nethemostsuccessful
typeashavingthereference
tness.Allothersaremeasuredasaproportionofthisreferencevalue.
Thus,

averagenumberofospringbyanystrategy

Fitness=W=
averagenumberofospringofmost
tstrategy

andfortheexamplewejustconsidered,

strategyBhadthegreaterabsolute
tnessat2.2;itbecomesthereference;

therelative
tnessofstrategyAistherefore1:7=2:2=0:77;

andtobecomplete,therelative
tnessofthereferencestrategyis2:2=2:2=1:0(ofcourse,it
willalwaysbe1.0).

Problem

6.Assumethatapopulationofasexuallyreproducingorganismspossesstwogeneticallydeter
mined
behavioralstrategies,AandB.Assumethat,takenasgroups,AandBstrategist's
tnesses
dieronlyasaresultofhowtheirbehavioraectstheirabilitytoreproduce(inotherwords,
theydieronlyinregardsofbeingAorBstrategists).Theseanimalsliveoneyearandthe
averagenumberofospringleftbyanAstrategistis0.85andforaBstrategistitis1.05.At
thestart,thereare850individualsofstrategyAand125ofstrategyB.

a)WhatarethefrequenciesofstrategiesAandB.
b)Whataretherelative
tnessesofstrategyAandB.
c)Whatwillbethefrequenciesofthetwostrategiesinthenextgeneration.
d)Atpresent,isthepopulationsizeasawholeincreasing,declining,orsteady.
e)Ifevolutioncontinuesandiftherelative
tnessesofthetwostrategiesremainthesame,

predictwhatwillhappentothepopulationasawhole|willitincrease,decreaseorremain
thesame.
Chapter5

Con
ictandOwnership:The
BourgeoisStrategy

Synopsis:ThischapterassumesthatyouhavethoroughlyinvestigatedtheHawksandDoves
game,thatyouunderstandthosestrategies,thatyoucancalculatepayomatrices,andthat
youunderstandpureandmixedESS.

Buildingonthisfoundation,wewillnowconsideranewstrategy,Bourgeois,whose
centralfeatureisthatownershipofaresourcedeterminesthebehaviorusedinaparticular
contest.IfaBourgeoisstrategistowns,itwilldefenditsownershipwithHawk-likeferocity
;
ifBourgeoisdoesnotown,itwillattempttoobtaintheresourceusingdisplaybutitwillnot
escalateto
ghting.

Youwillgetachancetocalculateapayomatrixforathreestrategygameandthenuse
thistoconsiderwhetherBourgeoisisstableagainsteitherHawkorDove.Thiswillprepare
youtousetothenextsimulationwhichlooksatevolutioninapopulationcontainingthese
threestrategies.

5.1Introduction

UsingtheHawksandDovessimulationyouexaminedthesolutionstoasimplegameinvolvingtwo
strategies.YouhavelearnedthatthisgameusuallyyieldsamixedESSbutundercertaincircum
stancescangiveapureESSforHawk
.Ontheotherhand,pureDoveESSsinvolvedunacceptable
assumptions.

IntheHawksandDovesgame,eachcontestinvolvedsituationswherethecompetingindividuals
either:

didnotpossesstheresourcepriortothecontest
and/oractedasifpreviousownershiphadnoeectontheoutcomeofagame.

Recallthattheconceptofownershipwasnotpartofthede
nitionofeitherstrategy.Now,there
arecertainlysituationswhereownershipisirrelevantforanumberofreasons.However,innu
merous
casesanimaldopossessresourceswhichothersmaysometimesattempttowrestfromthem.Inoth
er
cases,animalsseemtorespectownershipofaresource|theydonotbothertoattempttotakeit
fromanotherindividual.Aretheysimplybeingniceoristhisrespecttheresultofanevolutio
nary
calculationbasedonthebene
tsandcostsofrespectforownership.
Chapter5.Con
ictandOwnership:TheBourgeoisStrategy
42

Wecancertainlyusegametheorytoseeiftheresituationswherestrategiesthatrespectowner
ship
arestableorformamixedESSwithotherstrategiessuchasHawkandDove.Inthissectionwe
willde
nesuchastrategy,whichhasbeennamedBourgeoisbyMaynardSmithandconsideritin
gameswithHawkandDove.ThisadditiontoourHawksandDovespopulationwill

allowustoexamineastrategythatintroducessomeofthebehaviorsassociatedwithpossessio
n
ofresources,
giveusachancetoseewhathappenstopopulationwhenanew(previouslyunknown)populationar
rivesbymutationorimmigration
,and

itwillintroduceustosomeoftheevolutionaryintricaciesthatoccurwhenmorethantwo
strategiesarecompetingagainsteachother.
5.2De
nitionoftheBourgeoisStrategy

Bourgeoisisastrategyassociatedwithrespectfor\ownership"(i.e.,possession)ofaresou
rce.
Bourgeoisstrategists
ghttoholdontoresourcestheyalreadyown(i.e.,actlikeaHawk)and
theydisplayoverresourcesthattheydonotown.Inoursimpleexample,wewillthereforede
ne
BourgeoisashavingeitherHawk-likeorDove-likebehaviorcontingentonwhetheritortheoth
er
contestantownstheresource.Torecapitulate,ifBourgeois,then

whenanowner,
ghtlikeaHawktoholdterritoryandbewillingtoriskserious(
tnesslowering
|notdeath-causing)injury;

whennotanowner,donotriskinjuryandactlikeaDove.
ThisisincontrastwithbothHawkandDovestrategistswhoalwaysplaythesamestrategyregard
less
ofwhetherornottheyortheiropponentownaresource.

Problem

1.Doesownershipimplyterritoriality.

2.DoesBourgeoisseemtoyoulikeabehavioralstrategythatananimalmightreallyemploy.
Critiquethestrategy.

5.3PayosfortheBourgeois,Dove,andHawkGame

Wewillde
neBverysimplyinlightofHandDstrategies.WewillassumethatBhasa50.
chanceofowningaresourceinanycontest.Thus,inanycontestwithB,thereisa50%chancethat
itwillactlikeHawk(owns)anda50%chancethatitwillbeaDove(doesn'town).Tocontinueto
keepthingssimple,wewillassumethatasintheHandDcontest,youdon'tknowwhoyouare
playingagainsttillitstarts(elseHawkscouldavoidHawks,forinstanceandtheywouldbeapu
re
ESS).SeeTable5.1forthegeneralpayomatrix.

IfweinsertthesamedefaultpayostoHawksandDovesasusedwereusedintheprevious
exampleintotheequationsinthematrixabove,thenthepayomatrixforourthreestrategygame
isfoundinTable5.2.
Chapter5.Con
ictandOwnership:TheBourgeoisStrategy
43

Table5.1:ThegeneralpayomatrixfortheBourgeois,Dove,andHawkgame.

Hawk
Dove
Bourgeois
Hawk
E(H;H.
E(H;D)1
2[E(H;H)+E(H;D).
Dove
E(D;H.
E(D;D)1
2[E(D;H)+E(D;D).
Bourgeois
1
2[E(H;H)+E(D;H)]1
2[E(H;D)+E(D;D)]1
2[E(H;D)+E(D;H).

Table5.2:AparticularpayomatrixfortheBourgeois,Dove,andHawkgame.

Hawk
Dove
Bourgeois
Hawk
..25+50+12:5
Dove
0+15+7:5
Bourgeois..12:5+32:5+25

5.4IsBourgeoisaPureESS.

Noticethatthereisnosimplewaytoanswerthisquestionsincetheruleswelearnedearlierfor
comparingthepayosofdierentencounterswerefortwo-strategygames.However,athree-strate
gy
gamecanbebrokendownintosimplertwo-strategycontests.Sincewealreadyknowtheoutcome
ofHawkversusDove(forthesepayos),thecontestsofinterestareHawkversusBourgeoisandDov
e
versusBourgeois.IfinbothoftheseseparatelyBourgeoisisapureESS,thenitisreasonablet
o
concludethatBourgeoisisapureESSversusamixofHawkandDovesinceitcaninvadebothand
cannotbeinvadedbyeither.

ForBversusH:E(B;B)isgreaterthanE(H;B)(i.e.,+25isgreaterthan+12:5).Thus,for
anyfrequency,whenBinteractswithBthe
tnessconsequencestoBarebetterthanwhat
HreceiveswhenHinteractswithB(bothinteractionswilloccuratthesamefrequency).If,
forcompleteness,weturnthisaround:E(H;H)=..25whichislessthanE(B;H)=..12:5.
Thus,BisstableagainstH.

ForBversusD:E(B;B)=+25whichisgreaterthanE(D;B)=+7:5;ifturnedaround,
E(D;D)=+15whichislessthanE(B;D)whichequals+32:5.

Thus,BourgeoisisstableagainstbothHawkandDove.

You'llbeabletocon
rmthisresultforthesebene
tsandcostsbyusinganappropriateaccompanyingsimulation
.Youwillalsohavethechancetotryto
ndsetsofbene
tsandcostswhereB
isnotapureESS.Andyouwillbeabletodeepenyourunderstandingofhowapopulationmight
evolvewhenthreestrategiesarepresent.
Chapter6

AThreeStrategySimulation

Introduction

Synopsis:ThischaptercontainsadescriptionofhowtousethesimulationoftheBourgeois
versusHawksandDovesgame.Donotattemptthisgameuntilyouthoroughlyunderstand
the(seeChapter3)andyouarefamiliarwiththebasicoperationoftheHawksandDoves
simulation(seeChapter4.Thissimulationhasmanysimilarities.Alsobesurethatyou
understandthe\new"strategyBourgeoisinthepreviouschapter.Takethetimetoreview
thematerials
rstoryouwillnotgetmuchoutofthissimulationandyouwillprobablyhave
troubleansweringthequestions.

OnemajordierencebetweenthissimulationandtheHawksversusDovessimulation
isthatthereisnoplotof
tnessversusfrequency.Withthreestrategies,suchaplotis
diculttomake(requiringeitherthreeaxesor
xationofthefrequencyofonestrategy.
butmoreimportantly,unlikeatwostrategygame,theremaybenopureormixedESS
outcome.Dependingontheinitialconditions(payos,frequencies)anumberofoutcomes
arepossible|pure,mixedornoESS!So,wewillonlylookattheresultintermsofevolution.

ANotefromtheProgrammer:MaynardSmithandmanyothergametheoristsusuallyplotthree

strategygameevolutionresultsasbarycentricplots.Whileveryelegant,thesetakesomeget
ting

usedtoandsoIhavedecidedtousethemoreintuitiveplotsoffrequencyversustime.

6.1AbouttheSimulation

Onceyouhaveloadedthesimulationandhavemovedfromtheintroductorywindow,anewwindow

likeFigure6.1willappear.
Thiswindowisdividedintothreesections:

Theleftpanelcontainsabutton,\Instructions,"whichexplainshowtousethewindow.

Thecentralpanel,\InfoonCalc.ofPayos,"hasninebuttonseachlabeledwiththesymbolic
notationforaparticularpayo.Pressingoneofthesebuttonswilltellyouhowaparticular
payoiscalculated.

Note:AswiththeHawksandDovessimulation,youwillnotbeabletoaltertheactual
de
nitionofanystrategynorcanyoumodifytheformulaeusedtocalculatethepayosfora
Chapter6.AThreeStrategySimulation

Figure6.1:Theset-uppanelforthecostsandbene
tsintheHDBsimulation.

particularcontest.AsintheHawkandDovesimulation,theonlywaythatyouwillbeableto
modifythepayosisbychangingtheBene
tsandCosts.

Therightpanelcontainscontrolsforchangingbene
tsandcosts.Asusual,simplyenterthe
valuesyouwishfortheresourcevalue(GAIN)andtwotypesofcosts.Youshouldusethe
sameconventionsforassigningvaluestoGain,Injury,andDisplayaswithHawksandDoves
sincebothofthesestrategiesarefoundinthisgameandBourgeois.iscombinationofthe
twostrategies.

Thebuttonlabeled\ResettoDefault"willsetthegainsandcostsbacktotheirinitialvalues
whicharethesamedefaultvaluesthatweusedintheHawksandDovesgame.

Finally,pressingthered\Calc.Matrix"willsendyouontothenextwindowandwillcalculate
thepayomatrix.

Usethesameconventionsasbeforetoassignbene
tsandcosts:Bene
t0andCosts0.

Onceyouaresatis
edwiththebene
tsandcosts,presstheredbuttonandyouwillseethenext
windowFigure6.2whichreviewsthepayomatrix.

Theleftpanelgivesthreebuttons
{informationaboutusingthepage;
{abuttonthatwilltakeyoubacktothepreviouswindowtorevisethepayomatrixby
changingthebene
tsandcosts;
{Aredbuttonlabeled\Continue|SetFreqsandRun"|whichtakesyoutothenext
window.

Therightpanelcontainsthepayomatrix(gold)calculatedusingtheBene
tsandCostsyou
setonthelastpage.Thebluebuttonaboveeachpayocanbepressedtogiveyoutheformula
usedtocalculateeachpayo.
Chapter6.AThreeStrategySimulation

Figure6.2:ThepayomatrixfortheHDBsimulation.

Figure6.3:Thepanelforsettingtheinitialfrequenciesofthestrategies.
Chapter6.AThreeStrategySimulation

Whenyouaresatis
ed,presstheredbuttonwhichtakesyoutothenextwindow,Figure6.3.
Onceagainthiswindowisa\tryptic"(apologiestoallofthosegreatpaintersforappropriati
ng
theterm):

Theleftpanelhastwobuttons,
{informationonusingthepageand
{abuttonthatwillallowyoutogobacktoreviewthelastwindowincaseyouwantto
seethepayomatrixbeforeyousetstrategyfrequencies(fromthepreviouswindow,you
willbeabletoresetthebene
tsandcostsandtherebychangethepayosaswasnoted
above).
Thecenterpanelcontainsthreetext
eldsforenteringthefrequenciesofeachofthestrategies:

{Becertainthatthefrequenciesaddto1.0.Iftheydonotaddto1.0orifyouenternonnumerical
data
,youwillseeawarningwindowwhichyoushouldcloseandthenre-enter
yourdata.
{Ifyouwishtorunatwostrategygame(e.g.,HversusB),enteravalueof0.0forthe
strategyyouwishtoexcludebutbesurethattheothertwoaddto1.0.

Finally,therightpanelcontainstwocontrols,
{apull-downmenuthatallowsyoutosetthenumberofgenerationsintheevolution
simulation.Thedefaultis50,butexperiencewillshowthatinsomecasesyoumaywant
tousefewergeneration(togetabetterviewofthechanges)ormoregenerations(when
equilibriumhasnotyetbeenreached.
{andaredbuttonthatwhenpressedwilltakeyoutotheevolutionsimulation.
AswiththeevolutionsimulationintheHawksandDovesGame,therearetwoplots,seeFigure
6.4.Theleftisaplotoftherelative
tnessesandtherightisaplotofthestrategyfrequencies.
Akeyatthebottomgivesthecolorandsymbollabelsforeachstrategyandamessagewilltell
youhowmanygenerationswererequiredtoreachequilibrium(ifatall).
Afewthingstorememberwhenviewingtheplot:AswiththeHawkandDovegame,therearea
coupleofthingstonoticeabouttheplot.

First,
tnessisexpressedrelatively.Thus,italwayshasavaluebetweenzeroand1.0.

Second,beawarethattherearesomeroundingerrorsandsothegraphandhashmarksonthe
axeshavesomeslighterrors.Forinstance,the
tnessoftheHawklinedoesnotstartexactly
at1.0onthey-axisnordoesitendexactlyonthex-axisatafrequencyof1.0asitshould.
Nonetheless,theerrorsarenotlarge.
Third,belowthegraph,atextprintoutwilltellyouwhetherornotthereisapureormixed
ESSandifmixed,whattheequilibrialfrequencywillbe.UnliketheHawksandDovesgame,
thisprogramdeterminesequilibriumhasoccurredwhenthereisnochangeintwosuccessive
generationsinthefrequenciesofallthreestrategies.Pleasenotethatbiologically-speak
ing,
equilibriumwouldprobablyoccuratadierent,earliernumberofgenerations.Theprogram
doesnotassignequilibriumuntilthefrequencyofHremainsconstantto38placesfortwo
successivegenerations!
Finally,pleasenotethatwhenthegameisonlyplayedwithtwostrategies,e.g.,HandB,the
relative
tnessesofallthreestrategiesarestilldisplayed.Thisissimplytohelpyouenvision
whatwouldhappenifthethirdstrategywasadded.
Chapter6.AThreeStrategySimulation

Figure6.4:Relative
tnessandstrategyfrequencyplots.
Chapter6.AThreeStrategySimulation
49

6.2DierencesBetweentheAppletandApplication

Thereareafewdierencesbetweenthestand-aloneapplicationandtheweb-basedapplet.Here
theyare.

Launching:
{Applet:Simplypresstheappropriatelink.
{Application:Firstdownloadtheapplicationandbesurethatitisunpacked.Yourweb
browsershoulddothisautomatically,butfollowtheinstructionsthatcanbefoundon
thedownloadwindow.Onceit'sunpacked,doubleclickonitandit'lllaunch(provided
youhaveaJavainterpreterinstalledinyourOS|ifyouusesomeversionofWindows32
thismaybeabitmorecomplicated;seenotesonthedownloadpage).
Quitting:
{Applet:Simplycloseallwindows,thiswillexityoufromthesimulation.
{Application:GototheFilemenuandselect\Quit"(Mac)or\Exit"(Windows).
6.3QuestionstoAddressandThingstoTry

Thespeedofthissimulationwillallowyoutoanswerallofthesequestionsrapidly|taketheti
me
toconsidereachindetailandrecordyouranswersorthoughtsandquestionsinyourcoursenote
s
fordiscussioninclass.

Trytoanswerallofthequestionsbelow.Discussionmaterialisprovidedforsomeofthequesti
ons
intheAppendixattheendofthetext.Ifyouhavetroubleansweringotherquestions,askabout
theminclass.

1.a)SeehowBourgeoisdoesagainstjustHawk,justDove,and
nallyagainstboth.Usedefault
payovaluesandsetfrequencieseitherat50:50or0.33:0.33:0.33.
b)CouldyouusetherulespresentedearlierinSection2.4todetermineapureESSwithall
threestrategiesatonce.

2.Inasystematicmanner,startwithinitiallydierentfrequenciesofH,D,andB.
a)Forexample,tryHat0.9,Dat0.09and(therefore)Bat0.01.
b)ReversethefrequenciesofHandD.
c)TrynearlyequalfrequenciesofHandDandlowB.
d)SatisfyyourselfthatineachcaseBisstillanESS|canyoualterthevaluesofwinningthe

resource,injuryanddisplaycostsinanymeaningfulwaytopreventBfrombeingapure
ESS?Usethesamesortsofmodi
cationsthatyoumadeintheHandDgametomakeone
ortheother(inonecase,unrealistically)apureESS.
Chapter6.AThreeStrategySimulation

3.Moreaboutfrequencies:ReviewthesituationwiththedefaultpayomatrixandwithHat
0.9,Dat0.09,and(therefore)Bat0.01.
a)Setthenumberofgenerationsto10.DescribewhathappenstoH,DandBoverthistime.
b)Setthenumberofgenerationsto50.DescribewhathappenstoH,DandBbetween
generation10and50.
c)Setthenumberofgenerationsto150.DescribewhathappenstoH,DandBbetween
generation50and150.
d)RepeatthisexperimentwiththeinitialfrequenciesofHandDreversed(andthereforethe
sameinitialfrequencyofB).
e)Betweengenerations10and50,whatweretheapproximatefrequenciesofHandD.
f)WeretheythesameregardlessofwhetherornotyoustartedwithHorDat0.9.
g)Haveyouobservedthesefrequenciesbefore.
h)Whatisgoingonhere.
i)IfBisapureESSwhydoesittakesolongforitto
x.
j)WhatwouldyouneedtodotomakeB
xfaster,(givenastartingfrequency).

4.WhichstrategycouldyoumonitorinthisgametotellwhentheESSisreached.
Chapter7

SupplementaryMaterialfortheH
v.DandHv.Dv.BGames

7.1MoreontheHawkandDoveGame

WenowgeneralizetheanalysisofthecalculationsoftheHawk-DoveESSbymakingthecostsand
bene
tsinvolvedarbitrary.WemodifythepayosinTable3.2asfollows.

Table7.1:ThegeneralpayovaluesforHawkandDove.

Action
Bene
torCost(arbitraryunits.

GainResource
v
LoseResource
0
InjurytoSelf
i
CosttoDisplaySelf
d

Here,vispositiveandianddarenon-positivenumbersandthepayosareinunitsof
tness.We
nowcalculatethepayosinthevariousHawkandDovecontestsunderexactlythesameassumptions
asinSection3.4.Substitutingv,i,anddfortheirparticularvaluesof+50,..100,and..10in
(3.2)to(3.5)weobtain

11

E(H;H)=
v+
i

22
E(H;D)=1v..0=v
E(D;H)=0v+10=0
111

E(D;D)=
(v+d)+
d=
v+d:

222

SoforthisgeneralversionoftheHawkversusDovegame,thepay-omatrixis:
Chapter7.SupplementaryMaterialfortheH.DandHv.D.BGames
52
vv
Table7.2:ThegeneralHawkversusDovepayomatrix.

Opponent
FocalStrategyHawk
Dove
Hawk
1
2v+1
2iv
Dove
01
2v+d

PureESSs

RecallthatHawkisapureESSifeitherE(H;H)>E(H;D)orE(H;H)=E(H;D)andE(H;D).
E(D;D).UsingthevaluesinTable7.2,the
rstconditionbecomes

11

E(H;H)>E(H;D)(.

v+
i>0()v+i>0:

22

Thesecondconditionisequivalentto

11

E(H;H)=E(H;D)(.

v+
i=0()v+i=0

22

and

11

E(H;D)>E(D;D)()v>
v+d(.

v>d:

22
Butthislatterconditionisalwaystruesincevispositiveanddisnot.Thus,weconcludethatH
awk
isapureESSwheneverv+i0,thatis,wheneverthevalueoftheresourceisatleastasgreatas
thecostincurredbyinjury.

1
CanDoveeverbeanESS?WejustsawthatE(H;D)>E(D;D)because2v>d,soDovecan
neverbeapureESS.ThismakessensesinceanypopulationofDovescaneasilybeinvadedby
Hawks.
MixedESSs

InSection3.5,wesawthatforamixedESStoexist,bothstrategiesmusthavethesame
tness,
thatis,W(H)=W(D).IfhisthefrequencyofHawkatsuchamixedequilibrium,wedetermined
in(3.12)that

hE(D;D)..E(H;D.

.
:
(7.1.

1..hE(H;H)..E(D;H.

UsingthevaluesinTable7.2,thisbecomes

h1v+d..vv+2d..2v2d..v

=2==:
(7.2.

1..h11i..0v+iv+i
v+
22
Cross-multiplyingin(7.2)yields

h(v+i)=2d..v..h(2d..v.
h(v+i+2d..v)=2d..v
h(2d+i)=2d..v:
Chapter7.SupplementaryMaterialfortheH.DandHv.D.BGames
53vv
Nowsolvingforhweobtain

2d..v

h.
(7.3.

2d+i

ThisequationhowtheequilibriumfrequencyfortheHawkstrategyinamixedESSisafunction

ofthepayos.Thismakescomputingtheequilibriumfrequencyastraightforwardmatter.For

example,giventhepayosv=50,i=..100,andd=..10inthetext,weseethat

2(..10)..50)..70

h.
==0:583:

2(..10)..100..120

OfcoursethefrequencyoftheDovestrategywillbed=1..h=1..0:583=0:417.

ProblemsUseboththeHawksandDovessimulationandthematerialinthesectionaboveto
answerthefollowingquestions.

1.Usingtheinitialvaluesoftheresourcegainv=50,injurycosti=..100,anddisplaycost
d=..10,determinethatthepayomatrixinTable3.3inthetextiscorrect.Verifyusingthe
HawkandDovessimulationthattheequilibriumfrequencyforHawksinaMixedESSforthis
situationish=0:583aswascalculatedabove.

2.a)Nowincreasethevalueoftheresourcegaintov=60.Whathappenstoh.
b)Increasethevalueoftheresourcegaintov=80.Whathappenstoh.
c)Explaininonesentencewhythismakesbiologicalsense.

3.a)Resetthevalueofvto50.Nowincreasethecostofinjurytoi=..120.Whatistheeect
onh(comparedtothevalueofhinProblem1).
b)Nowincreasethecostofinjuryfurthertoi=..150.Whatistheeectonh.
c)Explaininonesentencewhythismakesbiologicalsense.
4.a)Resetthevalueofvto50andito..100.Nowincreasethedisplaycosttod=..20.What
istheeectonh(comparedtothevalueofhinProblem1).
b)Nowincreasethedisplaycosttod=..30.Whatistheeectonh.
c)Explaininonesentencewhythismakesbiologicalsense.

5.Leti=..100andd=..10,whatvalueofvwillproduceapopulationwith50%Hawksand50.
Doves?Usetheformulaforhin(7.3).Verifyyouranswerbyusingthesimulation.

6.a)Assumearesourcevalueisv=100,injurycosti=..120,anddisplaycostd=..20.Fillin
thevaluesinthepayomatrixbelowusingthegeneralformulasinTable7.2.Verifythese
valuesusingthesimulation.

Player2
Player1HawkDove
Hawk
Dove
Chapter7.SupplementaryMaterialfortheH.DandHv.D.BGames
54vv
b)WhyisHawknotapureESSinthisgame.
c)Determinehusingequation(7.3)verifyitusingthesimulation.
d)DeterminethesmallestvalueofvthatwillmakeHawkapureESSinthisgame.Whatish

ifyouusev..1inthesimulation?Whathappened.

7.a)ReconsidertheHawkandDovegame.Doublethesizeofallthecostsandbene
tsofthe
originalgame.Thatis,assumearesourcevalueofv=100,aninjurycostofi=..200,
anddisplaycostofd=..20.Fillinthevaluesinthepayomatrixbelowusingthegeneral
formulasinTable7.2.Verifythesevaluesusingthesimulation.

Player2
Player1HawkDove
Hawk
Dove

b)Eitherusethesimulationprogramoralgebraicmethodstodetermineh,theequilibrium
frequencyoftheHawkstrategyintheMixedESSforthisgame.
c)Comparethistotheequilibriumvalueofhintheoriginalgame(seeProblem1).Whatwas
theeectofthedoubling.

7.2MoreontheHawk,Dove,andBourgeoisGame

WecancarryoutthesamegeneralanalysisofHawk,Dove,Bourgeoiscontests.Againletvdenote
thevalueoftheresourcecontested,ithecostofinjury,anddthedisplaycost.Thepayosinthos
e
contestsinvolvingonlyHawksandDovesremainsthesameasinTable7.2.Recallthatweassume
thatBourgeoishasa50%chanceofowningaresourceanytimeitcompetessothatinanycontest
withBourgeois,thereisa50%chancethatitwillactlikeaHawk(owns)anda50%chancethat
itwillbeaDove(doesn'towntheresource).Therefore,thepayosingameswhereBourgeoisisa
contestantaregivenbelow.

.

11vi3vi

E(H;B)=[E(H;H)+E(H;D)].
++v=+

2
22244

hi
11vvd

E(D;B)=[E(D;H)+E(D;D)].
0++d=+

2
2242

.
11vivi

E(B;H)=[E(H;H)+E(D;H)].
+=+

2
22244

hi
11v3vd

E(B;D)=[E(H;D)+E(D;D)].
v++d=+

2
2242
11v

E(B;B)=[E(H;D)+E(D;H)].
[v+0]=:

2
22

Usingthesevalues,wecancreatethepayomatrixforthegeneralversionoftheHawk,Dove,
Bourgeoisgame(seeTable7.3).
Chapter7.SupplementaryMaterialfortheH.DandHv.D.BGames
55
vv
Table7.3:AgeneralpayomatrixfortheHawk,Dove,Bourgeoisgame.

Hawk
Dove
Bourgeois
Hawk
v
2+i
2
v
3v
4+i
4
Dove
0v
2+dv
4+d
2
Bourgeois
v
4+i
4
3v
4+d
2
v
2

IsBourgeoisaPureESS.

Aswasnotedinthetext,thereisnosimplewaytoanswerthisquestionsincetheruleswelearned
earlierforcomparingthepayosofdierentencounterswerefortwo-strategygames.However,
athree-strategygamecanbebrokendownintosimplertwo-strategycontests.Sincewealready
knowtheoutcomeofHawkversusDove(forthesepayos),thecontestsofinterestareHawkversus
BourgeoisandDoveversusBourgeois.IfinbothoftheseseparatelyBourgeoisisapureESS,
thenitisreasonabletoconcludethatBourgeoisisapureESSversusamixofHawkandDovesince
itcaninvadebothandcannotbeinvadedbyeither.

ForBourgeoisversusHawk:
vvivi

E(B;B)>E(H;B)(.

>
+
(.

..
>0:

24444

Thislastinequalityisalwaystruesincevispositiveandiisnon-positive.SoHawkcannot
invadeBourgeois.Further,

vivivi

E(B;H)>E(H;H)(.

+
>
+
()0>
+
()0>v+i:

442244

Thus,BourgeoiscaninvadeHawkwhenevertheinjurycostexceedsthevalueoftheresource.
Inthiscase,BourgeoisisstableagainstHawk.

ForBourgeoisversusDove:
vvdvd

E(B;B)>E(D;B)(.

>
+
(.

>
:
24242

Thisisalwaystruesincevispositiveanddisnot.Further,

3vdvvd

E(B;D)>E(D;D)(.
+
>
+d(.

>
()v>2d:

42242

Again,thisisalwaystruesincevispositiveanddisnot.Thus,Bourgeoisisstableagainst
Dove.

Thus,BourgeoisisstableagainstbothHawkandDoveindividuallyaslongas0>v+i,thatis,as
longastheinjurycostisgreaterthantheresourcevalue.You'llbeabletocon
rmthisresultand
thatBourgeoisisstableagainstcombinationsofthesestrategiesintheexercisesbelow.
Chapter7.SupplementaryMaterialfortheH.DandHv.D.BGames
56
vv
ProblemsUsingboththeHawk,Dove,Bourgeoissimulationandthematerialwedeveloped
above,answerthefollowingquestions.

8.a)Usingtheinitialvaluesoftheresourcegainv=50,injurycosti=..100,anddisplaycost
d=..10,verifythatthepayomatrixwecalculatedinclassforthissituationiscorrect.You
willhavetoresettheinitialcost-bene
tvalues.

b)VerifythattheBourgeoisstrategyisESS.

9.Inthepreviousproblemwesawthatifv=50,i=..100,andd=..10,thenBourgeoiswasa
pureESS.Thus,theBourgeoisshouldbeabletoinvadeapopulationofHawksorDoves,and
neitherHawksnorDovesshouldbeabletoinvadetheBourgeois.

a)TestthisbysettingtheinitialfrequenciesofHawkto0.999,Doveto0,andBourgeoisto
0.001.CantheBourgeoisinvadethispopulationofHawks?IsBourgeoisanESS?Ifso,how
manygenerationsdidittake.

b)NowsettheinitialfrequenciesofHawkto0,Doveto0.999,andBourgeoisto0.001.Can
theBourgeoisinvadethispopulationofDoves?IsBourgeoisanESS?Ifso,howmany
generationsdidittake.

c)Comparetheresults:DidtheBourgeoishaveaneasiertimeinvadingtheHawksorthe
Doves?Isthiswhatyouwouldhaveguessed.

d)NowsettheinitialfrequenciesofHawkto0.001,Doveto0,andBourgeoisto0.999.Can
theHawksinvadethispopulationofBourgeois?IsBourgeoisanESS?Ifso,howmany
generationsdidittake.

e)NowsettheinitialfrequenciesofHawkto0,Doveto0.001,andBourgeoisto0.999.Can
theDovesinvadethispopulationofBourgeois?IsBourgeoisanESS?Ifso,howmany
generationsdidittake.

f)DotheBourgeoishaveahardertimeinvadingamixedpopulation?Settheinitialfrequencies
ofHawkto0.49,Doveto0.49,andBourgeoisto0.02.CantheBourgeoisinvadethis
populationofHawksandDoves?IsBourgeoisanESS?Ifso,howmanygenerationsdidit
take?Wasitharder.

g)NowsettheinitialfrequenciesofHawkto0.495,Doveto0.495,andBourgeoisto0.01.Can
theBourgeoisstillinvadethispopulationofHawksandDoves?IsBourgeoisanESS?Ifso,
howmanygenerationsdidittake.

10.UsethegeneralpayomatrixfortheBourgeoisgametoanswerthefollowingquestions.
a)Supposethatthecostofdisplayisd=..10,asusual.WhatvalueofvwillmakeE(D;D).

E(B;D).
b)Doessuchavalueofvmakebiologicalsense.
c)Startingwithapopulationof0:33Hawks,0:33Doves,and0:34Bourgeois,isBourgeoisa

pureESSinthiscase?Ifnot,isthereamixedESS.

11.UsethegeneralpayomatrixfortheBourgeoisgametoanswerthefollowingquestions.
a)Supposethatthecostofdisplayisd=..10andthevalueoftheresourceisv=50.What

injurycostiwillmakeE(H;B)=E(D;B).
b)Doessuchavalueofimakebiologicalsense.
c)Startingwithapopulationof0:33Hawks,0:33Doves,and0:34Bourgeois,isBourgeoisa

pureESSinthiscase?Ifnot,isthereamixedESS.
Chapter7.SupplementaryMaterialfortheH.DandHv.D.BGames
57
vv
12.Supposewesetthedisplaycosttod=0butleavev=50andi=..100unchanged.DoDoves
fareanybetter?IstheDovestrategyanESS.

13.a)Setthevalueofvto100.Leavethecostofinjuryati=..100andresetthedisplaycostto
d=..10.IsthereapureESS?Ifso,whatisit?Ifnot,whatisthemixedESS?Howdoyou
tellfromthegraph?Whatproportionsofeachstrategyappear.

b)YoushouldhavefoundamixedESSinthepreviouspart.HowdoesthismixedESSdepend
ontheinitialfrequenciesofthevariousstrategies?SettheHawkandBourgeoisstrategiesto
0.5andtheDovestrategyto0inthefrequencydialogbox.Nowrunthesimulation.Doyou
getthesamemixedESS.

c)TrysettingtheHawkandBourgeoisstrategiesto0.4andtheDoveto0.2inthefrequency
dialogbox.Nowrunthesimulation.DoyougetthesamemixedESS.
d)ExplainbiologicallywhytherecanbedierentESSswiththesepayos.Hint:Lookbackat
thepayomatrix.

14.a)CanHawkeverbeapureESS?Wemightthinksoiftheinjurycostisnottoolargerelative
tothevalueoftheresource.UseTable7.2toshowthatE(H;H)>E(B;H)whenever
v+i>0.InthissituationHawkscannotbeinvadedtheBourgeois.

b)Underthesameassumptions,showthatE(H;B)>E(B;B).Thus,Hawkscouldinvadethe
Bourgeois.

c)Setthevalueofvto120.Leavethecostofinjuryati=..100(sov+i>0)andthedisplay
costatd=..10.Startingwithapopulationof0:33Hawks,0:33Doves,and0:34Bourgeois,
isHawkapureESS?Ifso,howmanygenerationsdidittake?Ifnot,whatisthemixedESS.

d)Whathappensifwesetthevalueofvto101.Leavethecostofinjuryati=..100(so
v+i>0)andthedisplaycostatd=..10.IsHawkapureESS?Ifso,howmanygenerations
doesittake?Doesthismakebiologicalsensewhencomparedtothepreviouspart.

15.a)Assumethatv=50,i=..100,d=..10.SupposethatyousettheinitialfrequenciesofHawk
to0.5,Doveto0.5,andBourgeoisto0.Whathappens?IseitherstrategyanESSoristheir
amixedESS.

b)Usetheequation
2d..v

h.

2d+i
wedevelopedfortheHawkandDovegametoestimatetheproportionofHawksinthemixed
ESS.Doesyourcalculationforhmatchtheequilibriumgraphforthisproblem.
Chapter8

WarsofAttrition:FixedCost
Strategies

Synopsis:Inthegameswehaveconsideredpreviously,weexaminedstrategiesthatused
ghting
(i.e.,conteststhatpotentiallyinvolvedinjury)tosettlesymmetricalcontests(e.g.,Haw
k
andsometimesBourgeois).WealsoconsideredthestrategyDove(andBourgeoiswhenit
didnot\own")whichsettledcontestswithotherDovesthroughdisplay.Indisplaysthereis
nochanceofinjuryalthoughtherecertainlyarecostsintermsofenergy,time,orriskof
beingpreyedon(injuryfromanon-contestant).

Inthischapterwewilllookathowsimplesymmetricalcontestsbetweenindividualsthat
onlydisplaymightbesettledwithoutresortto
ghting.Thesecontestsarereferredtoas

\symmetricalwarsofattrition."
Wewill
rstexaminethequestionofwhetherornotany
xedcostdisplaycanbeevolutionarilystable
.Wewillshowthat
xedcoststrategistsarenotevolutionarilystable.This
willleadustoaconsiderationofamixedESSsolutioninthenextsection.

8.1Introduction

Therearesituationswhere
ghtingdoesnotoccurinacontestoveraresource.Howthencould
ownershipbesettled.

Onerulewouldbesimplytocooperate:splittheencounters.However,thismightonlyseem
reasonablewhenthecontestantskneweachotherandkepttrackoftheircontestssothat
divisionswouldbeequal(fair).Thisisnotsomethingthatmostanimalscouldorwould
do.Wehumanscoulddothis,butwhatifyouwereunlikelytoeverencounterthesame
opponentagain?Gametheorymodelingofcooperativebehaviorlikethiscanbedoneusing
the\prisoner'sdilemma"game.Whatotherpossibilitiesarethere.
Anothersolutionwouldbetosettlethecontestbysomeasymmetry,usuallydetectableina
display.However,thissortofsettlementprobablyonlyworksiftherewassomeback-upto
thedisplay.Thus,thisleavesthechanceforescalation.Whatiftheloser(theonewith
thesupposedlyinferiordisplay)callsthewinner'sblu?So,ifweallowasymmetries,weare
leftwithamodelthatmayrequire
ghtingeitherasaresultoftheinabilityofthepartiesto
Chapter8.WarsofAttrition:FixedCostStrategies
59

discernthewinnerofanevenlymatchedcontestortobackupthe\honesty"ofthewinner's
display.

Isthereanotherwaytodecidewithoutresortingto
ghtingorsharing?Onesimpleandtime
honored(nottomakeajoke)solutionisawaitinggameorwarofattrition.Wehaveseen
ictseitherbyimmediate
anexampleofthisinDove.RecallthatDovestrategistssettlecon
withdrawalorbyunescalateddisplayingcontests.Thus,warsofattritionincludethemeans
DovesusetosettlecontestsagainstotherDoves(butnotagainstHawks!).

Inthissection,wewillreviewsomeoftheideasaboutsettlingcontestswithdisplays.

8.2WaitingGamesandtheirCurrencies

Inawaitinggame,thecontestantwhoiswillingtowaitthelongestwins.Thinkofthesilly,oft
en
tragicdramasofpeople(oftenpooranddesperate)whoentermarathondancecontests(didyoue
ver
seetheclassicmovieTheyShootHorses,Don'tThey?)towinprizemoneyorthosewhotrytowin
acarbykeepingtheirhandsonit,remainingawake,andstandinglongerthananyothercontesta
nt.

Suchwaitinggameshavealsobeendubbedwarsofattrition,althoughtheydonotneedto
bestrictlyanalogoustothehorrible\real"warofattritionwherethewinningsideistheonew
hose
armies,cities,andpopulationshaven'tbeenunacceptablydecimated.Inouranalysisofwars
of
attrition,wewillbeconcernedwithindividuals(actingasproxiesforstrategies)competin
gagainst
eachother.Wewillnotbeinterestedinsocietalorothergroupcompetitionasinthemilitary
concept(althoughthisanalysiscouldalsobeusedwithgroups).Typesofwarsofattritiontha
tare
meaningfultoabehavioristincludeconteststhataresettled

purelybywaiting(orsomeothertypeoftime-dependentdisplay)or

bydepletionofresourcessuchasenergy.Theassumptionisusuallymadethatwhilecosts
areinvolvedinwaiting,injury(intheconventionalsense)isnot|animalsdropoutofcontest
s
beforetheyareseriouslyharmed.
CurrenciesforWaitingGames

Thefundamentalcurrencyofwaitinggamesis,ofcourse,
tness.Butaswediscussedearlier,
tness
consequences,measuredaschangesinnumbersofgrandchildren,areusuallyhardtoassessfor
simple
behaviorssuchasdisplays.

Tosavetime,weusesomeotherfunctionsuchasnetbene
tornetvalue(gainafterallcosts
arefactoredin).Recallthatto
ndthenetvalue,weneedtohavesomemeasureofthevalueofthe
resourceandameasureofthecostsassociatedwithcompetingfortheresource.Inwaitinggame
s,
thesecostsareabsorbedbyboththewinnerandloser.

Costsastimeorenergy:Obviously,costsandbene
tsmustbeenumeratedthesameway;
theymusthaveacommoncurrency.Let'sstartourconsiderationwithcosts.Inawaitinggame,
theonlycostsaredisplaycosts.Thus,thereisnoescalationto
ghtingandnoinjury.

Whatarethesecosts?Anytimeananimalisdoingonething,suchasdisplaying,itisnotdoing
somethingelsethatmightbehelpingits
tness.Themoretimeitdisplays,thelesstimeitmight
haveforlookingforfood.Theremayalsobecostsduetoexposure{animalsthataredisplaying
areoftenfarmorevisibletotheirpredatorsorotherpotentialenemies.

However,wewillconsiderdisplaycostsastheextraenergy(comparedtodoingnothing)thatan
organismusestoperformthedisplay.Aswasdiscussedinthesectiononoptimality(review),t
hese
Chapter8.WarsofAttrition:FixedCostStrategies
60

costsareusuallyafunctionoftime.So,wecanmakethesimpleassumptionthatcostandtimeare
related("timeismoney"):

Costs/t;
(8.1.

wheretistime.Forourpurposeswewillassumethatcostsincreaselinearlywithtime.SO

Costs=kt
(8.2.

wherekisaproportionalityconstantequalingtheenergycost(x)oftheuseofoneunitoftime
(t).Alinearrelationbetweenenergycostandtimeisprobablythegeneralruleinanimalrepet
itive
animaldisplays.Agoodexampleiscallinginsectsandfrogs[ForrestandGreen,1991].Howeve
r,
notethattherearecaseswherecostisnotalinearfunctionoftime,butwe'llkeepthingssimpl
e
andstickwith(8.2).

Anexample:Let'slookatanexampleofhumanbehaviortounderstandtheideaofcontests
andcosts.Supposeyouarehikingandyouarelookingforasuitablesheltertospendthenight.
Ifyouarriveatashelterthatisalreadyoccupiedeitherbysomecritter{let'ssayabearora
rattlesnake,oryouandanotherhikerarriveatthesametime,aconteststartsoverwhogetsthe
shelter.Thesecontestsaresettledbydisplays|nokillings,snakebitesormaulingsallowed
.You
trytoscareoutthebearorsnakewhilekeepingarespectfuldistanceoryoudothetypicalhuman
thingstotrytogettheotherhikertoleave(butlet'snotbetoohuman|no
ghts!).

Let'sfocusonthecontestwithanotherhumansincethecostsaremostlikelytobesymmetrical
andsincegamesareusually(butnotalways)consideredascostsbetweenconspeci
cs.Thecostof
thecontestisyourtimeandpatienceasyoudiscussorpostureoverwhoisgoingtogettostay.
Eventuallyoneyouquits.Youhavebothpaidthesamecostinthecontest.Andwecouldhave
measuredthiscosteitherintermsoftimeorenergy.See(8.2).

Now,whataboutbene
t?Sincewemeasuredcostastimeorenergy,weneedareasonableway
toevaluatetheshelterinoneofthesecurrencies.Assumethesheltersareequallyspacedinte
rms
ofthetimeittakestoreachthem.Occupyingasheltermeansthatyouhaveavoidedthecostof
havingtowalktothenextshelter.Soinasimplisticbutusefulsense,thevalueofashelterequ
als
thecostyouwouldhavepaidtohiketothenextshelter.AfamousquotefromthevenerableBen
Franklincrystallizesthisidea:"apennysavedisapennyearned".

Avoidtheerrorofthinkingaboutthecostsofsearchingascontestcosts.Thecostsareonlytho
se
associatedwiththeactualcontest|theyinvolvethetimeandenergyandperhapsriskinvolved
in
givingthe"evileye"tothebear,rattlesnake,orotherhiker.Searchcostsareusedonlytoobt
aina
reasonable,easilymeasuredvalueoftheshelter.Noticethattheactualsearchoccursoutsid
eofthe
contest|theconteststartswhenthesearchends.

8.3IsthereanESSforaWarofAttrition.

Toanswerthisquestion,wewillmakethefollowingassumptions(identicaltothoseweassumed
for
Doveearlier):

Wewillassumethattherearenoasymmetriesinthecontestantsbeyondthefactthatone
mightbepreparedtowaitordisplaylongeror,expendmoreenergy(theseareallrelated,as
wesawearlier).Thus,animalsdonotgiveupbecausetheyperceivethattheiropponentis
strongerasaresultofthedisplay.Sowewillrefertothesecontestsassymmetricalwarsof
attritionorsymmetricalwaitinggames.
Chapter8.WarsofAttrition:FixedCostStrategies
61

Wewillassumethatanimalshavenoknowledgeofhowlongtheiropponentsaregoingto
display|strategiesaresetatthestartofthegameandplayedout.
Theysimplywaitordisplayuptoacertainperiodoftimeorburnuptoacertainamountof
energy.
{Ifafocalanimal'sopponenthasnotyetquitwhenthefocalanimalreachesitslimit,it
loses.
{Iftheopponentquitsbeforethelimitisreached,thefocalanimalwins.
{Sincebothanimalsquitatessentiallythesametime(determinedbythelosersincethe
winnerispreparedtogoonlonger),bothwillpaythesamecostinthecontest(although
forthewinneritwaslessthanitwaswillingtopay).

{Thismeansthatthevalueoftheresourceisdiscountedtothewinnerbythecostof
displayingforit.Wehavediscussedthisideaearlierwhenwedevelopedageneralformula
fordeterminingpayosinacontest(see(2.1)).
Forfurtherinformationontheseassumptions,theinterestedandmathematicallyinclinedin
dividualshouldconsult
[BishopandCannings,1978].

Theanalysisofthisgameisrathercomplicatedmathematically.Theinterestedreaderisadvi
sed
toconsult[Smith,1974and1982]and[BishopandCannings,1978]forelegant,detailedexplan
ations
oftheproblem.Whatfollowsisasynopsisoftheirworkwithcommentaryandexpansiondesigned
toaidastudentwhoisnewtogametheoryandmathematicalmodeling.Ihavedealtwiththe
mathematicsbypresentingboththecalculusequations(withaexplanationsandinsomecases,
derivations)andaparallelsetofdiscretesolutionswhichmybemorecomfortabletostudents
who
arenotfamiliarwithcalculus.Theyhavetheaddedadvantagethatastudentcaneasilyusethem
onaspreadsheetto
ndanswersthatapproximatethosegivenbythecalculus.

8.4CanaFixedWaitingTimeStrategybeaPureESS.

CostsandBene
ts

First,let'sde
nethecostsandbene
ts:

ThecontestedresourcehasacertainabsoluteorgrossvaluewhichwewillcallV(following
MaynardSmith).SinceIliketothinkofthingsintermsofenergy,let'ssaythatitisacertain
valueinjoules.

Wewillusetwosymbolsforcost.
{Anycostissymbolizedasx.Theseaccumulateaccordingtosomefunctionoftime(see
(8.3)immediatelybelow).

{Thecostthateachcontestanthaspaidatthemomentthecontestendsism.Thiscost
isdeterminedbytheloser,butbothpayit.
Wewillassumethatdisplaycosts,x,increaseinalinearmanner:
DisplayCosts=kt
(8.3.

Caution:FollowingMaynardSmith,wewillconsidercostsaspositivevaluesandsubtract
themfromthegrossresourcevalue.ThisisadierentconventionthenweusedintheHawks,
Doves,andBourgeoisgamesbutthe
nalmathematicsarethesame.
Chapter8.WarsofAttrition:FixedCostStrategies

Thus,ifwehavetwocontestants,AandBwhoarewillingtodisplayfortimest(A)andt(B),
thenwecande
netheirdisplaycostsas:

x(A)=kt(A)andx(B)=kt(B.
(8.4.

ThegaintothewinnerofanycontestwillbethevalueoftheresourceVdiminishedbythecost
ofgettingit.Rememberthatwewillsymbolizethecumulativecostpaidattheterminationofth
e
contestasm:

NetGain=V..m:
(8.5.

Now,recallthattheloserpaysthesamedisplaycostasthewinnersincetheloserdetermineswh
en
thecontestwillend,i.e.,whenhequitsx=mandsotheloserpays:

Loss=..m;
(8.6.

Payos

Wecannowconstructalistofpayosfordierentcontests.Thisisnotthesameasthepayo.
matriceswehaveseenbefore,butwewillusetheinformationinittoconstructsimilarmatrice
sa
bitlater.

Table8.1:Payosforvariouscontests.

StrategyandOutcomeChangein
tnessforAChangein
tnessforB
m(A)>m(B),thereforeAwins
V..m(B.
..m(B.
m(A)<m(B),thereforeBwins
..m(A.
V..m(A.
m(A)=m(B),thereforeStalemateResourcepossessionisdecidedrandomly,eachwinshalfofth
etime
0:5V..m(B.
orequivalently
0:5V..m(A.
0:5V..m(B.
orequivalently
0:5V..m(A.

Hopefully,Table8.1makessense.EachrowcorrespondstothepayosforplayersAandBfor
agivencost(lengthofdisplay)thatthecontestantsarewillingtopay.

Thus,inthe
rstrowstrategyAiswillingtopaymore(displaylonger).Awillwinevery
contest.

ThecostthatApaysinwinning(andBinlosing)isdeterminedbyB.Thus,bothcontestant's
costsare..m(B).

Thesecondrowissimplytheconverse|hereBwinsbybeingwillingtodisplayforalongerperiod
oftimeandnowthecostthatispaidiswhatAwaswillingtopay.Thethirdrow|atie,istheonly
onewithanynewtricksinit.WhatifbothstrategiesAandBpickthesametime(cost)topay.
Forthesakeofsimplicity(butnotrealityashepointedout),MaynardSmithstipulatedthatth
e
winnerwillbedeterminedatrandom.Thus,eachcontestantwouldwin50%ofthetime.Inevery
Chapter8.WarsofAttrition:FixedCostStrategies

contestboththewinnerandloserwillpaythecostm(A)=m(B).Substitutingintotheformula
forassigningpayosthatwelearnedearlier,

payoifbothdisplaythesame=0:5(V..m(A))..0:5m(A.
=0:5V..0:5m(A)..0:5m(A.
=0:5V..m(A.

whichistheexpressiongiveninTable8.1.

AnalysisofaGameBetweenFixedCostStrategies

OK,let'sconstructthepayomatrixtoseeifacertainstrategyisapureESS.Wewilluseourusual
procedureofassumingthatonestrategyisestablishedandtheotherinvadesinverylownumber
s.

Situation1:Aninvaderwillingtopaymorearrives!

De
netwostrategies,AandB.LetBbewillingtopayaslightlyhighercostthanA,i.e.,
m(B)=m(A)+m
(8.7.
wheremisthesmalladditionalcostthatBiswillingtopay.Thus,
m(B)>m(A.
(8.8.
MakeAthecommonstrategyandBarareinvader.Noticethatthisisexactlythesamescenario
wehavealwaysdiscussedindeterminingwhetherornotastrategyisapureESS.Thus,

nearlyallofstrategyA'sinteractionsarewithotherAstrategistswhile
nearlyifnotallofstrategyB'sinteractionsarewithAstrategists.
Let'sconstructapayomatrixusingtheformulaeinTable8.1above.
WheneitherAmeetsAorBmeetsB,contestsaresettledatrandomsincebothcontestants
arewillingtopayeitherm(A)orm(B),respectively.
BalwaysbeatsAintheircontests,thus,E(A;B)=0andE(B;A)=V..m(A).
ThepayomatrixisgiveninTable8.2.

Table8.2:PayomatrixforanInvaderBwillingtopaymorethanA.

A
B
A0:5V..m(A)0
BV..m(A)0:5V..m(B.

UsingRule1for
ndingApureESS1
,weseethanAcannotresistinvasionbyBandtherefore
ybetemptedtoconcludethatB
AisnotApureESS.Lookingatthematrixabove,youmaybrie
isanESS.Butlookcloser.

1SeeChapter2,Section4.
Chapter8.WarsofAttrition:FixedCostStrategies
64

Situation2:Sameoldsameold:Aninvaderwillingtopaymorearrives!

OK,assumethatBhastakenoverandisverycommon.Nowthebigquestion:Whatifanother
strategy(we'llcallitC)thatwaitsjustabitlongerthanBshowsup?Theanswer,ofcourse,is
thatwewillhavearepeatofthesituationwhenBinvadedA!Thus,Cwillnowsuccessfullyinvade
BandsoBisnotapureESS.Ifyoucontinuetofollowthislogic,youmaycometotheconclusion
thatastrategythatiswillingtopayanin
nitecostwouldbeapureESS.Notsofast.

Situation3:TheseQueuesareGettingTooLong!

Imaginethatourpopulationcontinuestobeinvadedbyindividualsthatarewillingtowaitlon
ger
towin.Accordingto(8.3),thecostsareincreasingwithlongerwaits,butthevalueofthereso
urce
isstillthesame.Thus,thenetgainforwinningisbecominglessandlessthelongeronewaitsto
win.

Imaginethatwe
nallygettoawaitingtimethatissolongthatitisgreaterthanhalfthevalue
oftheresource,i.e.,

m(Long)0:5V:
(8.9.

Now,thisisstillawinningvaluewithrespecttotakingtheresourcecomparedtoanytimethati
s
shorterthanitis.Let'ssayanewmutantappearsthatdoesnotwaitordisplayatall.

ProblemsAnswerthesequestionsbeforegoingon:

1.Constructapayomatrixforagameoflongwait(wherem(LongWait)0:5V)versusno
display.Explainhowyouworkedouteachpayo,referringtothepayosinTable8.2when
appropriate.

2.ExplainwhetherE(Long,Long)willbepositiveornegativenumber.

Backtothefuture

AversionofthepayomatrixthatyoushouldhavegottenisinTable8.3.

Table8.3:PayosforLongwaitversusNoDisplaywhenm(Long)>0:5V.

LongNoDisplay
Long<0(negative)V
NoDisplay0
0:5V

OK,nowifLongrepresentsastrategywheredisplaytimesaremorecostlythan0:5V,willit
bestableagainstinvasionbyindividualswhosimplydonotdisplay?Onceagain,Longiscommon
,
NoDisplayisrare.Lookingdownthe
rstcolumnofTable8.3whichshowsthemostcommon
interactionsforeachstrategy,wecanseethatNoDisplaycaninvadeoncethedisplaysgetcost
ly
(long)enough.

OneotherpointaboutNoDisplay.Noticethatif0:5Vm(Display),thenthematrixlookslike
Table8.4insteadandApopulationofNoDisplaycanbeinvaded!
Chapter8.WarsofAttrition:FixedCostStrategies

Table8.4:PayosforDisplaywaitversusNoDisplaywhenm(Display)<0:5V.

DisplayNoDisplay
Display>0(positive)V
NoDisplay0
0:5V

Conclusions

ThisexercisehasshownusthatthereisnopureESSinthewaitinggame.Wehaveseenthatno
displayscanbeinvadedbyincreasinglymorelengthy(costly)displaysuntilthepointwheret
hecost
ofthedisplayexceeds0.5,theresourcevalueatwhichpointnodisplaycaninvadeagain!

Rock,Scissors,Paper

Itisoftenpointedoutthattheoutcomewehavejustseenhascertainsimilaritiestothechild'
sgame
Rock,Paper,Scissors.Recallthatinthatgame(whichyoumayhaveplayed)therearethreepure
strategies(rock,paper,orscissors).Herearetheirde
nitions:

RockbreaksScissors:E(R;S)=+1,E(S;R)=..1,andE(R;R)=0.
ScissorscutsPaper:E(S;P)=+1,E(P;S)=..1,andE(S;S)=0.
PapercoversRock:E(P;R)=+1,E(R;P)=..1,andE(P;P)=0.

ThepayomatrixisgiveninTable8.5.

Table8.5:PayosforRock,Paper,Scissors.

RockScissorsPaper
Rock0+1..1
Scissors..10+1
Paper+1..10

Aswiththewaitingtimeswehavejustinvestigated,clearlynoneofthesestrategiesarepure
ESSs(usethe\lookdownthecolumn"rule.Doyouremember(fromyourchildhood)thebestway
towinoratleastsurviveinthisgame?We'llcometoitinamoment.

8.5AMixedESSSolutiontotheWaitingGame

So,howaboutour\warofattrition"game?Fromtheprevioussectionitshouldbecleartoyou
thattherearesituationswhereanypurewaitingtime(purestrategy)canbeatanyother(butno
t
allothers).Thus,thereisnopureESSsolutiontothewarofattrition.However,couldtherebe
a
mixedESS.

Foranoverviewoftheanswertothisquestion,let'sstartoutwiththeRock,Scissors,Paper
game.Itssolutionwillhavesomeparallelstheoneforourwarofattritionwhichwewillseeont
he
Chapter8.WarsofAttrition:FixedCostStrategies
66

nextpage.Butitwillalsohaveoneveryimportantdierence,whichwewillexplaininthenext
section.Nevertheless,let'scontinuewithRock,Scissors,Paper.

IfyouplayedRock,Scissors,Paper,asachild,youmayrememberthatyoucouldnotwinifyour
opponentsknewwhichstrategyyouweregoingtopick.Forexample,ifyoupickRockconsistentl
y,
allyouropponentwouldneedtodoispickPaperands/hewouldwin.Achilddiscoversquickly
thatifsheorhedoesn'tknowwhattheopponentwillpick,thenthebeststrategyistopickRock,
Paper,orScissorsatrandom.Inotherwords,theplayerselectsRock,Paper,orScissorswitha

1
probabilityof3.Thiswasprobablyhowyouconsistentlybeatinexperienced,youngerplayers
(who
tendtoemploythesamepurestrategyrepeatedlyuntiltheycatchon).

Itshouldbeobviousthatifyoudoknowwhatyouropponentislikelytodo,thenpickinga
strategyatrandomwithaprobabilityof1isnotbethebestthingtodo(unlessthatisthestrate
gy
3
youropponentisusing!).

1
IfyouplayedRock,Scissors,Paperwithaprobabilityof3strategyasachild,youmayremember
thatwhenyouplayedthegameagainstanothersavvyplayer,youonlywonhalfthetime.Butthe
otherplayerdidnotwinanymoreoftenandifsomeoneelsetriedadierentstrategy,heorshe
didnotdoaswell.PlayingeitherRock,Paper,orScissorsatrandomwithaprobabilityof3
1in
eachgameisamixedESS.Itismixedsinceitinvolvesplayingthreedierentstrategiesata
xed
(equilibrial)probability.ItisanESSsinceitisbothofhigher
tnessthananyalternativepureor
mixedstrategy.

Nowbacktoourwaitinggame.UnlikeRock,Scissors,Paper,potentiallythereareanin
nite
numberofpurestrategies(eachadierentwaitingtime)insteadofjustthree.Nevertheless,in
the
nextsectionwewillseethatthesolutionhasoneimportantparalleltoRock,Scissors,Paperi
nthat
thesolutionrequiresamixedstrategy.
Chapter9

AMixedESSSolutiontotheWar
ofAttrition

Synopsis:Thischapterwillshowthataparticularmixedstrategythatiscomposedofall
possibleacceptablecosts,eachtobeplayedatauniquefrequencyisevolutionarilystablein
thesymmetricalwarofattritionagainstanypurestrategy(uniquemaximumcost)orother
mixofpurestrategies.Wewilltermthestablemixedstrategyvar.Wewillseethatvaris
characterizedby:

aconstantprobabilityofcontinuing(orquitting)fromonecosttothenext,

theprobabilityofcontinuingisgovernedbythevalueofthecontestedresource,
theresultofaconstantrateofcontinuing(orquitting)isanegativeexponentialdistributi
on
ofquittingcosts|mostvarstrategistsquitatrelativelylowcosts.

Theapproachinthischapterwillbeto
rstreviewtheideaofamixedESS,
thenshow(usingsomesimpleandfullyexplainedcalculus)howwediscoveranequationthat
describesanequilibrialmixofallpossiblemaximumcosts.

Finally,usingthebasicruleswelearnedearliertodetermineanESSandsomesimplecalculus
andgraphs,wewillshowthatthisequilibrialmixisalsoevolutionarilystable.
Pleasenote:Thisisthemostmathematicalchapterofthetext.Itmustbesobecausewe
willneedtoderiveanequationthatdescribespotentiallyanin
nitenumberofbehaviors(an
in
nitenumberofdierentmaximumacceptablecosts).In
ndingthisequationandlaterin
showingthatvarisanESS,wewillmakeuseofsimpledierentialandintegralcalculus.I
havetriedtoexplainwhythesetechniquesareusedandfurther,toexplainhowtheyareused
sothatanyinterestedstudent,regardlessofwhetherornottheyarefamiliarwithcalculus,
shouldbeabletofollowthearguments.Asimportantly,Ihopetoconvincestudentsofthe
bene
tstoanybiologistgainedbyunderstandingbasiccalculus.
Chapter9.AMixedESSSolutiontotheWarofAttrition
68

9.1TheBasicsofaMixedESSintheWarofAttrition

Inthepreviouschapterwesawthatinthesymmetricalwarofattrition,eachuniquecostxthat
ananimalispreparedtopay(ortimeitiswillingtodisplay)isapurestrategy.Thus,thereare
potentiallyanin
nitenumberofpurestrategieseachde
nedbyadierentcostx.Wealsolearned
thatnopurestrategyisanESSinthewarofattrition.Giventhis,couldtherebeamixedESS.

InlookingforthismixedESS,wemustrealizethatanypurestrategyisacandidateforinclusio
n
inthemixedESS.Infact,weexpectthateverypossiblepurestrategyshouldbelongtothemix(i
.e.,
allpossiblemaximumacceptablecostsshouldsupportthemix).Thereasonforthisissimple|w
e
learnedearlierthatundertherightcircumstances,any
x(cost)strategycanincreaseand/ormixes
ofthesestrategiescanappear|it'sjustthatnoneoftheseareevolutionarilystable.So,wee
xpect
thatanystablemixwillcontainallpossiblestrategiesassupportingstrategies.

De
nitions:

Apurestrategyisde
nedassomeuniquemaximumacceptablecostbetweenzeroandin
nity.
Supportingstrategiesareallpurestrategiesthataremembersofanequilibrialmix.See[Bis
hop
andCannings,1978].Asynonymforsupportingstrategyiscomponentstrategy.

Incharacterizingamix,wemustknowthelikelihoodthatagivenplayermightencountereach
ofthesesupportingstrategies.Whileitispossiblethatthesefrequenciesarethesameforea
ch
supportingstrategy,itwouldseemfarmorelikelythatmany,ifnotall,supportingstrategie
swould
occurattheirownuniquefrequencies.Theonlyrulesarethat

allofthesefrequenciesmustaddupto1.0(sincetheyformthewholepopulation.

andofcourse,thefrequenciesforeachsupportingstrategyaresuchthateachendsupwith
thesame
tness.
Thus,wecansummarizethemixas

mix=prob(cost(a1))+prob(cost(a2))++prob(cost(an))=1
(9.1.

whereeachaiisasupportingstrategyandprob(cost(ai))iseitherthefrequencyofthestrate
gyin
thepopulationortheprobabilitythatamixedstrategist\adopts"thatparticularcostinagi
ven
contest.

Noticethelastpoint|aswelearnedearlierwhenweconsideredtheHawksandDovesgame,
therearetwowaystoproduceanequilibrialmix.Tothislist,we'lladdathird.Apopulationth
at
isevolutionarilystablecouldbe

apopulationofpurestrategists,eachpurestrategyisatitsappropriateequilibrialfreque
ncy,
or
apopulationofmixedstrategists,eachofwhomcanpotentiallyplayallstrategiesofthe
equilibrialmixattheappropriatefrequencies.Thus,inagivencontestamixedstrategist
usessomemechanismtoadoptaparticularmaximumacceptablecostatthecorrectfrequency.
uenceswhatitwilldothenexttime,or
Whatitadoptsinonecontestinnowayin
apopulationthatisamixofsupportingpurestrategists(eachattheappropriateequilibrial
frequency)andmixedstrategists(sincetheyplayeachsupportingcostattheequilibrialfre
quency
).Totakethisastepfurther,themixedstrategistscouldevenbe\incompletemixes"
solongastheycomplementedeachotherandthenetresultwasthat<B>inthepopulationas
awhole,thechanceofanyindividualbeinginacontestwithanystrategysupportingthemix
wasalwaystheequilibrialvalueforthatstrategy.
Chapter9.AMixedESSSolutiontotheWarofAttrition
69

Thislastpointisveryimportant,solet'smakeitonemoretime.Allthatmattersforapopulati
on
tobeevolutionarilystableisthat

the
tnessesofeachsupportingstrategymustbeequal.Asalways,iso
tnessinnowayrequires
thateachsupportingstrategyactuallyhasthesamefrequency!

themixisimmunefrominvasion.
Itdoesn'tmatterhowtheappropriatemixisobtained|whetheritisfrommixedstrategyindivi
duals,
purestrategyindividualsinthecorrectfrequencies,orsomecombinationofthetwo.

9.2SupportingStrategyProbabilitiesatEquilibrium

Aswestarttolookforawaytodescribethemix,weseemtofaceadauntingtask.Weexpect
allpossiblecoststobemembersofthismix.Thus,thereareanin
nitenumberofsupporting
strategieseachpotentiallyatitsownuniquefrequency.So,wewillnotbeabletousethesimpl
e
techniqueto
ndthemixthatwelearnedwithHawksandDoves.Insteadofonlyneedingacouple
oflinearequationsto
ndtwofrequencies,weneedafunctionthatcangiveusthecorrectfrequency
foranin
nitenumberofdierentsupportingstrategies!Whatfollowsisageneraldescriptionofthe
methodsusedbyMaynardSmith[1974]to
ndthisfunction.

Pleasereadthissectioncarefully.Itsetsthefoundation,establishesterminology,and
reviewsthemathematicsusedthroughouttherestofourtreatmentofthewarofattrition.

Weshallusethepayothataspeci
csupportingstrategyexpectstoreceivewhencompeting
againstthemixto
ndthefunctionthatgivesustheequilibrialfrequencyofeachstrategysupporting
themix.So,westartwithapurestrategythatisamemberofthemix.

Thisfocalsupportingstrategyiswillingtopayuptocostx=m.

So,we'llrefertoitas
x(x=m).

Now,imaginethat
x(x=m)isabouttoplayaseriesofcontestsatrandomagainstother
individuals(supportingstrategies)fromthatmix.So,
x(x=m)'sopponentinanycontestcanbe
understoodtobemixitself.

Remember,itdoesn'tmatterwhether
x(x=m)'sopponentisapureormixedstrategist:in
eithercaseweknowtheresultisthatonlyonestrategycanbeplayedbyanopponentinagivengam
e
andthechancethataparticularstrategy(maximumcost)willbefacedisgivenbythecharacter
istics
oftheequilibrium.

Let's
ndanequationforthepayo
x(x=m)receivesagainstanyothersupportingstrategy
inthemix,E(
x(x=m);mix).Starting,ingeneralterms

E(
x(x=m);mix)=LifetimeNetBene
tstoFocalStrategyinWins
..LifetimeCoststoFocalStrategyinLosses(9.2.

Areminder,gentlereader:Rememberourpurposeinwritingequationsforlifetimenetbene
tand
costwillbetoextractafunctionthatpredictsthefrequencyofeachcomponentstrategyofthe
mix.

In
ndingtheseequations,let'smakeoneotherimportantassumption:theresourcehasaconstant
valueinanygivencontest
.Youmaythinkthatitisobviousthataresourcevalueshouldbe
constantinanycontest.Therecertainlyaremanyifnotmostsituationswherethisistrue.But
,
thinkforamomentandyou'llrealizethatitisquitepossibleforaresourcetobecomedepleted
duringacontest.Forexample,individualsmaybecontestingaresourcethatoneofthemalread
y
Chapter9.AMixedESSSolutiontotheWarofAttrition
70

isusingorthatnaturallydepletesinvalueovertimeindependentofanythingthecontestants
are
doing.Or,whiletwoindividualscontestforaresource,itispossiblethatanotherindividua
l,perhaps
amemberofadierentspeciesdepletesit.So,whilereasonableformostsituations,theassumpt
ion
thatforacontestVisaconstantmaynotalwaysbejusti
ed.

FindingExpectedLifetimeNetBene
ts

Bene
tsareonlyobtainedbythefocalstrategistwhenshewins|i.e.,whenthefocalstrategistis
willingtopayahighercostthanheropponentfromthemix(x<m,wheremisthecostthefocal
strategistwillpay).

NetBene
tto
x(x=m)inawin=V..x;
(9.3.

whereVistheresourcevalueandxisthecosttheopponentfrommixiswillingtopay.

Unfortunately,(9.3)isnotsucientforourneeds.Thecomplexityofthewarofattrition
intervenes!Recallthatthemixiscomposedofanin
nitenumberofcomponentstrategies.
x(x.
m)onlyfacesoneofthesesupportingstrategiesinanygivencontest.Thus,(9.3)onlydescrib
es
thenetgaininonespeci
ccontest.Youshouldrealizethatthisparticularcontestwillprobablybe
quiteraregiventhemanydierentstrategiststhat
x(x=m)couldfacefromthemix.Thus,one
particularcontestanditsbene
tswillhavelittleifanyimportantlifetimeeecton
x(x=m)'s
tness.Singlecontestscannotdescribethenetbene
tthatthefocalsupportingstrategyexpectsto
gainfromalargenumber(alifetime)ofcontests.

Togetanaccuratemeasurementoflifetimenetgains,weneedtotakeintoaccountalltypes
(costs)ofconteststhat
x(x=m)willwin(i.e.,thosewheretheopposingstrategyxisatmostm.
andtheprobabilityofeach.

NetBene
t=[(V..x)(Probabilityoffacingx)]:
(9.4.

0xm

Notethatthisisanin
nitesumbecausethereareanin
nitenumberofdierentcosts(strategies.
y
xbetween0andm.Tohandlethistypeofsum,wewillneedtousecalculus,whichwenowbrie
consider.Wewillreturnthequestionoflifetimenetbene
tsoncewehaveintroducedtheappropriate
calculustechniques.1

9.3AnIntroductiontoIntegration

Theprobabilityoffacingaparticularstrategyxisdeterminedbyaprobabilitydensityfunct
ion
orpdf.Wewillillustratetheideabyexamininganumberofdierentsituations.

Situation1:

Supposethatthereareanin
nitenumberofstrategiesxthatplayersadoptwith0<x1.Assume
furtherthateachstrategyxwith0<x0:5isjustaslikelytooccurasanyotherinthisinterval.
Finally,assumethateachstrategyxwith0:5<x1istwiceaslikelytooccurasanystrategyx
with0<x0:5.Ifweassignapositiveprobabilityptoanystrategyxin(0;0:5],thensincethere
areanin
nitenumberofotherequallylikelystartegies,they,too,wouldhaveprobabilityp.But
thensummingtheseprobabilitieswouldproduceanin
nitelylargevalue,not1.Forthisreason,
theprobabilitythatanyparticularstrategyxisencounteredmustbe0.

1MaterialinthenextsectionwaswrittenbyKevinMitchell;don'tblameKennethPrestwich.
Chapter9.AMixedESSSolutiontotheWarofAttrition
71

Abetterquestiontoaskiswhattheprobabilityisoffacingastrategyxthatfallswithina
certaininterval.Forexample,letpdenotetheprobabilityofencounteringastrategyxthatl
iesin
theinterval(0;0:5].Iclaimthatp=1=3.Here'swhy:

Thestrategiesin(0:5;1]aretwiceaslikelytobeencounteredasthosein(0;0:5],sotheyhave
atotalprobability2p,i.e.,twicetheprobabilityofthosein(0;0:5].

Butsincetheonlystrategiesarethosebetween0and1,wemusthave1=p+2p=3p,or
p=1=3.

Suppose,next,thatwewantedtoknowtheprobablityofencounteringastrategyxwith0.
x0:25?Well,sinceallthestrategiesin(0;0:5]areequallylike,andsincetheirtotalprobabi
lityis
p=1=3,andsince(0;0:25]isexactlyhalftheinterval(0;0:5],weconcludethattheprobablit
ythat
0<x0:25mustbehalfofp,thatisp=2=1=6.Inthesamewayyoushouldbeabletoshowthat
theprobablitythat0:75<x1is1=3.

Figure9.1:Ageometricrealizationorprobabilitydensityfunctionp(x)forsituation1.

4=3
.
................................................................................
..............................................
2=3.
................................................................................
..............................................
0

Thefunctionp(x)inFigure9.1givesageometricalrepresentationofsituation1.Ithasthe
followingproperties.
p(x1)=p(x2)foranytwostrategiesx1andx2in(0;0:5]becausetheyareequallylikely.
p(x1)=p(x2)foranytwostrategiesx1andx2in(0:5;1]becausethese,too,areequallylikely.
Butp(x2)=2p(x1)foranystrategyx2in(0:5;1]andx1in(0;0:5]becausestrategyx2istwice
aslikelyasx1.
Theregionunderthegraphofp(x)ontheintervalfrom0to0.5isrectanglewhoseareais

121

233Whichistheprobablitythatxisin(0;0:5].

=
Theregionunderthegraphofp(x)ontheintervalfrom0.5to1isrectanglewhoseareais

142

233whichistheprobablitythatxisin(0:5;1].

.
Theregionunderthegraphofp(x)overtheentireintervalfrom0to1is33

12=1whichis
+
theprobablitythatxisin(0;1].

Moregenerally,theareaunderp(x)ontheinterval[a;b]representstheprobabilityofastrat
egy
xwhereaxb.

Thefunctionp(x)isanexampleofaprobabilitydensityfunctionorpdf.Suchfunctionsmust
satisfytwoconditions:

1.p(x)0forallx,

0.00.51.0
Chapter9.AMixedESSSolutiontotheWarofAttrition
72

2.thetotalareaunderp(x)mustbe1.

Foragivenxthefunctionp(x)measuresthecomparitiveorrelativelikelihoodofstrategyx.T
hisis
whythegraphofpistwiceashighon(0:5;1]asitison(0;0:5].

Siutation2:

Assumethattheonlystrategiesxarethosesuchthat0x2andthatallofthesestrategiesare
equallylikely.Sincetheyareallequallylikely,p(x)mustbeconstanton[0;2],sothatthere
gion
underthegraphisarectangle.Sincetheareaunderthepdfover[0;2]mustbe1,thetheconstant

1
height(fortherectangle)mustbe2becausethebaseis2(seeFigure9.2(a)).

Figure9.2:(a)Theprobabilitydensityfunctionp(x)whenallstrategiesbetween0and2areeq
ually

1
likely.Theareaunderthegraphfromatobis2(b..a).

1.0

1.0

0.5.
................................................................................
................................................................................
................................................................................
............
................................................................................
................................................................................
................................................................................
...........
..
....
.
..
....
.
0.5
.
....
..
...
....
...
...
...
...
...
...
...
...
...
...
...
...
...
...
..
....
..
...
....
...
...
...
...
...
...
...
...
...
...
...
...
...
...
.
0.0

0.0

012
0ab2

Sinceallstrategiesareequallylikely,theprobabilitythataparticularstrategyxliesin[
0;1]is

1
just2orhalfthetotalprobablity.Ofcoursethiscorrespondsexactlytotheareaunderp(x)in
Figure9.2(a)overtheinterval[0;1]:itisa11rectangle.Infact,ifbisanynumberin[0;2],
2
thentheprobabilityofencounteringastrategyxintheinterval[0;b]istheareaunderthecur
ve
from0tob,thatis,b1
b
.Theprobablitythatoneencountersastrategywithwaitingtimeless
=
22
thanorequaltoxiscalledthethecumulativeprobabilitydistributionofquittingtimes,
andisdenotedbyP(x).NotetheuppercasePforthecumulativefunctionandthelowercasepfor
thedensityfunction.Inthisexample,P(x)isjusttheareaoftherectanglefrom0toxwithheig
ht

,so

P(x)=x=2;0x2:

Ifwewantto
ndtheprobabilityofencounteringastrategyxbetweenaandb,wecould
nd

1
b..a
theareaoftherectanglefromatob.Itsbasewouldbeb..aandtheheight2,soitsareais2.
ButthisprobablitycanbeexpressedmoregenerallyusingP(x).Theprobabilitywewantisjust
theareafrom0tobminustheareafrom0toaasinFigure9.2(b).Butthisisjust

bab..a

P(b)..P(a)=
..
=:

222

Situation3:

Supposeagainthattheonlypossiblestrategiesxarethosesuchthat0x2andthatthepdfis
p(x)=x=2.Noticethatthisisalegitimatepdfsincep(x)0andtheregionunderthegraphofp

1
isatrianglewhosearea221=1(seeFigure9.3(a)).

WhatisthecumulativedistributionP(x)forthissituation?Well,P(x)isjusttheareaunderp
from0toxwhichisjustatrianglewithbasexandheightp(x)(seeFigure9.3(b)).Thus,

11xx2

P(x)=
xp(x)=
x
=
:

2224
1
2
Chapter9.AMixedESSSolutiontotheWarofAttrition
73

Figure9.3:(a)Theprobabilitydensityfunctionp(x)=x=2for0x2.(b)P(x)=x2=4isthe
areaunderpfrom0tox.

1.0
.
................................................................................
................................................................................
................................................................................
........................................1.0
.
................................................................................
................................................................................
................................................................................
........................................
....
...
.
.
....
..
...
...
...
...
...
...
...
...
...
...
...
...
...
...
...
...
...
...
...
...
...
...
..
0.5

p(x.

0.0

0.0
0.00.51.01.52.0
0
x2
Forexample,theprobabilitythatastrategyxliesintheinterval[0;0:5]is

(0:5)2
0:25

P(0:5)===0:0625;
44

whiletheprobabilitythatxisintheinterval[0:5;1:5]is

(1:5)2
(0:5)2
2:250:25

P(1:5)..P(0:5)=..=..=0:5:

4444

Situation4:

Supposenowthatthepossiblestrategiesarerestrictedto0x1andthatp(x)=3x2(see
Figure9.4(a)).Clearlyp(x)0,sotoshowthatp(x)isapdf,weneedtoshowthatthearea
underthiscurveis1.Buthowdowe
ndtheareaofacurvedregion.

Figure9.4:(a)Theprobabilitydensityfunctionp(x)=3x2for0x1.(b)P(x)isthearea
underpfrom0tox.

.
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0

0
010101

Wecan'tdirectlyusetheareaformulaofarectangletodeterminethearea,butwecanuseit
indirectlytoapproximatetheareaunderthecurve.Supposewedividetheinterval[0;1]into
ve
subintervalsofequalwidth

1..0

x==0:2:

5
Chapter9.AMixedESSSolutiontotheWarofAttrition
74

Approximatetheareaunderthecurveoneachsubintervalbyusingarectanglewhoseheightis
determinedbyevaluatingpattheright-handendpointofthesubinterval(seeFigure9.4(b)).
These
veright-handendpointswillbex1=0:2,x2=0:4,x3=0:6,x4=0:8,andx5=1.Soin
thiscase,the
veheightswouldbep(x1)=p(0:2),p(x2)=p(0:4),p(x3)=p(0:6),p(x4)=p(0:8),
andp(x5)=p(1).Thus,approximateareais

p(xk)x=p(0:2)0:2+p(0:4)0:2+p(0:6)0:2+p(0:8)0:2+p(1)0:2

k=1

=0:2[p(0:2)+p(0:4)+p(0:6)+p(0:8)+p(1).

=0:2[3(0:2)2+3(0:4)2+3(0:6)2+3(0:8)2+3(1)2.
=0:2[0:12+0:48+1:08+1:92+3.
=0:2(6:6.
=1:32:

Ifweusedtenrectanglesinsteadeachwithwidthx=0:1(seeFigure9.4(c)),theapproximation
isevenbetter.Thistime,theareais

10

p(xk)x=p(0:1)0:1+p(0:2)0:1++p(0:9)0:1+p(1)0:1

k=1

=0:1[p(0:1)+p(0:2)++p(0:9)+p(1).

=0:1[3(0:1)2+3(0:2)2++3(0:9)2+3(1)2.
=0:1(11:55.

=1:155:
Thesameprocessusingone-hundredrectangles(nodrawingforthis!)yieldsanapproximation
of
1.01505andwithone-thousandrectanglestheapproximationis1.0015005.Itappearsthatthe
se
approximationsaregettingcloseto1asthenumberofrectanglesgetslarge.

Mathematiciansde
netheexactareaunderthecurvetobethelimitofthisrectangularapproximationproccessast
henumberofrectanglesnbecomesin
nitelylarge,

limp(xk)x:

n!1
k=1

Wedenotethelimitofthissummationprocessmorecompactlyas

Z
1

p(x)dx:

Thisisreadas,\Theintegralofp(x)from0to1."Theintegralsign,,isanelongatedS,a
remindertousthatanintegralisreallyasum.Thelowerandupperlimitsofintegration(here0
and1,respectively)arethebeginningandendingpointsoftheintervalwherethesumistaking
place.Theexpressionp(x)dxismeanttoremindusofp(x)x,theareaofarectangleofheightp(x.
andwidthx.Thinkofp(x)dxasbeingtheareaofin
nitesimallythinrectangleofheightp(x).

Inourparticularcase,withp(x)=3x2
,itappearsthat

Z
Z
1
3x
2dx=1;
0
Chapter9.AMixedESSSolutiontotheWarofAttrition
75

andthisis,infact,correct.TheFundamentalTheoremofCalculustellsusthatundercertain
circumstancessuchintegralscanbeeasilyevaluatedusingfunctionscalledantiderivative
s.Infact,
thecumulativedistributionfunctionisjustanantiderivativeofp(x).Inthisparticularsi
tutation
usingcalculus,P(b),thatis,theareaunderp(x)=3x2overtheintervalfrom0tob,isgivenbyt
he
formula

Z
b

P(b)=3x
2dx=b3:
0

SinceP(b)=b3
,thenthecumulativedistributionfunctionisP(x)=x3
,whichcalculusstudents
willrecognizeasanantiderivativeofp(x)=3x2
.Usingmethodsdevelopedinintegralcalculus,such
antiderivativescanbefoundforawidevarietyoffunctions.

Usingthenotationofintegrals,wecanexpresstheprobabilityofencounteringastrategyxin

R
theinterval[a;b].Thisisjusttheareaunderthecurvep(x)onthisinterval,or
ab
p(x)dx.Butwe
sawthatearlierthatthisisjusttheareafrom0tobminustheareafrom0toa(asinFigure9.2
(b))so,

ZZZ
bba

p(x)dx=p(x)dx..p(x)dx=P(b)..P(a):
(9.5.

a
00
b
ItiscustomarytousethesymbolP(x.
a
todenotethedierenceP(b)..P(a).Sowewouldwrite
b
Z
Z
b

p(x)dx=P(x)
.
:

a
a

Inthecaseofsituation4,withp(x)=3x2andP(x)=x3
,theprobabilitythatxliesinthe
interval[a;b]is
b
Z
Z
b
23=b33
3xdx=x
..a:

a
a

Forexample,theprobabilitythatxliesintheinterval[0:2;0:8]is

Z
Z
Z
b
0:8
0:8
3x
2dx=3x
2dx=x
3
.
=(0:8)3
..(0:2)3=0:512..0:008=0:504:
a
0:2

0:2
Amorerealisticsituation:

Generallyspeaking,strategiesarenotrestrictedto
niteintervalssuchaswehaveusedinthe
previousexamples.Thatis,xcantaketakeonanynonnegativerealvalue,x0.Thatmeans
thatapdfp(x)mustbede
nedonthein
niteinterval[0;+1),notjustsome
niteintervalsuch
as[0;1]or[0;2].Ontheotherhand,theareaunderthiscurvemuststillbe1sinceitrepresents
the
probabilityofencounteringsomestrategy,thatis,

Z
1

p(x)dx=1:

R
Suchexpressionsareevaluatedusinglimits.We
rstevaluatetheexpression0
b
p(x)dx.Thenwe
seewhathappenstothisexpressionasbgetsin
nitelylarge.Thenotationforthisis

Z
b

limp(x)dx:

b!1
0
Chapter9.AMixedESSSolutiontotheWarofAttrition
76

Here'sasimpleexample:Supposethatthepdfwasp(x)=1=(x+1)2forx>0.Toshowthat
thisisapdf,weneedtoshowthattheareaunderthecurve(seeFigure9.5)is1.Thatisweneed
toshowthat
Z
11
dx=1:

0(x+1)2

Figure9.5:Theprobabilitydensityfunctionp(x)=1=(x+1)2forx0.Thetotalareaunderthis
in
nitelylongcurveis1.

0
.
..........
......
................................................................................
................................................................................
................................................................................
................................................................................
................................................................................
................................................................................
................................................................................
................................................................................
................................................................................
................................................................................
....................................
01234
Itturnsoutthatthecumulativedistributionfunction(orincalculusterms,anantiderivati
ve2
forp(x)isP(x)=1,
1
.Usingthisandlimitstoevaluatetheintegralabove.

x+1
ZZb
11
b
1
1
dx=lim
dx=lim1..
.
(9.6.

0(x+1)2
b!10(x+1)2
b!1x+10

.

11

=lim1....1..:(9.7.

b!1
b+10+1

1
Nowasbgetslargeb+1approaches0.So

Z
Z
1
.

1
11

dx=lim1....1..=(1..0)..(1..1)=1

0(x+1)2
b!1
b+10+1

asexpected.
Ourexcursiontocalculusisnowcompleteandwereturntothediscussionoflifetimebene
ts.

FindingExpectedLifetimeNetBene
ts,Continued

Beforeourcalculusinterlude,wewereattemptingtogetanaccuratemeasurementoflifetimen
et
gains,weneedtotakeintoaccountalltypes(costs)ofconteststhat
x(x=m)willwinandthe
probabilityofeach.Letmbethespeci
cmaximumcostthatourfocalcontestant(
x(x=m).
willpay.Thenourfocalcontestantonlywinsthoseconstestsinwhichtheopponent'sstrategy
xis
between0andm.So

NetBene
t=[(V..x)(Probabilityoffacingx)]:
(9.4.

0xm

Wesawthatwecouldapproximatesuchsumsbydividingtheinterval[0;m]upintonpieceseach
ofwidthx,so(9.4)becomes

NetBene
t=(V..xk)p(xk)x:

k=1

1
..2
..1
2Calculusstudentsnotethatthegeneralantiderivativeforp(x)==(x+1)isP(x)=..(x+1)+c.

(x+1)2

WemustchoosecsothatP(0)=0,becausetheprobabilitythatanopponentquitingatcostlessth
an0is0.
Chapter9.AMixedESSSolutiontotheWarofAttrition
77

Here,asinourearliersituations,p(xk)xrepresentstheapproximateprobabilityofencounte
ring
opponentstrategieswillingtopayroughlycostxk.ThisprobabilityismultipliedbyV..xkto
obtain
thenetbene
taccruingfromsuchanencounter.Togettheexactnetbene
t,wetakethelimitas
thenumberofsubintervalsnbecomesin
nitelylarge.

lim(V..xk)p(xk)x:

n!1
k=1

Fromourcalculusexcursion,werecognizethislimitisanintegral.Sowecanre-express(9.4)
,the
equationforthenetbene
ttofocalsupportingstrategyversustheequilibrialmixas

Z
m

NetBene
t=(V..x)p(x)dx:
(9.8.

ExpectedLifetimeCostsforLosses

Bene
tswerethehardpartoftheE(focalsupportingstrategist;mix)equation.Now,themuch
simplerequationforlifetimecoststofocalstrat
x(x=m)incontestsitlosestothemix(i.e.,a
mixedstrategyopponent).

Asbefore,thelogicissimplE:c
x(x=m)loseswheneverx,thecosttheopponentfrommixin
anyparticularcontestiswillingtopay,isgreaterthanm.Allofthesecontestsendwithacost
of
m.Therefore,foranyonelosingcontest,

Costto
x(x=m)ofaLoss=..m:
(9.9.

So,unliketheequationfornetbene
t,thecostsinanylossarealwaysthesame.But,we'renotdone
becauseaswithnetbene
ts,weneedtotakeintoaccounttheproportionofthetime
x(x=m.
encountersanopponentthat(inthiscase)itlosestointhemix.So

CostofLosses=mprobabilityoflosingtostrategieswithxm

Sincelosesoccurwhenever
x(x=m)encountersastrategyxwithx>m,thentheprobability
that
x(x=m)losesis
Z1
Q(m)=p(x)dx:
m

Thus,thelifetimecostsoflosingtothemix,i.e.,losingtoamixedstrategistis

Z
1

CostofLosses=mQ(m)=p(x)dx;
(9.10.

wheremisthemaximumcostthatourfocalsupportingstrategywillpayandthefunctionQ(x).

R
1

mp(x)dxgivesthelifetimeproportionoftimesthat
x(x=m)losestoanothermemberofthe
mix.Noticethataswithnetbene
ts,thefunctionp(x)iscentral.

ExpectedLifetimePayo.

So,togettheexpectedlifetimepayoto
x(x=m)vsersustheequilibrialmix,wesimplysubstitute
theequationsfornetbene
t,(9.8),andcost,(9.10),andobtain

.
m
.

Z
E(
x(x=m);mix)=(v..x)p(x)dx..mQ(m):
(9.11.

0
Chapter9.AMixedESSSolutiontotheWarofAttrition
78

OK,wehavethepayoandcostequations,(9.8)and(9.10),thatcontainthepdffunctionp(x).
Howdoesone
ndthecorrectfunctionp(x)forthewarattrition?Itisnotterriblydicult,butthen
neitherisitcentraltoourstory.Ifyouareinterested,youshouldtakealookattheAppendixa
t
theendofthischapter.Itisabeautifulapplicationofanumberofideasfromcalculus.Butfor
the
moment,we'llproceeddirectlytothenextsectionwherewe'llintroducetheresultthatMayna
rd
Smithobtainedforp(x)andwe'lldiscussitinconsiderabledetail.

9.4TheMathematicsoftheMixedEquilibrium

MaynardSmith'sgoalwasto
ndafunction,p(x),thatwouldsupplythefrequenciesofeach
supportingstrategy(i.e.,cost,x)foranequilibriuminthewarofattrition.Togetp(x)heus
ed
(9.11)andobtainedthefollowingresult,

p(x)=V1
e..x=V;
(9.12.

wherep(x)istheprobabilitydensityfunction(dimensionsofprobabilityperunitcost),xis
cost,V
isresourcevalue,andeisthebaseofthenaturallogarithmfunction3(e2:713).SeeFigure9.6.
NegativeexponentialfuncitonsareanexampleofaveryimportantgroupoffunctionscalledPo
isson
distributions4
.

1
..x=V
Figure9.6:Theprobabilitydensityfunctionp(x)=V
ewhenV=1.Theareaunderthiscurve
from0to1is1.
1

0
.
................................................................................
................................................................................
................................................................................
................................................................................
................................................................................
................................................................................
.................................................
0123
Ifyou'vetakencalculusthefollowingsortofargumentwillbefamiliar;ifnot,justgettheid
ea
thatweapplyappropriateantiderivativeandlimitrulestogettheexpectedresult.Fromcalc
ulus,

e..x=V1
3Rememberthat=x=V.Negativeexponentsarethesamethingasthemultiplicativeinverseofth
e

e
expression.

4Poissondistributionsaremathematicaldescriptionsoflargenumbersofrandomevents.Sta
rtingfromafew
simpleequationsthatdescriberandomevents,theseequationsgeneratepredictionsforthel
argescalepatternsthat
wouldresult.Theseresultingdistributionshaveanumberofdierentshapesthataredetermine
dbythetypeof
processthatisbeingmodeled.

OneexampleofanaturalphenomenonthatcanbemodeledusingaPoissondistributionisradioac
tivedecay.We
knowisthatthereisacertainchancethatanunstablenucleusofacertaintypewillemitenergy
eachmomentintime.
Thus,decaysappeartoberandomeventsthathaveacertainchanceofhappeningeachunitoftime
.Usingatypeof
Poissondistributionknownasanexponentialdecay,whichisinformidenticalto(9.12)(theb
asisforourdescription
ofthebehaviorofthestablemixedstrategyvar),wecaneither:predicttheprobabilitythata
givennucleuswill
decayaftersomestartingtimeor,ifwehaveapopulationofnuclei,wecanpredictthenumberof
decaysthatshould
occurperunittime.

Closertohome,thedistributionofquittingcostsusedbyavarstrategistwillalsohaveanega
tiveexponential
decay.Andthatisbecausejustliketheradioactivenuclei,thereisacertainprobabilityofc
ontinuing(quitting)for
eachincrementofcost.

Poissondistributionsareextremelyimportantinscienceingeneralandinbiologyinparticu
lar.Otherversionsof
thedistribution,forexample,formthebasisfordeterminingwhetherornotpatternsweobser
veinnaturearerandom
ascomparedtogrouped.
Chapter9.AMixedESSSolutiontotheWarofAttrition
79

1..x=V
anantiderivative(readcumulativedistributionfunction5)forp(x)=Veisjust

..x=V
P(x)=1..e:

1..x=V
Nowwecanshowthatthetotalareaunderthedensityfunctionp(x)=Veis1.Firstuse
limitstoexpressthein
nitelylongintervalofintegrationandthenusetheantiderivativeofp,

ZZb
1
b

p(x)dx=limV1
e..x=Vdx=lim1..e..x=V
.
:
b!1
b!1
0
00

Nextdotheevaluatoin,
b

.
1

..x=V
..b=V
0=V
lim1..e
.
=lim1..e..(1..e)=lim1..:

eb=V
b!1
0
b!1
b!1

Nowastheexponentbgetsin
nitelylarge,theentiredenominatorgetsin
nitelylargeforcingthe
fractiontoapproach0.So

lim1..=1..0=1:

eb=V
b!1

Puttingthisalltogether,

Z
Z
Z
1
b
1..x=V
p(x)dx=limVedx=1:

b!1
00

Let'salsoseehowtointegrate(9.12)togetanexpressionthattellusthechancethatanindivi
dual
playsuptoacertaintime.First,let's
ndanexpressionforthetotalproportionofindividualsin
themixwhoareexpectedtohavequitbetweencostsbetweenzeroandcostx=m.Thisofcourseis
thesameasgivingthechancethatamixedstrategistwillquitbycostm.Thisisthecumulative
probabilitydistributionofquittingtimesP(x)thatwesawearlier.Inthiscase,whenx=m,

1..x=V
theprobabilityweseektheareaundertheprobabilitydensitycurvep(x)=Vebetween0and
m,or

ZZ
m
mm
1..x=V..x=V
P(m)=p(x)dx=Vedx=1..e
.

0
00

..m=V
=1..e..(1..1.

..m=V
=1..e:
(9.13.

Toreiterate:Whenweevaluate(9.13)foranyparticularcostx,theresultwillbethetotal
proportionofapopulationofmixedstrategistswhowouldhavequitasofcostx.Thatis,P(m).
1..e..m=Vincludesthosequittingatcostx=mandallthathavequitbeforecostm.

Figure9.7showsplotsofP(x)forthreeresourcevalues(V)overarangeofcostsbetweenx=0
andx=10.Noticethatinallcasesthechanceofhavingquitis(ofcourse)initiallyzero.Ascon
test
costsaccumulate,itbecomesmorelikelythatonewillhavequitsincecostsstarttoexceedthe
maximumdierentsupportingstrategiesarewillingtopay.Note:Wehavetalkedaboutindividua
ls
whoquitatcost0;assumethatwhatreallyhappensisthattheyquitafteranin
nitesimallysmall
cost,0+dx,ispaid).

Anotherwaytothinkabouttheseplotsistoimagine1000identicalmixstrategistsstartinga
displaygame.Attimezero,allareplayingsozerohavequit.Ashorttimelatersomehavequit,a
s

e..x=V..e..x=V
5Thegeneralanitderivativeofp(x)=V1
isjustP(x)=+c,buttoaccountforthefactthat
P(0)=0,wemustsetc=1inthecumulativedistribution.
Chapter9.AMixedESSSolutiontotheWarofAttrition
80

Figure9.7:P(x)forthreeresourcevalues(V)overarangeofcostsbetweenx=0andx=10.

..
..................

..
....
................................................................................
................................................................................
................................................................................
................................................................................
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..
1.0
1.0
.
..............
..............
..............
.............
............
.
...................
.
.
.
...............
.
.
0.8
0.8
.
.............
.
.............
.
.
.
.
.
.
.
..............
.

0.6
0.6
.
.............
.
.
.
.
.......
....
...
.

.
.
..
...
...
...
..
0.4.
.
.
........
...
..
.

0.2.
..
..
....
..
...
.
.
.
.
|V=0:5,--V=1:0,V=2:0
.

0.0
.
..
..
..
..
...
..
..
..
...
..
..
..
..
...
..
..
..
..
...
..
..
..
..
...
..
..
..
...
..
..
..
..
...
..
..
..
..
...
..
..
....
...
....
.....
..
....
..
...
..
..
..
..
..
.......
......................
................................................................................
....
...
....
..
........
..
..
.
.
012345678910
timegoesonagreaterandgreaterproportionhavequitandsotheoverallchancethatanindivid
ual
whostartedthegamewillhavequitgraduallyincreases.

TheotherthingtonoteistheeectofVonquitting.AsVgetslarger,individualsquitatlower
rate(fewerquitperincreaseincostx).Thisshouldmakesense|acontestantshouldbelesslik
ely
togiveupoveravaluableresource.Infact,therateofquittingisproportionalto1=V;moreab
out
thisbelow.

Hopefullythisisallstartingtomakealotofsense.Nowlet'slookattheconverseofthe
cumulativeprobabilityofhavingquitascostx(alternately|thetotalfrequencyofquitters
asof
costx).Theconversewouldbethecumulativeprobabilityofthosewhohavenotquitasofcost
x.Equivalently,thisistheprobabilityofnothavingquit,ortheprobabilityofenduringtoa
certaincostx.WecallthisQ(x)andwesawitearlierin(9.10)fornetcosttoanysupporting
strategyversusmix.

OK,ifP(x)isthecumulativechancethatanindividualwillhavequitassomecost,thenthe
probabilityofenduringuptoacertaincostxis

..m=V..m=V
Q(m)=1..P(m)=1..(1..e)=e;
(9.14.

wherewehaveused(9.13)forP(m).Wecould,ofcourse,
ndQ(m)byintegratingpfrommto
in
nity.(Tryit!)SeeFigure9.8foraplotofQ.

Figure9.8:AgraphofQ(x),theprobabilityofenduring,whenV=1.

1.0
0.8
0.6
0.4
0.2
0.0
.
....
...
...
...
..
...
..
......
....
..
...........
.....
..
......
..
.......
..
....
...
..
..
...
..
...
..
...
..
...
.........................
.......
.......................................
.............
................................................................................
................................................................................
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................................................................................
................................................................................
................................................................................
................................................................................
......................
012345678910
Chapter9.AMixedESSSolutiontotheWarofAttrition
81

Problems

Beforegoinganyfurther,let'sbesurethatyoucancalculatethecumulativeprobabilitydist
ributionequationP
(x).Tosolvethisproblem,youwillneedacalculatororspreadsheetwithnatural
exponentialfunction(exponentiationofe,oftencalledexp)oralogtable.

1.Whatisthecumulativechanceofquittingbetweenacostof0andin
nityifV=1?V=5.
V=0:5.

2.Whatisthecumulativechanceofquittingbetweenacostof0and0:6ififV=1?V=5.
V=0:5.

3.AswithP(x),ifweevaluateQ(x)in(9.14)foraseriesofvaluesofcostswecangetaplotof
thecumulativechanceofenduring(notquit)asofanycostx.ReconsidertheplotforP(x)in
Figure9.6.HowmustthisgraphalteredtoformQ(x).

CumulativeProbabilityoveranInterval

Now,forourlastequationbeforeweuseeverythingwe'velearnedtosummarizethecharacteris
tics
ofourmixedstrategy(i.e.,thestablemix).Noticethat(9.13)and(9.14)bothgivecumulativ
e
probabilities.Thismeansthatbothgivefrequencies/probabilitiesstartingatzerouptoso
mecost
x.Thus,ifthatcostxisin
nite,thenthecumulativechanceofhavingquitbythatcostis1.0and
thecumulativechanceofnothavingquitis0.

Butwhatifwesimplywanttoknowtheprobabilitythatanindividualwillquitoversomespeci
c
costrange|forexample,betweencostx1=0:50000andcostx2=0:50001.Thisisespeciallyusef
ul
inunderstandinghowacomputersolvesthewarofattritionsuchasinthewarofattritionsimul
ation
thataccompaniesthispage.

Allweneedtodoissubtractthecumulativefunction(P(x)orQ(x))valuesfortwodierent
costs.WewillcallthisprobabilityP(m)orP(m1xm2),thatis,theprobabilityof
quittingbetweenthespeci
ccostsm1andm2.

WecanalsogetP(m)bysimplyintegratingbetweenanytwolimits.(Wealreadysawthis
ideain(9.5)..

Zm2

m
m
P(m)=P(m1xm2)=V1
e..x=Vdx
(9.15.
1
m2
..x=V
m2=V
..m1=V..m1=Vm2=V
=..e
.
=..e..(..e)=e..e:

m
m
1

Noticethatusing(9.13)givesthesameexpression,

m2=V..m1=V..m1=V
..m2=V
P(m)=P(m2)..P(m1)=(1..e)..(1..e)=e..e:

So,wehavenowgoneovertheequationsthatcangiveusvariousprobabilitiesorfrequency
distributionsinthewarofattrition.Andmostimportantly,allofthesearethe\children"of
(9.12),
theprobabilitydensityfunctionthatMaynardSmithderivedstartingwith(9.11).Wewilluse
these
functionsinthediscussionsbelow.

Inthenextsection,wewilldiscusswhat(9.12)reallymeans:whatdoesitsayaboutmixed
strategiesinthewarofattrition.Afterwehaveafulldescriptionofthismix,wewillturntoo
ur
naltask|provingthatthemixisanESS.
Chapter9.AMixedESSSolutiontotheWarofAttrition
82

Problems

4.Nametheprobabilitydistributionsthatwesawearlierthatgive
a)theprobabilityofcontinuingtoacertaincost;
b)theprobabilityofquittingasofacertaincost.

5.If(9.15)givesthechanceofcontinuingforaunitofcost,
ndanexpressionthatgivesthechance
ofquittingperunitcost.

9.5ADescriptionoftheMixedEquilibrium

Wearenowatapointwherewecanunderstandthecharacteristicsofthemixedequilibrium.As
mentionedpreviously,thisequilibriumcouldconsistofeither

amixofindividualswhoplayeddierentpurestrategies(singlemaximumcosts)butwherethe
frequencyofeachpurestrategytypewasequilibrial(asultimatelydescribedby(9.12),

apopulationconsistingentirelyofmixedstrategists|thatis,individualswhowerecapable
ofplayinganystrategyinagivencontestsolongastheprobabilityofplayingaparticular
maximumcostwasultimatelygivenbyby(9.12),or
somemixofthetwoabove,includingperhapsalternativeversionsofmixedstrategistssolong
astheoverallfrequencyofeachsupportingstrategyinthepopulationasawholewasinline
with(9.12).
Thus,(9.12)hasakeyroleindescribingtheequilibrium.Inthissectionwewillfocusonthe
characteristicsoftheequilibrium.Howshouldmembersofapopulationatthisequilibriumac
t.

ImportantConvention:Forconveniencewearegoingtothinkaboutourpopulationinterms
ofthesecondpossibilityjustdiscussed|wewillregardtheequilibrialpopulationasconsis
tingentirelyofmixedstrategists
,allofwhomarecapableofplayinganymaximalcostwithaprobability
ultimatelydescribedby(9.12).Sinceothermixesarepossiblewe'llgivethisparticularmix
aname
varforvariablecoststrategist.

ANoteAboutStrategyNames

Someofthisisreiterationofwhatwasjustsaidbutpleaseglanceoveritsothatyouarefamilia
r
withthestrategynamesandde
nitionswewillusefromhereonout.

Thenamesandsymbolswewilluseforthestrategiesareabitdierentthanthoseusedby
MaynardSmith[1974]andBishopandCannings[1978].Theyaremeanttobemoredescriptive
andthereforeeasierforsomeonetoremember;hopefullythisusewillnotresultinanyconfusi
on.
tothosefamiliarwiththeseauthor'swork.Idothiswithsomereluctancebuthavefoundthatmy
studentsseemtohaveaneasiertimethiswayascomparedtousingsymbolssuchasIandJor
mix.So:

Asjustmentioned,we'llcalltheevolutionarilystablemixdiscoveredbyMaynardSmithvar
forvariabledisplaycost.varconsistsofallpossiblecostsplayedatfrequenciesdetermine
dby
theprobabilitydensityfunction,(9.12).varwillbethecenterofmostofourdiscussion.
Thetermmixwillapplytoanymixedstrategy|i.e.,astrategythatconformsto(9.1).
Chapter9.AMixedESSSolutiontotheWarofAttrition
83

Thename
x(x)willapplytoanystrategywhoseplayersselecta
xedmaximumdisplaycost
(time)x.Thus,therearepotentiallyanin
nitenumberofversionsof
x(x),eachcharacterized
bydierentmaximumcostsbutallsharingthecharacteristicthatoveralifetimetheyhave
butonemaximumcost(incontrasttovar).Wehavepreviouslyconsidered
x(x)indetailin
thepreviouschapterandshownwhytheyarenotevolutionarilystable.

Fortherestofourtreatmentofthewarofattrition,wewillregard
x(x)strategistsnot
assupportersofthevarequilibriumbutinsteadascompetitors,i.e.,potentialinvaders.Ju
st
thinkofthemasattemptingtoinvadeapopulationconsistingentirelyofmixedstrategists;t
he
additionofany
x(x)strategistwillhavetheeectofchangingthefrequencyofaparticular
maximumacceptablecost(whichcanbegeneratedbyeitheravarstrategistorthis
x(x.
invader)fromtheequilibrialvaluegivenby(9.12).We'regoingtolearnwhetherornotthis
alterationwillbepermanent.

Thecharacteristicsofthemixedstrategyvar

1.Likeotherstrategies,varishighlysecretive!Therecanbenoinformationtransferfromva
r
toitsopponentthatmightsignalwhenvarwillquit.

Thus,theopponentofavar(variabledisplaycoststrategist)neverknowsnornevercan
knowexactlywhenthevarstrategistwillquit.Nofactor(e.g.,physiologicalcondition
orsomeintentionmovement)canbeallowedthatmighttipotheopponentastovar's
intentions.

Obviously,ifsuchinformationtransferoccurred,itwouldbeeasytocreateastrategy
againstvar(out-waitvarinanycontestuptomV=2,quitatm=V=2).
Thisisoneofthefewimportantcharacteristicsofvarthatisnotsubsumedby(9.12).
Butnotethatitisalsoacharacteristicthatanystrategyshouldpossess.Forinstance,if
a
x(x)strategisttipsitshand,itwouldalsoplaceitatadisadvantage.

2.varstrategistsmaypotentiallyplayanycost|fromnocostto(theoretically)anin
nitecost.
Wediscussedthereasonsforthisintheintroductiontothischapter.

3.varstrategistshaveaconstantrateofcontinuingovereachunitofcost.Thechanceof
continuingisproportionalto1=V.Thisquantityisalsoknownastherateconstant.

ProbabilityofContinuingPerUnitCostx=e..1=V:
(9.16.

ThisisthefunctionQ(x)in(9.14)withx=1.Thus,withregardtothechanceofvar's
continuingofadisplay:

theexponentx=Vofein(9.12),(9.13),(9.14),and(9.15)isnothingmorethana
cost/bene
tratio.Lookingatcostandbene
tsseparatelyisalsoinstructivE:c

{Thelargerx=Vis(thecost/bene
tratio),thesmallerthechanceofcontinuing.
(Thisshouldbeclearbecausetheexponentx=Voccurswithanegativesigninthe
expressione..x=V..

{So,sincethechanceofquittingistheinverseofcontinuing,thelargerx=Vis(the
cost/bene
tratio),thegreaterthechanceofquitting.

Thus:
Chapter9.AMixedESSSolutiontotheWarofAttrition
84

{thechanceofcontinuingisdirectlyproportionaltotheresourceV.Thisshould
makegoodintuitivesense|themorevaluabletheresourcethelesslikelyacontestant
shouldbetoquitinagivenincrementofcost.
{Thechanceofcontinuingisinverselyproportionaltothecostorcostincrement|the
greaterthecost,thelowertheprobabilityofcontinuing.
4.Now,sincethebehaviorofavarstrategistisdeterminedbyacertainchanceofquittingwith
eachunitofcost,andsincevarnevertipsitshand,youshouldrealizethatanopponent
willneverknowexactlywhenavarstrategistwillquit|anymorethanyou,meoranyonecan
alwayscorrectlyguesswhena\fair"coinwillturnup\heads."Thus,knowingwhensomething
willhappenisquitedierentfromknowingthechanceofsomeevent.Thisistheessenceof
theproblemvar'sopponentsface!

5.Anotherresultofaconstantchanceofcontinuingperunitcost(i.e.,aconstantchanceof
quittingpercost)isthatthechanceofacceptinggreatercosts(i.e.,ofplayingfromthestar
t
throughtocostx)decreasesexponentially(foranyvalueofVlessthanin
nity,i.e.,forany
e..x=V1:0).Theeectofthisisthatthereisvirtuallynochancethatavarstrategistwill
bewillingtopayacostthatisverylargecomparedtoV.

Whatthismeansisthateventhoughthechanceofremainingorquittingisalwaysthe
sameforthosewhoarestillplayingthegame,thenumberofplayerswilldropmost
rapidlyatthestartandthenmoregraduallyasthenumberofplayersapproachzero.We
havealreadyseenthisinProblem3intheplotofQ(x)(thechanceofplayingfromthe
starttoaparticularcostx)vsersuscost.(SeealsoFigure9.8..
Asmentionedearlier,wecallthistypeofplotanexponentialdecay.Examplesof
exponentialdecayincludeprobabilitydensityin(9.12),Q(x)in(9.14),andP(x)in
(9.15).
6.So,tosummarize,theopponent:

canhaveageneralideaofwhatavarstrategistwilldoinacontestforagivenresourceif
ithasaknowledgeofthedistributionofvarstrategistswillingtopaydierentmaximum
costs,Q(x).
However,theopponentcanneverconsistentlypredictvar'sactionsinanyparticular
contest.Thatisbecausevar'sactionsatanycostaretotallyindependentofanything
thatitdidinpreviousgames|whetheritcontinuesfromonemomenttothenextis
simplyamatterofaconstantchancefactor.
Thus,varis\predictablyunpredictable."

Thelaststatementisperhapsthemostcrucialinunderstandingthebehaviorofvarstrategist
s.
Centraltoitaretheideasofconstantprobabilityofcontinuingthegameandindependenceof
decisionsfromonemoment(cost)tothenext.Youwillalsoexplorethisingreatdetailwhen
yourunthesimulations.

Problems

6.Comparewhatacontestantseeswhenitconfrontsapopulationconsistingentirelyofvarstr
ate-
gistsascomparedtoapopulationthatisanequilibrialmixofpuresupporting
x(x)strategies.
Wouldthecontestantseeanydierenceinthesetwosituations.
Chapter9.AMixedESSSolutiontotheWarofAttrition
85

7.Howwouldyouexpresstheideaofconstantrateofquittingwithrespecttoapopulationofpur
e
strategistswhotogetherproduceanequilibrium.

8.Whyisitcrucialthatnoinformationastovar'sintentiontocontinueorquitacontestbepas
sed
ontoitsopponent.

9.Howdoyouestimatetheprobabilitythatavarstrategistwillwinacontestofcostx.

10.Howdoyouestimatetheprobabilitythatavarstrategistwillloseacontestofcostx.

11.Howdoyouestimatetheprobabilitythatavarstrategistlosesbypayingacostbetweenxand

x+x.

12.These
nalquestionscallforsolutionstoequationsderivedfrom(9.12),theprobabilitydensity
functionthatdescribesvar.Youwillneedacalculatororspreadsheet.

a)Shouldthechanceofencounteringamemberofthe\stablemix"withaquittingcostbetween
0.60and0.61begreaterorlessthanencounteringanindividualwithaquittingcostbetween
0.60and0.62?Explain.

b)Whatisthechanceofencounteringamemberofthestablemixwithaquittingtimebetween
acostof0.60and0.61ifV=1?V=0:5?Comparetheseanswerswiththenextquestion.

c)Whatisthethechanceofencounteringamemberofthemixwhoquitsbetweenacostof
1.0and1.01ifV=1?V=0:5?Comparetheseanswerswiththelastanswers.Whythe
dierence|thesizeofthecostintervalisthesame.

9.6ProvingthatvarisEvolutionarilyStable

RequirementsofProof

Wenowknowthegeneralcharacteristicsofthemixedstrategywecallvar|therangeofitsmaxim
um
displaycosts,theprobabilityofplayingeachofthesecostsandtheirrelationshipofthesep
robabilities
totheresource,etc.Andweknowthattheequation(9.12)whichdescribesvar'sbehaviorsprun
g
fromtheassumptionthat

E(any
x;var)=E(anymix;var)=E(var;var)=aconstant

Finally,weknowthatBishopandCannings[1978]haveshownthatthisassumptionmustbecorrec
t
foranyESSinthesymmetricalwarofattrition(seeBishop-Canningstheorem).

However,simplyshowingthatthevarstrategyhassomebehaviorconsistentwithbeinganESS
isnotthesamethingasshowingthatitisanESS.Recallthetwogeneralrulesfor
ndingESSswe
learnedaboutearlier.varisanESS(cannotbeinvadedifsucientlycommon)if

Rule1.Commoninteractions:E(var;var)E(
x(x);var).

Rule2.Commoninteractions):a)IfE(
x(x);var)=E(var;var),then(rareinteractions):b.
E(var;
x(x))E(
x(x);
x(x)).

Now,inthecaseofvarweareonlyinterestedinRule2sincewealreadyknowthatRule2
a):E(
x(x);var)=E(var;var)istrue.varisderivedfromthis!AndofcourseRule2isnot
consistentwithRule1.ButjustbecausevarisderivedfromRule2a)doesnotmeanthatitmust
beconsistentwithRule2b).Andifvarversusany
x(x)isnotconsistentwithRule2b),then
varisnotanESS.
Chapter9.AMixedESSSolutiontotheWarofAttrition
86

IfvarwerenotanESS,whatwouldItbe?Ifvarversusany
x(x)isonlyconsistentwith
Rule2a),itisequilibrial.ThisisbecauseifE(var;
x(x))E(
x(x);
x(x))isfalse,thenthe
onlyinterpretationthatisalsoconsistentwithRule2a)isthatE(var;
x(x))=E(
x(x);
x(x)).
So,thecommoninteractionswouldhavethesame
tnessconsequencesoneachparty(noadvantage
toeither)andtherareinteractionswouldalsogivenoadvantagetoeitherstrategy.Notethat
the
payosincommonversusrareinteractionswouldnothavetoequaleachother,theonlyequality
neededisthatcommonareequalforbothasaretherare.Theresultisthatselectioncouldnot
changethestrategyfrequenciesandwewouldsaythatthepopulationwasequilibrial.

So,toshowthatvarisanESSallweneedtodoistoshowthatRule2b)holds,i.e.,
E(mix;
x(x))E(
x(x);
x(x)).

WhatwillfollowisamathematicalproofthatRule2b)is,infact,trueandthereforethatvaris
anESSinthewarofattrition.Onceagain,therewillbeabitofcalculustoenhancetheargument
butanyoneshouldbeabletofollowatleasttheoutlineoftheproof.Asbeforethecalculusisal
l
explained,furthermore,muchofitisverysimilartowhatwehaveseenearlier.And,tomakethe
conceptsclearer,graphswillbepresented.

TheProof

Onceagain,toshowthatvarisanESSrequiresthat:

Rule2b)E(mix;
x(x))E(
x(x);
x(x).

istrue.So,wewillneedto
ndexpressionsforE(var;
x(x))andE(
x(x);
x(x))anddetermine
whetherornotthedierencebetweenthetwoisalwaysapositivenumber,i.e.,

E(mix;
x(x))..E(
x(x);
x(x))0:
(9.17.

Now,recallfrom(9.2)thatthepayotoagivenstrategyinacertaintypeofcontestisalways

E(
x(x=m);mix)=LifetimeNetBene
tstoFocalStrategyinWins
..LifetimeCoststoFocalStrategyinLosses.(9.2.

So,let's
ndthenetbene
tandcostequationsforE(mix;
x(x))andE(
x(x);
x(x))andthen
substitutetheminto(9.2)before
nallysolvingtoseeifwehaveanESS.We'llusethesamegeneral
symbolsandoperationsthatweusedin
ndingE(
x(x);var)earlier.

PartOne:CalculationofNetBene
ts

Bene
tsneededtocalculatethesepayosareeasyto
ndandsotheyrepresentagoodplaceforus
tostart.First,recallthatweassumethatthevalueoftheresourceisconstantinanygivencon
test;
furtherweassumethatithasthesamevaluetobothcontestants.Asusual,wewillsymbolizeita
s
V.Herearethenetbene
tsforeachtypeofinteraction.

Netbene
tstovarincontestsversus
x(x)Rememberthatvardoesnotenteracontest
possessingaparticularmaximumcostthatitiswillingtopay(asdoesa
x(x)strategist).Instead,
ateachinstantithasaconstantprobabilityofquittingproportionalto1=V.Thus,itisunpre
dictable
astoexactlywhenitwillquit.

Recallthatinwarsofattrition,winners,likelosers,paycosts.Thesecostslowerthenet(re
alized.
valueoftheresourcetothewinner.We'llcallthemaximumcostthe
x(x)strategistiswillingto
paym.So,againstagiven
x(x=m)strategist,varwinswheneveritiswillingtopaymore(i.e.,
Chapter9.AMixedESSSolutiontotheWarofAttrition
87

wheneveritcontinuestoplayafter
x(x=m)quits.Thus,whenvarwins,itwillalwayswinV..m.
Butrememberthatitisnotcertainthatvarwillplaytoahigher(winning)costthan
x(x=m.
sincevarusesaprobabilityfunctiontodeterminewhentoquit.So,varexpectstoget

NetBene
t=[(V..m)(Probabilityoffacingx)](9.4.

0x<m

Recallfromearlierthatthechancethatvarhasnotquitasofpayinganycostx=m(i.e.,the
chancethatvarhascontinuedlongenoughsothatthecostitiswillingpay,m,isgreaterthanit
s
opponent)is

Z
Z
1

Q(m)=p(x)dx:
(9.18.

Theequationsaysto
ndthechancethatvarhasnotquitasofcostmbyaddingupallofthe
probabilitiesofvarquittingatcostsgreaterm.Obviously,thissumisthetotalproportiono
ftimes
whenvarhadnotquitasofagivencostm.WesawthisequationforQ(x)earlier.

So,sinceforanycontestthatendsatacostofmthatvarwins,varwillreceiveV..m.And,
varwillwinatafrequencygivenby(9.18).Thus,thebene
ttovaris

Z
Z
1

Bene
t=(V..m)p(x)dx:
(9.19.

NoticethatV..misoutsideoftheintegralbecauseinthecaseofvaragainstagiven
x(x=m),
varcanneverexpecttowinanythingexceptV..m.So,V..misaconstantforacontestthat
canlastuptoagivencostm.Andvaronlywinswhenithasnotquitasofm.varwillofcourse
quitdisplayingwhenitwinsatcostm.Rememberthiswhenyousee(9.23)forthecostvarpays
inlosses|theremayonlybeonewaytowinagainstagiven
x(x=m)butaremanywaysforvar
tolose.Inthatcase,thecostwillremainwithintheintegral.

Substituting(9.18)into(9.19)yields

Bene
t=(V..m)Q(x):
(9.20.

andby(9.14),Q(m)=e..m=V
,so

..x=V
Bene
t=(V..m)e:
(9.21.

Netbene
tsin
x(x)versus
x(x)contestsInthiscontestwehavetwoidentical
x(x.
strategistsfacingeachother.Thus,theyplaytoexactlythesamecostx=m.Sinceweassumeno
otherasymmetries,thenitisbesttoassumethattwoidenticalindividualswilleachwin50%of
the
time|theywillineectsplitthenetbene
ts.Thus,

Bene
tfor
x(x)versus
x(x)=0:5(V..m)=0:5(V..x.
(9.22.

PartTwo:CalculationoftheCostofLosing

Calculationofcosttovarstrategistsinlossesto
x(x)Forcontestsinvolvingvar,the
calculationswillbeabitmorecomplicatedthanthosefornetbene
t.Asmentionedabove,the
reasonisthatvarcanlosetoagiven
x(x=m)manyways!Here'sanexample.

Supposethatavarstrategistrepeatedlyplaysa
x(x=m=1)strategistincontestswhere
V=1.Whathappensintermsofcosts.
Chapter9.AMixedESSSolutiontotheWarofAttrition
88

{Weknowthatvarlosesanytimeitquitsbeforepayingacostslightlygreaterthan1.
{Therearemanywaysthatavarstrategistcanlosetoa
x(x=1)strategistovera
repeatednumberofgamesbecausevarcanplayapotentiallyin
nitenumberoflosing
costs(i.e.,costsbetween0and1)against
x(x=1).
{Thereisadistinctprobabilityassociatedwitheachoftheselosinggambits(costs).

So,overalifetime,thecostthatavarstrategistexpectstopaywhenitselectsalosingcostwi
ll
beequaltothesumoftheproductofeachuniquelosingcosttimestheprobabilityofplaying
thatlosingcost.
Thisideaisexpressedmathematicallyusingintegration:

Z
m

CosttovarLosingto
x(x=m)=xp(x)dx:
(9.23.

Let'sbesureweunderstandwhat(9.23)means:

xisthecostvarpaidasofthemomentofquitting,and
p(x)dxisthechanceofquittingbetweencostxandthenextin
nitesimallysmallincrement
incost.Thus,

theproductofxandp(x)dxistheexpectedlifetimecosttovarofplayingtoaparticularcost
xandthenquitting.

Now,sincetherearemanywaystolose,thereforewemust
sum(integrate)thevaluesexpectedforeachcontestcostxp(x)dxbetweenx=0andx=m,
thecostthe
x(x=m)opponentiswillingtopay.

Thissumisthethelifetimecostvarexpectstopayinlosingcontestswheretheopponentis
willingtopayacertainamountm.
Wecansolve(9.23)byinserting(9.12)forp(x)andintegrating6
,

Z
Z
m
Zm
CosttovarLosingto
x(x=m)=xp(x)dx=xV
1e..x=Vdx=V..e..m=V(V+m):

00

(9.24.

Calculationofcostto
x(x)strategistswhenversus
x(x)Onceagain,thisisaveryeasy
calculation.Thecontestantsareidentical,botharewillingtopaycostx=m.Aswesaidinour
considerationofbene
ts,wesimplyassumethateachindividualwins50%ofthetime.So,halfthe
timetheylostandpaycostx=m

Costpaidby
x(x=m)inlosingto
x(x)=0:5x=0:5m:
(9.25.

6Calculusstudentswillrecognizethatthisisanintegrationbypartsproblem,since

R
1
e..x=V..xe..x=Ve..x=V..xe..x=V
..x=V
x
dx=+
dx=..Ve:
V
Chapter9.AMixedESSSolutiontotheWarofAttrition
89

PartThree:PayoEquations
SectionA:E(
x(x=m);
x(x=m))We'llreversethingsnowandstartwith
x(x)contests
thatendinties(sincethey'reeasy).Now,recallfrom(2.1.
Payo(toStrategy1,whenversusStrategy2)=(Bene
tfromwin)..(Costofloss).(2.1.
Ifwesimplysubstitutetheequationsforbene
tinwinning(9.22)andcostinlosing(9.25)into(2.1.
weobtain
E(
x(x=m);
x(x=m))=0:5V..m:
(9.26.

SectionB:E(var;
x(x=m))Thistimewesubstitute(9.19)and(9.23)into(2.1.
Z1
Zm
E(var;
x(x=m))=Bene
t..Cost=(V..m)p(x)dx..xp(x)dx:(9.27.
m
0
Ifweintegratethisequationweobtainasin(9.21)and(9.24),then
E(var;
x(x=m))=2Ve..m=V
..V:
(9.28.

TheMixedStrategyvarisEvolutionarilyStable

RecallfromabovethattoprovethatvarisevolutionarilystablethatweneedtoshowthatRule2
b)iscorrect.

FindinganequationforthedierenceinpayosStartingwithRule2b),wemustshow
that

E(mix;
x(x=m))E(
x(x=m);
x(x=m)):
orequivalently,wemustshowthat

E(var;
x(x=m))..E(
x(x=m);
x(x=m))0:

Now

E(
x(x=m);
x(x=m))=0:5V..m
andfrom(9.28.

..m=V
E(var;
x(x=m))=2Ve..V:
Sowemustshow

..m=V
E(var;
x(x=m))..E(
x(x=m);
x(x=m))=2Ve..V..0:5V..m0:

Thissimpli
estoshowingthat

..m=V
E(var;
x(x=m))..E(
x(x=m);
x(x=m))=2Ve..1:5V+m0:(9.29.

Nowthebigquestion|is(9.29)alwayspositiveasitmustbeifvarisanESS?Wecouldstartout
bysimplygraphingit.IfwedosoforV=1wewillseethatthereisnoplacewhereE(var;
x(x)),
E(
x(x);
x(x))0.SeeFigure9.9.

Thus,itwouldappearthatvarisstable.Butnotsofast|thisisforonlyonevalueofV.Isit
possiblethattherearevaluesofVwherevarisnotevolutionarilystable?Afterall,Vdoesaect
var'sbehavior.

Aswith
ndingthefrequencyofeachmaximumacceptablecost(whenwelookedforp(x)),
solvingforeverypossibleVmightappeartobeadicultproblem(andapproachedthatway,it
is!).However,onceagainabitofelementarycalculuscancometoouraidandcomfort.
Chapter9.AMixedESSSolutiontotheWarofAttrition
90

Figure9.9:ThegraphofE(var;
x(x))..E(
x(x);
x(x))isalwayspositivewhenV=1.(Looks
likethe\swoosh"doesn'tit!.

4
3
2
1
0

................................................................................
................................................................................
................................................................................
................................................................................
................................................................................
.........................................................................
012345
CostmpaidbyWinner(unitsofV)

MathematicalProof

Toshowthatnopointon(9.29)islessthanorequaltozero,weneedto
ndtheminimumvalueof
atpartofthegraphabove;onthat
(9.29).Thisoccurswheretheslopeofthegraphiszero(the
graphithappensatavaluesomewherenearcostm=0:7).

To
ndthispointforanyV,weusethecalculustechniqueofdierentiation.Itwillgiveus
anequationfortheslopeateverypointofaplotof(9.29).

Ifwethensolvethis\equationofslopes"forthecostmwheretheslopeequalszerowe
nd
thatthisalwaysoccursatVln20:693V.

Now,allthatremainstodoistosubstitutethevalue0:693Vbackinto(9.29)andsolvefor
E(var;
x(x=m))..E(
x(x=m);
x(x=m)).Theresultisthethatminimumdierence
isalways(ln2..0:5)V0:193V.

Thus,varisanESS!

GraphicalIllustrationoftheProof

Ifyouarenotfullycon
dentthatyouunderstandtheproof,youwillprobablybereassuredifyou
lookatthegraphsbelowof(9.29)fordierentvaluesofV.Remember,theminimumdierencein
tnesswillalwaysbeapproximately0:193Vandwillalwaysoccuratcostm0:693V.Thus,asV
getslargertheminimumdierencebetweenthetwopayosincreases.

Foranycostmpaidbythewinner,E(var;
x(x))..E(
x(x=m);
x(x=m))>0.Consequently
,E(var;var)=E(
x(x);var)andvarisevolutionarilystableagainstany
x(x)!

Problems

13.Writeanexpressionforthelifetimecosttoavarstrategistofquittingatacostofexactly
x.
Chapter9.AMixedESSSolutiontotheWarofAttrition
91

Figure9.10:Theevolutionofapopulation.

14.Writeanexpressionforthelifetimecosttoavarstrategistforlosingcontestswherethew
inner
waswillingtopaym.

15.WhatisE(var;
x(0))inthecaseofatie.

ThingstoRememberAboutthevarStrategy

Perhapsthemoststrikingthingaboutthevarstrategyisthatitsopponentnevercanknowwhen

itwillquit.Wehaveseenthattheoverallpatternofquittingisdescribedbyanexponentialde
cay

typeofPoissondistributionwitharateconstantequalto1=V.Thus,anopponentcouldlearn7i
n

generaltermswhatitsvaropponentwoulddo.Itcould\know"thatitwasmostlikelytoquitearl
y

inacontestandthatthechanceofquittingperunitofcontestdisplaycostise..1=V
.Fromthis,it

ispossibletocalculate(orlearnfromexperience)theexpectedoutcomeofcontestsofvariou
scosts.

Evenifitknewthesethings,itcouldneverknowwhetherornotvarreallywouldquitwiththenex
t

incrementofcost.Thus,noamountofexperiencewithvarstrategistswillallowanopponentan
y

edgeoverit.

Theotherthingtoreiterateaboutvaristhatthereisalogictoitsquitting.Itistiedtothe

resourcevalue|thegreaterthatvalue,thelesslikelythatvarwillquitatanyparticularcos
tandas

aconsequenceitispotentiallywillingtoacceptahighercostcontest.Also,sincevaralways
quits

mostfrequentlyearlyincontests,thechancethatitwillpaylargecostsrelativetoaresourc
evalue

arelow.

Problem
7Iusethetermlearnloosely|itcouldmean\learn"intheusualsenseoflearningandmemoryori
tmaybethat
wearesimplytalkingaboutmakinganappropriateevolutionaryresponse|selectionforrespo
nsesthatworkagainst
a
xedwaittime.Ineithercase,anappropriateresponsearisestoaparticular
xedstrategy.
Chapter9.AMixedESSSolutiontotheWarofAttrition
92

16.\AreYouFeelingLucky,Punk?"IntheclassicClintEastwoodthriller,DirtyHarry,theEas
twood
characterasksanaer-do-welltopredictthefutureandguesswhetherornottherebulletsleft
inEastwood'sgun.Sowhatdoyouthink?Areyoufeelinglucky?Thechanceofgettingkilled
inascheduledcommercialairlinecrashisroughlyontheorderofoneinseveralmillion.About
thesamechancetheearthhasofbeinghitbyalargemeteor,smallasteroid,orcomet.Discuss
ightattendantorpilot)for
iescommercialairlinesdaily(e.g.,a
whetherornotsomeonewho
ighttobeinafatalaccident.Likewise,theearthhas
yearsismorelikelyonherorhisnext
notbeenhitbyareallybigoneforabout65millionyears.Arewemorelikelytobehitnow
thanweweresay60millionyearsago(5millionafterthelastone).Areyoumorelikelytowin
onyournextlotteryentry(taxonstupidity)ifyouhaven'twoninthepastandlesslikelyifyou
havewon?Whatdoesallofthishavetodowiththewarofattrition.

TestingtoseeifAnimalsareUsingavar-likeStrategy

Thereareanumberoffamousexamplesofanimalsthatappeartobeplayingsimplewaitinggames.
Wewillnotgointothemherebecausetheyarewellpresentedbothintheliteratureandinjust
y,Scatophagastercoraria,
abouteveryanimalbehaviortextbook.Perhapstheclassicisthedung
studiedheavybyParker[1970a,b]andParkerandThompson[1980].Theinterestedreaderisurg
ed
toconsultthesepapersoranynumberofbehavioralecologytexts.Wewill
nishthispage,however,
withthefollowingquestion(whichwasaddressedbyParkerandThompson).

Problem

17.Supposethatsomeonedemonstratedthatanimalwaitingtimescorrespondedtothosepredic
ted
by(9.14).DoesthatconstitutesucientproofthatamixedESSdescribedby(9.14)exists.
Explain.
Chapter10

References

Bishop,D.T.andC.Cannings.1978.Ageneralizedwarofattrition.J.Theor.Biol.70:85{125.

Forrest,T.G.andD.M.Green.1991.Sexualselectionandfemalechoiceinmolecrickets(Scapt
eriscus
:Gryllotalpidae):modelingtheeectsofintensityandmalespacing.Bioacoustics3:93.

Gould,S.J.1990.WonderfulLife:TheBurgessShaleandtheNatureofHistory.W.W.Norton.
NewYork.(Afascinatingbookdealingwiththemeaningofearlyanimalfossilsfromthe
BurgessShales;whilesomeofGould'sinterpretationsmaybeo,thisbooklaysouthisnotions
ofcontingencyandhistoricalaccidentclearly.Thesethemesarealsocontinuallyfoundinhi
s
monthlyarticlesforNaturalHistorymagazine).

Hamilton,W.D.1967.ExtraordinarySexRatios.Science156:477{488.

Lewontin,RC.1961.Evolutionandthetheoryofgames.J.Theor.Biol.1:32{403.

Luce,R.D.andH.Raa.1957.GamesandDecisions.Wiley,NewYork.

icts.J.Theor.Biol.47:209{221.
MaynardSmith,J.1974.Thetheoryofgamesandanimalcon

.1982.EvolutionandtheTheoryofGames.CambridgeUniv.Press.

ict.Nature246:16{18.
MaynardSmith,J.andG.R.Price.1973.Thelogicofanimalcon

Mayr,Ernst.1954.Changesingeneticenvironmentandevolution.InHuxley,J.,A.C.Hardyand
E.B.Ford(eds.).EvolutionasaProcess.AllenandUnwin.pp157{180.(Dealswithfounder
eect..

.1982.TheGrowthofBiologicalThought.BelknapPress.(Chapter2isrelevanttothe
discussiononmethodology,buttheentiremassiveworkiswonderfullythought-provokingand
anyseriousstudentofbiologyshouldmakethetimetoreadandponderit..

Parker,G.A.1970a.ThereproductivebehaviourandthenatureofsexualselectioninScatopha
ga
stercorariaL.(Diptera:Scatophagidae).II.Thefertilizationrateandthespatialandtemp
oral
relationshipsofeachsexaroundthesiteofmatingandoviposition.J.Anim.Ecol.39:205{228
.
.1970b.ThereproductivebehaviourandthenatureofsexualselectioninScatophaga
stercorariaL.(Diptera:Scatophagidae).IV.Epigamicrecognitionandandcompetitionbetw
een
malesforthepossessionoffemales.Behaviour37:113{39.

Parker,G.A.andE.A.Thompson.1980.Dung
ystruggles:atestofthewarofattrition.Behav.
Ecol.Sociobiol.7:37{44.
Chapter10.References
94

Popper,K.1972ObjectiveKnowledge.CambridgeUniv.Press.

Prestwich,K.N.1994.Energyandconstraintstoacousticcommunicationininsectsandanuran
s.
AmZool.94(6),625{643.Downloadthismanuscriptat:

http://science.holycross.edu/departments/biology/kprestwi/pubs/index.html

_
Riechert,S.E.andP.Hammerstein.1983.Gametheoryintheecologicalcontext.AnnRevEcol.

Syst.14:377{409.
Ryan,M.J.1985.TheTungaraFrog.Univ.ChicagoPress.(Chapters7and8.
Slobodkin,L.B.andA.Rapoport.1974.Anoptimalstrategyofevolution.Q.Rev.Biol.49:
181{200.

Wright,S.1931.EvolutioninMendelianPopulations.Genetics16:97{158.

VonNeumann,J.andO.Morgenstern.1953.TheoryofGamesandEconomicBehavior.Princeton
Univ.Press.
Chapter11

Glossary

Adaptation.Anaspectofthephenotype,whetherbehavioralormorphologicalthatisheritabl
e
andthatconfersareproductiveadvantageonitspossessor(s)ascomparedtosomealternative
trait(s).Thus,theampli
cationofthegenespartiallyresponsibleforthesetraitsisdueto
Darwinianevolution.Traitsthatarerelativelydisadvantageousaretermednon-adaptive
ormaladaptive.Notethatthetermnon-adaptiveisalsooftenusedfortraitsthathaveno
signi
cantadvantageordisadvantagewithrespecttoeachotherasoccursinWrightianevolution
(geneticdrift).Perhapsabetteradjectiveforthesetraitsisneutrallyadaptive.

Asymmetry.Wheretherearedierencesinthecompetitivenessoftheplayers.Thus,theoutcome
ofacontestisnotsimplyamatterofchance.Asymmetriescanbeduetomanyfactors,for
instancedierentability,experience,motivation,presentownership,and/orcondition.

Boundary.Inthemathematicalsenseusedinanumberofmodelsinthistext,aboundaryisa
frequencyaboveorbelowwhichselectionforceschangefromfavoringsometraittofavoringan
alternative.

Contest.Thegametheorytermforthecompetitiveinteractionthatoccurswhentwoormore
individualsattempttoobtainthesameresourceitem.

Continuousvariable.Avariablewiththepropertythatbetweenanytwovalues,therearean
in
nitenumberofothervalues.Synonym|analogvariable.

Cooperativecommunication:.Whereactionbythereceiverofasignalincreasesthe
tnessof
boththesignalerandreceiver.Alsoseehonestcommunication.

Currency.Unitsthateitherdirectlyorindirectlymeasure
tness,examplesaregrandchildren,
ospring,eggs,energy,time,chanceofdeath,etc.Inbothgamesandoptimalitymodels,bene
ts
andcostsmustbestatedinacommoncurrencyandthatcurrencymustusedthroughout
themodel.Attemptingto
ndthecorrectcurrencyisoneofthemostimportantaspectsof
modeling.

Cumulativeprobabilitydistribution.Examples:Q(x)orP(x).Thecumulativechanceofsome
eventwithrespecttotheindependentvariable.Forexample,iftheindependentvariableisco
st,
thenP(x)mightindicatethetotalproportionoftimes(orindividuals)whohavequitasofa
certaincost.Thus,thiswouldvarybetweenzero(noonehasyetquit)and1.0(everyonehas
quit).

Q(x).Forourpurposes,theprobabilitythatamixstrategistsuchasvarwillpayaparticularco
st
xandthenquit.Itcanbecomputedbydeterminingtheprobabilityofpayingcostx(Q(x))and
costx..x(wherexissomesmallincrementofx)usingthecumulativeprobabilityfunction.
ThedierenceQ(x..x)..Q(x)isQ(x)andisequaltothechancethatanindividualpaid
Chapter11.Glossary
96

costxandquit.

Discretevariable.Avariablethatcanonlypossesscertainexactvalues|intermediatevalue
sare
notpossibleandareroundedtothenearestpermittedvalueorinrarecasesaresimplyignored.
Synonym:\digital"value.

Dishonestcommunication.Whereactionbythereceiverofasignaldecreasesthereceivers
tness
butincreasesthesender's
tness.Pleasenotethatdishonestisusedasashorthand;noethical
ormoraldimensionsarenecessarilypartofthisde
nition.Synonyms:deceit,non-cooperative.

Display.Abehaviorthathasbeenmodi
edtoserveasaformofcommunication;usuallynodirect
physicalcontactisinvolvedandifthereissuchcontactitishighlycontrolledandisshortof
ghting.
e.Thebaseoftheofnaturallogarithmfunction,approximatelyequalto2.7183.
EquilibrialStrategy.Inourcontext,astrategythatisNOTincreasingrelativetoanyothera
s
aresultofselection.Thus,allequilibrialstrategieswithinapopulationmusthavethesame
tness.Equilibrialstrategiesmayormaynotbeevolutionarilystable.
Equilibrium.Inevolution,constancy,stasis.Apopulationisinequilibriumwhenthereisno
changeinthefrequencyofoccurrenceofcompetingphenotypes(e.g.,behavioralstrategies)
or
allelesinapopulation,measuredovergenerationaltime.

ictmovestoincreasinglydirectconfrontationandperhaps
Escalate.Whenacon
ghts.Anoften
ictoccursinreddeer|twowell-matchedmalesinitially
citedexampleofanescalatedcon
displayacousticallybyroaring,thismaybefollowedbyvisualdisplaystypi
edbyparallel
walksandmay
nallyescalateinto
ghtswithantlers.

Evolutionarystablestrategy(ESS).Astrategythatcannotbedisplacedbyanyotherknown
strategy.Putanotherway,whenitsfrequencyis1.0otherstrategiescannotenterthepopulat
ion
andinsituationswhereitappearsasamutant,itincreasesto
xation.Itisastaticcondition.
ESS'scancomeintwogeneraltypes|pureandmixed.

exp(x).Thenaturalexponentialfunction:exp(x)=ex
,wheree2:7183.

Exponentialdecaydistribution.Adistributionthatresultsfromapplyingaconstantrateof
continuingtothemembersofsomeinitialpopulation.So,foreachvalueofx,thechancethat
anindividualwillchosetocontinueisconstant.Now,sincenotallcontinuefromonevalueof
xtothenext(thosethatdon'tcontinuequitandarenolongerinthepopulationofplayers),
thepopulationdecreaseswithx.Sincethepopulationisdecreasingandsinceeachmemberof
italwayshasthesamechanceofcontinuing,thegreatestnumberquitat
rst,withfewerat
eachsubsequentstep.SeeFigure9.8.

Fitness.Thesumofdirectandindirect
tness.Forourpurposes,itislargelydirect
tnesswhich
isusuallyde
nedasthenumberofgrandchildren.However,manyotherde
nitionsorstand-ins
for
tnessareoftenused{forexample,fecundity,numberofmates,numberofeggs,territory
size,territoryquality,etc.

Fixed.Apopulationgeneticsterm|astrategyandthe\gene"thatcausesitaresaidtobe
xed
whentheirfrequenciesare1.0(i.e.,noalternativesexistinthepopulation).
x(x).A
xedcost(or
xeddisplaytime)strategistequivalenttothe
xedcoststrategiesMaynard
Smith[1982]termeda,b,c,:::,etc.Inthenotationweareusingherexreferstothecostthat
xiswillingtopay(itsvalueofm).Thus,xcanhaveanyvaluebutwhencontestsarewritten
as
x(x)versus
x(x)assumethatxhasthesamevalueforeachcontestant(i.e.,eachhasthe
samem).

Frequency-dependence.Whentherelative
tnessofsomephenotypictraitsuchasabehavioral
strategydependsonhowcommonlycertaineventsorinteractionsoccur.Inthecontextof
games,theseeventswouldbecertaintypesofcontests.Imaginethatananimalwithaparticula
r
behavioralstrategy(focalanimal)canexperiencetwoparticulartypeofinteractions.Furt
her,
Chapter11.Glossary
97

imaginethatoneoftheseinteractionsisbene
cialtofocalanimalwhiletheotherisdetrimental.
Clearlythe
tnessofthefocalindividualwilldependontherelativefrequenciesofeachtypeof
interaction|thisisfrequencydependence.

Game.Aseriesofcontestsbetweenallstrategiesinproportiontotheirfrequency.Thesummed
outcomeofthesecontestsdeterminestheoutcomeofagame;agamecouldbeviewedasthe
interactionsthatoccuroveronegenerationalthoughotherde
nitionsarecertainlypossible.

Honestcommunication.Ashorthandforcooperativecommunication,pleasenotethatintent
andmoralityarenotimplied.

m.Somemaximumcostthatacontestantiswillingtopay.Thus,itisaspeci
cvalueofx(cost.
ort(time).Onmanyoccasionsmisusedsimplytosymbolizethemaximumcostthatsome
arbitraryfocalindividualinagivencontestwillaccept.Thatisthede
nitionweuseonthis
page|mostcommonlyasamaximumcostthatapurestrategyiswillingtopayascomparedto
whatavariablecost(mix)strategistmightpay.Itisusedinconsistentlyintheliterature|o
ften
misthecostpaidattheterminationofacontest.Thus,sinceconteststerminatewhenthe
costexceedswhatoneplayeriswillingtopaybyatinyamount(dx),thenthiscost(which
representsthemaximumacceptablecosttotheloser)becomesm.Sorryfortheconfusion.

MixedESS.AnESSwhereindividualseither(a)playdierentstrategiesa
xedportionofthe
time(e.g.,Hawk60%andDove40%|termedamixedstrategy)suchthatnoothermixwould
beanymoresuccessfulorwhere(b)certainportionsofindividualsplayonestrategyallthe
time(e.g.,alwaysHawkorDove)suchthatthe
tnessofpractitionersofeachstrategyisequal.
Alsoseepurestrategy.

Mixedstrategy.Whenanindividualplaystwoormorebehavioralstrategies,usuallyasamatte
r
ofprobability.Thus,selectingarandomamountoftimetodisplayisanexampleofamixed
strategy.Bycontrast,apurestrategyinvolvesselectingaparticularstrategy,forexample
,
always\displayfortimet."

Optimality.Anoptimalbehaviorisonethatmaximizesthedierencebetweenbene
tandcostin
somecommoncurrency|forinstance
tness,energy,time,rate,etc.Optimalitytheoryisused
topredictthebestwaytoperformabehaviorforagivensetofenvironmentalandphysiological
conditions,itmakespredictionsthatareindependentofthebehaviorsbeingusedbyother
individuals.

Pairwisecontests.MaynardSmith[1982]wroteaboutpairwisecontestsasgameswheretwo
individualsfaceoagainsteachotheroversomeresource.Theoutcomesofthesecontests,if
anindividualengagesinmorethanone,haveadditiveeectsontheindividuals
tness.The
gamesconsideredinthistextinvolvingsequentialinteractionsofHawk,DoveandBourgeoisa
re
pairwisecontests.TherulesfordeterminingwhetherornotanESSexistsinpairwisecontests
aresomewhatdierentthanthoseofaanothermodelofinteraction\playingthe
eld."

Payo.Thenetbene
tofpayoofsingletypeofcontestorinteraction.Inthehonestversus
dishonestcommunication,thepayostothereceiverwereeitherBorCwhileintheHawk
versusDoveeachstrategy(DoveorHawk)hadtwopossiblepayos(onewhenplayingagainst
thesamestrategy,e.g.,DversusD,andtheotheragainsttheoppositestrategy,e.g.,Dversus
H.Multi-strategygameshaveevenmorepayos,theirnumberdependingonthenumberof
strategiesbeingplayed.Inthecaseofwaitinggamesthepayodependsonthetimespent
waitingandwhatothersdo.

Player.Ingamestheory,anindividualengagedinacontest,sometimesusedbroadlyasasynony
m
forastrategyinacontest.

Playingthe
eld.MaynardSmithusedthistermtodescribesituationswhereanindividualis
notengagedatacertainmomentinacontestwithjustoneotherindividual(whoemploysa
certainstrategy)asinpairwisecontests,butinsteadwithmanyindividuals.Agoodexample,
Chapter11.Glossary
98

MaynardSmithpointsout,isaplantthatcompetesnotusuallywithoneotherplantbutwith
manyneighbors,simultaneously.TherulesfordiscoveringwhetherornotthereisapureESSfo
r
anexampleofthistypeofcontestaresomewhatdierentthanforpairwisecontests.Thistext
doesnotconsiderplayingthe
eldmodels,theinterestedreaderisurgedtoconsultMaynard
Smith[1982]asastartingpoint.

Pseudorandomnumber.Theresultofa\random"numbergenerationbyacomputer.Thecomputerus
esanalgorithmtogenerateanumbers
;numbersgeneratedbythismeans
tmodelsfor
randomlydistributednumbers.However,sinceade
nedsetofmathematicaloperationsproduce
these\random"numbers,theyarenotrandominthetruestsenseoftheterm.Manymathematician
sandcomputerscientistshavepointedoutthattherearesubtledi
erencesbetween
numbersgeneratedbycomputeralgorithmsandthosegeneratedby,forinstance,observing
motionofmolecules(orevenmixingballsinalotterymachine!).However,forourpurposes,
pseudorandomnumbersarejust
ne|wewillnevernoticethedierence.Thetermisused
simplytoremindyouthatcomputergeneratedrandomnumbersarenottrulyrandom!

PureESS.AnESSwhereonestrategyis
xedandallknownalternativesareunabletoinvade
sincetheyhavelower
tnesses(seemixedESS).

p(x).Theprobabilitydensityfunctionofcost(x).Thefunctionthatcanbeusedto
ndthe
probabilityofanysupportingstrategyinthemix;
ndingthisfunctionwasMaynardSmith's
maintaskindescribingamixedESStothesymmetricalwarofattrition.ImportantNote:this
functiongivestheprobabilityperunitcostandmustnotbeconfusedwithafunctionthatgives
probabilityperse.Inthewarofattrition,aprobabilitydensityfunctionisusedasacentral
elementofthedescriptionofavariablecoststrategist.

P(x).Thecumulativeprobabilitydistributionfunctionofcost(x).Thisgivesthecumulativ
e
probabilityofsomeevent(forexample,quittingdisplay)asafunctionofsomeindependent
variable,inthiscase,cost(x).Itiscalculatedastheintegraloftheprobabilitydensityfu
nction.
WeuseP(x)toindicatethecumulativeproportionofapopulationwhohavequitasofsome
costx.

Q(x).Thecumulativeproportionofindividualswhohavenotquit(arecontinuinginthecontes
t.
asofsomecostx.

Rateconstant.Aconstantintheexponentofanequationofexponentialdecaythatdetermines
thehowfastthedependentvariable(forexample,chanceofquitting)changeswithrespectto
theindependentvariable(forexamplecost).Forthevarstrategist,therateconstantis1=Vs
o
thelargerthevalueofthecontestedresource(V),thesmallertherateconstantandtherefore
thelesstheindependentvariable(e.g.,probabilityofquitting)changesperunittime.Thus
,for

..rateconstantx
cumulativeprobabilitydistributionP(x)=1..eandsinceinthiscasetherate

..x=V
constantis1=VthenP(x)=1..e.

Resource.Anyenvironmentalfeature(bioticorabiotic)ofimportancetoanorganism's
tness.
Examplesincludefood,nestingsites,shelter,mates,symbionts,orspeci
cplacesintheenvironmentthatarefavorablephysiologicallyorforbehavioralreasons
.Contestsarewagedover
resources.

Satellite.Usuallyusedindiscussionsofsexualselectioninregardstoadvertisementbehav
iors
(generallybymales).Theclassicalexampleisfromacousticsignalingwheresatellitesarei
ndividualswhoremainsilentbuttakeupapositions
(usuallyhidden)nearanactivelyadvertising
individuals.Theyattempttointerceptfemalesthatapproachthecaller.Thus,theydonotpay
aslargecostsasdotheadvertisers.Satellitingmaybeanevolutionarystablestrategy(wher
eat
somefrequencyitproducesthesamelifetimereproductivesuccessasalternativestrategies
such
asadvertisement)orasimplecontingentbehaviorinducedby,forexample,poorphysiologica
l
state.
Chapter11.Glossary
99

Stasis.Equilibrium,nogenerationalchangeinallele(orthephenotypedeterminedbytheall
eles.
frequency.

Strategy.Abehaviororsetofbehaviorsusedbyanindividualtodealwithanimportantlifehis
toryproblem
(forexample
ndingamate,rearingyoung,obtainingfood,etc.).Aswithother
de
nitions,thehumantermstrategythatimpliesconsciousthoughtisusedasashorthand;no
consciousplanningisrequired,eventhoughitmightappearthatthebehaviorsarerationalan
d
plannedinthehumansense.Theuseoftheword\strategy"issimplyashorthandthatexpresses
theappearanceoftheresultofsomebehaviors.Itisgenerallyassumedthatinmostspecies
strategiesarelargelyinnate,areproducedbytheusualgeneticanddevelopmentalmechanism
s,
andareactedonbynaturalselection.However,strategiescanalsobelearned,eveninrelativ
ely
simpleanimals.

Supportingstrategy.Anypurestrategy(uniquecostinthewarofattrition)thatisamember
ofthemixedESS.Alternatively,itisanyuniquecost(inthewarofattrition)thatamixed
strategistplays.Agoodsynonymiscomponent(ofthemix)strategy.Forexample,inthe
HawksandDovesgame,ifinjurycostisgreaterthanV,amixwithsupporting(component.
strategiesHawkandDoveresults.

Symmetry.Equalitywithrespecttocompetitiveabilityasde
nedinaparticulartypeofcontest.
Anunlikelysituation.Inmostofthemodelsweconsider,weassumesymmetryasasimpli
cation
.Ifcontestantsaretrulyofequalability,weassumethateachhasa50chanceofwinning
ictwithnoresorttofurtherescalation.Inrealsituations,thecloserthecompetitive
thecon
ictwilloccur.
abilitiesoftwocontestants,themorelikelythatahighlyescalatedcon

t(time).Acostmeasuredintermsoftimespentdisplaying.Whenthesymboltisused,itis
meanttorefertoauniverseofpossiblevaluesofdisplaytimes.Agivent(e.g.,t1)referstoa
speci
ctime.Ausefulmetricsincedisplaytimeiseasytomeasureandunderstandandsince
tnesscosts(x)areusuallyasimplefunctionoftime.

x(cost).Anydisplaycostinsomesortofunitsthatcanbeconvertedto
tness.Normallyused
interchangeablywithtimeofdisplay(t)sincex=f(t),wheref(t)canbeanyfunctionthat
convertstimetocost.Wealwaysassumethatcostisalinearfunctionoftime(x=mt+bwhere
mistheslopeandbthey-intercept)butthereisnoreasontoassumethatthiswillalwaysbe
so.

V.Thevalueofthecontestedresource;itsreciprocalequalstherateconstantintheprobabil
ity
densityfunctionandcumulativeprobabilitydistributionsforvar.

var.Avariablecost(variabledisplaytime)strategistequivalenttothemixedstrategyMayn
ard
Smith[1982]termedI.Itiscomposedofallpossiblecosts(equivalentofallpossible
xedcost
strategies)eachplayedwithfrequencydeterminedultimatelybyaprobabilitydensityfunct
ion.

Zero-sumgame.Whenthereisa
niteresourcethatdierentstrategiescompetefor;itisdivided
betweenallcompetitorsaccordingtotheircompetitiveability.Whiletherecertainlyarema
ny
examplesofwhatareessentiallyzerosumsituations,therearealsocaseswhereonealternati
ve
behaviorallowsitspossessorstoexploitaresourcenotpreviouslyavailable(i.e.,notavai
lable
toalternativestrategies)inwhichcaseitisanon-zerosumgame.
Chapter12

Appendix:DiscussionandSelected
Answers

12.1AnswersforChapter2

1.Recallthat
tnessisarelativemeasure|thesegameshavetodowithcompetingstrategies
andsoitmakesnosensetoevenconsiderasituationwherebotharenotatleastpotentially
present.Rememberthat
tnessisarelativemeasurebecauseoverthelongrun,individuals
(strategies,genes)thatleavemoreospring(copies,whatever)ofthemselvescometodominate
thepopulation.Ifeveryonehasthesamestrategy,then,withrespecttotheevolutionofthis
strategy,theyenjoyequal
tnessbene
tsordecrementsandsothereisnoevolutionwithrespect
tothisstrategy.Thus,the
tnessescalculatedfromthepayomatrixandfrequencyofdierent
strategiesonlyhavemeaninginthecontextofcompetition.Ifyouhaveproblemswiththis,
reviewthesectionsof
tnessandcompetition.
Oneadditionalpointhowever.Itwouldbecorrecttoassumethatthesmallerthe
tness
valuecalculatedbyeitheroftheseequations,thesmallerthenumberofospringtothatstrateg
y.
Likewise,ifthe
tnesscalculationyieldedanegativenumber,thatwouldmeanthatthestrategy
wouldbedecliningfromonegenerationtothenext.Whilenegative
tnessescouldnotgoon
inde
nitely,ifonestrategy's
tnesswaslessnegativethantheother,itwouldincreaserelative
totheothereventhoughoverall,numbersaredropping!

2.ThepayotoanindividualplayingBagainstoneplayingstrategyA.

3.EachindividualplaysA80%ofthetimeandB20%ofthetime.

4.Apurestrategyisasetofbehaviorsthatanindividualwillemployinagivensetofcircumsta
nces.
ApureESSisasinglestrategythatcannotbeinvadedbyanyotherknownstrategy.
Amixedstrategyisonecomposedofseveralpurestrategycomponents.MaynardSmith
[1982]statesthatthereisarandomcomponentintheorganism'sbehavior(intermsofwhich
behavioralcomponentitwillemployinagivensituation).Wesawexamplesofmixedstrategies
intwo-strategygamesthathadnopureESSandinthe\warofattrition."Bycontrast,amixed
ESSinvolvesmorethanonebehaviormakingupanequilibrium.Thiscouldamixedstrategyor
anequilibriumbetweenindividualsofdierentstrategies.
Chapter12.Appendix:DiscussionandSelectedAnswers
101

5.a)ThefrequencyofstrategyBisB=1:0..a=1:0..0:9999=0:0001.
b)ThefrequencyofAversusAinteractionsintheentirepopulatonisa2=0:99992=0:9998.
FromthepointofviewofanAstrategist,thefrequencyisa=0:9999.
c)ThefrequencyofBversusBinteractionsisb2=0:00012=0:00000001.Fromthepointof

viewofaBstrategist,thefrequencyisb=0:0001.
d)ThefrequencyofAversusBinteractionsE(A;B)isab=0:99990:0001=0:0000999.
e)ThefrequencyofBversusAinteractionsisthesame:ba=0:00010:9999=0:0000999.

f)Fromthelasttwocalculationsabove,youcanseethatthetotalfrequencyofthosepayos
intheentirepopulationisequal.Butthatisnotwhatmatterswhenconsideringwhetheror
notAorBarepurestrategies.Weneedtoknowhowcommoneachparticularinteraction
is.Andthatissimplygivenbythefrequencyofthestrategywithwhichthefocalstrategy
interacts.

OK,let'sseewhatthismeans:ForpayostoA:99.99%ofthemwillbewithotherA
strategistsand0.01%willbewithBstrategists.Thus,9999tiemsmoreinteractionswill
occuragainstA;theBinteractionswouldnotseemtobeveryimportant.Forpayosto
B:Onceagain,99.99%ofthemwillbewithotherAstrategistsand0.01%willbewithB
strategists.

Thus,theimportantpayosforcalculatingthe
tnessofAandBrespectivelywhen
BisrareareE(A;A)andE(B;A)whichaccountfor99.99%oftheinteractionsforboth
strategists!

6.E(B;B)>E(A;B)orifE(B;B)=E(A;B)thenE(B;A)>E(A;A).

7.Fromthepayomatrix,E(B;B)=0:5andE(A;B)=1:0.Thus,Bisnotstabletoinvasionby

A.
8.Yes,providedthatE(A;A)=E(B;A).

9.Absolutelynot.RememberthatapureESSisnotinevitable.Inatwostrategygame,ifone
strategyisnotapureESSthenyoumusttesttoseeiftheotherisaswell.Ifitisn't,thenthe
solutionisamixedESS.WewilllaterseeinthreestrategygamesthatifnopureESSisfound
usingtheruleswehavejustlearnedtheneitheramixedESSornoESSatallarethepossible
solutions.

10.Probablynot.Nocostswouldseemtoimplyaverybriefcontestwithinjuriesonlygoingtoth
e
loser.Thatimpliesacontestthatisprobablyveryasymmetrical|thewinnerisabletoquickly
impressitssuperiorityontheloser.Yethawkvs.hawkcontestsaresupposedtobesymmetrical
.
Ithasfrequentlybeendocumented(inmanygametheorybasedstudies)thatanimalsthatare
evenlymatchedtendto
ghtlongerandinjuryismorelikelytooccur.Insuchcases,atleast
minorinjurieswouldbeexpectedeventothewinner.Thenthereareenergy,timeandperhaps
evenpredationcoststhatwouldbeexpectedtobeincurred.Forinstanceinmylab,Ihavefound
thatthecostsofstrugglesinspidersareveryhigh,especiallywhencomparedtowalkingor
othermoreroutineactivitiesthatmightcloselyapproximatedisplays.Moreover,thesestru
ggles
involveanaerobicmetabolism(whichtakesspidersalongtimetorecoverfrom)anddepletion
ofstoresofcompoundsveryimportanttorapidmotion.Andsometimesstrugglescanlastfora
considerableperiodoftime.
Chapter12.Appendix:DiscussionandSelectedAnswers
102

11.ForH:E(A;A)isE(H;H)=..25.E(B;A)isE(D;H)=0.E(H;H)<E(D;H),therefore
HisnotanESS.ForD:E(D;D)=+15andE(H;D)=+50,thereforeDisnotanESSsince
E(D;D)<E(H;D).ThereisnopureESS

12.Iftheanimalhasareasonableexpectationofcontinuedreproduction,ifitpassesbythepr
esent
ghtforthisparticularresource,andifinjuriesarelikelytobesevereandlowersigni
cantly
itsfuture
tness,thenCwouldbelargecomparedtobene
t.Ayoungmaleelephantsealwith
littleprospectofactuallyholdingasectionofbeachoccupiedbyfemalesandgreatchanceof
injuryagainstalargerexperiencedmalewouldbeadecentexampleofthissortofsituationwhe
re
C>B.

13.ThefrequencyofHawkish=0:58.Thus,thehefrequencyofDoveisd=(1..h)=0:42.

12.2AnswersforChapter4

1.d)Onesituationtolookatiswherethepayotowinningtheresourceislessthanzero.But
youneedtoaskyourselfthequestion,\Wouldanyanimalworkforasuchapayosinceit
lowersits
tnessfromwhatitwouldgetwithoutexhibitingthebehavior?"Anothersituation
wouldbetosettheinjuryordisplaycoststoalow(nearzero)value.Thinkaboutwhatthese
sortsofpayosmean|couldyoutranslateanyofthemintosituationswehavetalkedabout
inclassthatdealwiththebehaviorsofrealanimals.

2.Atvaluesgreater1.0orlessthan0.0|inotherwords,atimpossiblefrequencies!

4.a)Ittellsyouthatmeasuresofrelative
tnessarejustthat|relative.Evolutionisagame
ofrelativeadvantage(assumingthatthereisatleastenoughreproductionsothatthereis
anothergeneration!).Inthistwostrategygame,wheneverHawkismore
tthanDove,ithas
arelative
tnessof1.0(andthesameistrueofDove).Also,eitherstrategycanmaintaina
relative
tnessof1.0eventhoughitsabsolute
tnessdecreaseswithachangeinitsfrequency
(forexample,theabsolute
tnessofHawkdecreaseswithincreasingnumbersofHawks).All
thathastobetrueisthatitismore
tthanthealternative.Thus,usingthedefaultpayos,
Hwillincreaseatlowfrequencieseventhoughwheneveritincreasesitactuallyresultsina
loweringoftheaveragereproductionofthenextgenerationofHawks.

c)Ifthestrategyisatagreaterfrequencythanitsequilibrium,its
tnessisloweredasaresultof
arelativelylargenumberofunfavorableinteractions|forinstance,usingdefaultpayos,H
versusHinteractionsarehighlyunfavorabletoH.Thus,asthefrequencyofHdecreasesand
fewerofthesecontestsoccur,the
tnessofHincreases(proportionatelyithasmorefavorable
interactionswithDoves).Thevaluewhererelative
tnessstopsincreasingisthemixedESS
point|inthecaseofHawk,theproblemswithrunningintootherHawksareexactlybalanced
bythebene
tsofintimidatingDoves!
Chapter12.Appendix:DiscussionandSelectedAnswers
103

5.a)Thisindividualwillhavea
tnessthatisavery,verynegativenumber.Itdoesn'tmatter
howlonganindividuallives,ifitdoesn'treproduce(orsomehowgainindirect
tnesswhich
isaseparateissue),its
tnessiszero.NoticethatthisisaDove-likestrategythatcannot
work!Itisalsonotaveryrealisticonesincethereareusuallyplentyofwaysforananimal
togainacriticalresourceshortof
ghting,evenwhen
ghtingiscommon.
Noticealsothatweareinabitofamathematicalquandaryhere.Byadoptingthe
conventionthatbene
tsarepositivenumbersandcostsnegative,wecannoteasilyassignthis
strategythe
tnessitdeserves|zero.Recallthatinthesystemweareusing,apayoof
zerosimplymeansnoeecton
tness,notzero
tness.Zero
tnessesarepayosthatare
in
nitelynegative.Thisproblemisobviatedsomewhatbyusingsomepositivenumberasa
baseline
tnessandassigningbene
tsinadditiontothatnumberandcostsasvaluesbelowit
withzeroasthezero
tnesspoint.But,ifyouthinkaboutitamoment,you'llseethatnow
theproblemexistsinassigningcostswhicharenowconstrainedbetween0andthebaseline
positivevaluefor
tness.

b)ThisisaHawk-likestrategy.Therearesomeproblemswiththemathinthiscase(seethenext
partbelow)butlet'suseittomakeanimportantpoint.Deathdoesnotmattertotheallele
responsibleforthestrategysolongassomeonecarryingtheallelesucceedsinreproducing
anddoessoatasleastofgoodofarateasthealternatives.So,aslongassomeindividuals
dowinandreproduce,eventhoughtthecostsarehigh,thispayowillbehigherthanthe
payostothosewhonever
ghtandneverreproduce.

c)Generally,no.Themathematicsofthegameassumesthatindividuals
ghtanumberof
sequentialcontests.Iftheydieinthecontests,thentheirstrategy'sfrequencydecreaseso
ver
time.So,inourHawkandDovegame,ifHawkskilleachother,thefrequencyofDove(and
thereforecontestsinvolvingDove)willincreaseovertimeduringthegame.Noticethatwhen
wedeterminedthe
tnessofastrategy,wemultipliedthepayoofeachtypeofencounterby
thefrequencyoftheencounter,whichwetooktobeconstant.Thus,thegameassumesthat
noonedies.Injuryjustlowerssuccessatreproduction.

Now,thereisonewaythatdeathcanbeallowedinagame,shortofrecalculatingstrategy
frequenciesaftereverycontest.Ifeveryonesimplyengagesinonecontestwithanopponent
pickedatrandomandifnofurthercontestsoccurafter(sothatstrategy
tnessesarethe
aggregatesoftheseonetimeencounters),thenindividualscandieandthegamewillstill
work.Inthiscase,deathissimplyequatestoa
tnessofzero(orsincenoteveryonedies,
avery,veryhighinjurycost).Alternatively,thenumberoflivingindividualscouldbeused
torecalculatethestrategyfrequenciesaftereachcontest,butthatisnothowwesetupthe
mathematicsofthegame.

d)Inasimpleworldwhere
ghtingwastheonlywaytogainaccesstoamate,thenmalesthat
loseordonotengagein
ghtswouldhaveno
tness.Ifwede
nedtwostrategies,\
ght"
(Hawk-like)anddisplayor\don't
ght"(Dove-like)thepayosforthe\don't
ght"strategy
wouldtrulybezeroor,usingthenumerationschemewehaveselected,thebene
twould
bezeroandthecosts(exclusionfrommating)in
nitelylargeandnegative.Mostofthe
ghterswouldnotsucceedandwecouldde
nethecostsoflosingaslarge.Butthebene
ts
towinningwouldbeimmenseand\
ght"wouldbeapureESSsinceE(
ghtervs.
ghter.
wouldbegreaterthanE(don't
ghtvs.
ght).

However,itshouldbeeasytoenvisionastrategycouldinvadeapopulationof
ghters.
Theycouldalsoeschewall
ghtingbuttrytosneakmatings.Aslongastheyweresuccessful
sometimes,eventhoughthebene
tstheyreceivedwouldbedierentthanthoseofthe
ghter,
neverthelesstheywouldnotsueranyinjuriesandtheymightbeabletoinvade.
Chapter12.Appendix:DiscussionandSelectedAnswers
104

Innorthernelephantsealsthingslikethishappen.Largemalesdodefendsectionsofthe
beachusingdisplaysand
ghtsthatcanescalatetodeath.However,othermaleswilltryto
sneakmatings,atleastduringcertainpartsoftheirlives.Now,astowhetherornotthis
sneakingstrategyisanESSorsimplyamatteroftryingtodothebestyoucangiventhat
youdon'townthebeachwilldependonwhetherornotthe
tnessesareequal.Comparedto
oursimpleHawkandDovegame,thissituationismuchmorecomplexandifitwereanESS
(noonehaslookedatit,tomyknowledge)itmightwellinvolvemixesofstrategiesovera
lifetime(notoneortheotherforanentirelifetime,althoughthisprobablyhappensinsome
otherspeciessuchassalmon)thatwereconditionalonananimal'ssize,age,andgeneral
physicalcondition.SosimplegameslikeHawkandDovemayhelpustolearnaboutsome
ofthefactorsinvolvedwithanimalcontests,buttheyneedlotsofre
nementbeforewecan
reallyunderstandtheoftenhighlysophisticatedbehaviorofanimals.

6.a)Thereare850individualsofstrategyAand125ofstrategyBatthestartforatotalpopulat
ionsizeof
850+125=975.Aswithanycalculationoffrequency,

numberofmembersofthegroup
frequencyofsomegroup.

totalpopulationsize

so
frequencyofstrategyA=850=975=0:872

andsincethereareonlytwostrategies,then

frequencyofstrategyB=1:0..frequencyofstrategyA=1:0..0:872=0:128:

Checkingusingtheformulaabove,frequencyofstrategyB=125=975=0:128.
b)StrategyBleavesanaverageof1.05ospringversusonly0.85forstrategyA.Thus,using
theformulaforrelative
tness,

relativeW(A)=0:85=1:05=0:81
relativeW(B)=1:05=1:05=1:00

c)Sinceweknowtheaveragenumberofospringproducedasexuallybymembersofeach
strategy,wesimplymultiplythatnumbertimesthenumberofindividualstogetthenumber
inthenextgeneration.ForstrategeA

absoluteW(A)(numberofAparents)=0:85850=722:

ForstrategyB

absoluteW(B)(numberofBparents)=1:05125=131:

Thetotalospring(sizeofF1generation)is722+131=853.Therefore,

newfreq(A)=722=853=0:846

and
newfreq(B)=1:0..0:846=0:154;

BisincreasingandAdecreasing(nosurprise|afterallAisless
t).
Chapter12.Appendix:DiscussionandSelectedAnswers
105

d)Thepopulationisdeclining|from975to853.Whilethepopulationisdeclining,atthesame
timeitisevolvingmoretowardsstrategyB!

e)ThepopulationwillcontinuetodeclineuntilmostAstrategistsaregone(sincetheyonly
leave0.85ospring),WhenthefrequencyofstrategyB(whichleaves1.05ospring)reaches
acertainpointthepopulationwillbeginincreasingagain.Inourexample,thishappensat
aboutgeneration15.(SeeFigure12.1.Noteinthisgraph,thetotalpopulationsizeis
expressedrelativelytothe
rstgeneration..

Figure12.1:Theevolutionofapopulation.

Notethatthisparticularpatternisnotrequiredforevolutiontowork|itispossiblethat
apopulationcouldbeincreasingtheentiretimeonestrategywasout-competinganother!

12.3AnswersforChapter5

1.Ownershipisabroaderanddierentconceptthanisterritory.Theexactmeaningofterritoryi
s
muddiedandbeyondthescopeofthisdiscussion.Forourpurposes,let'sjustsaythatterritor
y
impliesownershiporpredominantuse(somethinglessthanownership)ofsomephysicalspaceo
f
theenvironment.Italsoimpliesownershiporpredominantuseofatleastsomeoftheresources
inthisspace(resourcesarede
nedwithreferencetoindividualsneedingtomakeuseofthem).
However,ananimalcan\own"aresourcewithoutholdingwhatisusuallyconstruedasaterritor
y.
Oneexamplemightbeamaleguardingitsmate,forinstancethroughprolongedcopulationas
iscommonininsects.
WhendiscussingtheBourgeoisstrategyitiscommontousethewordsterritoryandowner-
shipinterchangeablyalthoughtheyarenotreallyexactlythesamething.
Chapter12.Appendix:DiscussionandSelectedAnswers
106

2.AretheBourgeoisiesophisticated?WithapologiestoVirginiaWoolfandleavingdiscussio
nsof
high,low,andmiddlebrowaside,wecanaskthequestionofwhetherornotBourgeoisrepresents
areasonablysophisticated(i.e.,realistic)behavioralstrategy.
Bourgeois,likethebehaviorsonwhichitisbased,HawkandDove,isarathersimple-minded
strategy.Forinstance,anindividualpracticingBourgeoisdecideswhetherornotto
ghtentirely
onwhetherornotitownstheresourceundercontention.Bourgeoishasthepropertyofbeing
anuncorrelatedasymmetry.Theasymmetryinacontesttracestothefactiteitherowns
ordoesnotown(theopponentowns)aresource.Thestrategyisuncorrelatedsincecondition,
ghtingability,andlikelihoodofvictoryhavenothingtodowiththedecisionofwhetherornot
to
ght.Thedecisionisbasedentirelybywhetherornottheindividualownstheresource.
Thisis
neasfarasitgoes|animalsthatholdaresourcearemorelikelyto
ght.Butit
isnotuncommonforananimaltoconsiderthelikelihoodthatitwillprevailinagivencontest.
Fightsaremostoftenescalatedaairsthatfollowconsiderableamountsofassessment|ithas
beenrepeatedlydocumentedthatthemostserious
ghtsoccurwhenthepartiesareevenly
matched.Thus,amoresophisticatedtreatmentwouldcorrelatelikelihoodofescalationtoa
ghtandcontinued
ghtingwithfactorslikethevalueoftheresourceandlikelihoodofinjury
(whichisdeterminedinlargepartbyassessmentofthe
ghtingabilitiesofbothcontestants).

Nevertheless,eventhoughitissimple-minded,itstillisanadvanceoversimpleHawkand
Dovestrategies|animalsthatownresourcesoftenaremorelikelyto
ghtandthosethatdonot
areoftenmorelikelytodisplayoravoidanescalated
ghtwithanowner.

12.4AnswersforChapter6

1.No,butyoucanconsiderallthepossiblecombinations(HawkversusDove,HawkversusBourge
ois,
DoveversusBourgeois).Inanycase,ifonestrategyisstableagainstseveralotherknown
strategiesinpairwisecompetition,thenbyde
nitionitcannotbeinvadedbyanyofthemand
itcaninvadeallofthem.Thus,itisapureESSwithrespecttothesestrategies.Youshould
satisfyyourselfthatBourgeoisbeatsbothHawkandDoveaccordingtothestandardcriteria,a
t
leastwiththepayosthatwehavedescribed,andthentryplayingthesimulationallpossible
ways.

2.AbriefdiscussionofsimulationsusinginitiallydierentfrequenciesofH,DandB:
Foranyreasonablesetofpayos,BourgeoisisapureESSversusHawkandDove.Theinitial
frequencieshavenothingtodowithwhetherornotastrategyisapure(orforthatmattera
mixed)ESS|whendealingwithanESStheinitialfrequenciesonlydictatehowlongitmighttake
togettoequilibrium.Theyalsomightcausesomeinterestingstrategy
uctuationsingettingto
theESS.
Forinstance,youshouldhavenotedsituationswhereBourgeoisandoneoftheotherstrategies
(whichonedependsonthepayomatrixyouareusing)initiallybothincreaseastheother
decreases.Sometimesthechangesinfrequencyvaryovertime.Anespeciallyinterestedexamp
le
ofthisoccurswiththedefaultmatrixstartingwithf(Dove)=0:9andf(Hawk)=0:09.However,
eventuallyineverycaseBourgeoiswinsout|afterall,thatisthede
nitionofapureESS.
Chapter12.Appendix:DiscussionandSelectedAnswers
107

3.a{g)Youshouldnoticethatifyouusedthedefaultpayomatrix,HawkandDovequicklycome
tofrequenciesthatarenearthoseofamixedHawk/DoveESSthatwestudiedearlier.

h)Whatisgoingonhere?RegardlessofwhetheryoustartwithahighfrequencyofHorD,
thesameapproximateequilibriumisreached.Thisshouldn'tsurpiseyou|Bisatsucha
lowfrequencyinitiallythatitissimplynotaplayer|thusthevaluesoff(B)E(H;B)and
f(B)E(D;B)arenearlynegliable(HandDhardlyeverencountertherareB)andtherefore
apseudomixedESSisreachedbetweenthosetwo.However,notethatBisstillmore
tthan
either(see
tnesscurve)anditcontinuedtoincreaseatasteadyrate,eventuallyremoving
bothHandDwithDdisappearing
rst.

Carefullyexaminethe
tnesschangesthatoccurinthe
rstfewgenerationsandhowthey
happenwhenyoustartwitheitherHawkhighorDovehigh.

i)Therearethreepartstotheanswertothisquestion.(i)Bourgeoisisveryrareinitiallyand
evenifitisdoublingeachgeneration,doublingsomethingextremelyrareitstillremainsrar
e.
(ii)SinceBourgeoisissorare,Bourgeois'srelative
tnessisnotinitallyverymuchgreater
thantheothertwostrategies,mainlybecauseafterafewgenerationsBourgeoisstillhardly
encountersitself(agoodpayo)whileDoves(alsoagoodpayo)quicklybecomerarerthan
Hawks(abadpayo).And(iii)sinceBisahybridofHawkandDove,itspayosarenotvery
dierentthantheirs.

j)Thereislittleyoucando:Bisahybridstategyandanythingthathelpsitwillhelponeof
itscompetitors(tryit!!).Butitstillalwayswinsout!

4.AnytimeonestrategyisknowntobeapureESS,itstheonlyoneyouneedtomonitor.Sointhis
case,monitoringBwilltellyouwhenequilibriumisreached,whilstmonitoringHorDcouldbe
deceptive{onemightgoextinctwhiletheothercontinuesandthereforeneverreachequilibri
um.

12.5AnswersforChapter7

2.hincreasesasvdoesbecauseitbecomesmoreworthwhileto
ghtfortheresource.

3.hdecreasesasiincreasesbecauseitbecomesmoreriskyto
ght.

4.hincreasesasddoesbecomesitbecomesmorecostlytodisplay.

5.v=40.

6.a.

Player2
Player1HawkDove
Hawk
..10100
Dove
030

b)BecauseE(D;H)=0>..10=E(H;H).
c)h=0:875.
d)v=120;ifyouusev=119,HawkisstillapureESS.Apparentlytheprogramroundso.

certainvalues.
Chapter12.Appendix:DiscussionandSelectedAnswers
108

7.a.

Player2
Player1HawkDove
Hawk
..50100
Dove
030

b)h=0:583.
c)histhesameasintheoriginalgame.

9.a)Yes;ittook62generations.
b)Yes;ittook61generations.
c)Aboutthesame;IwouldhaveguesseditwouldhaveamuchhardertimeagainsttheHawks.

d)BourgeoisisstillapureESS;1generation.

e)BourgeoisisstillapureESS;1generation.

f)Bourgeoishasamuchhardertimeagainstamixtureofstrategies;ittakes172generations
toreachanESS.
g)Probably.ItdoesnotreachanESSin200generations,buttheBourgeoisareincreasing
rapidlyinthe
nalgenerations.

10.a)v=..20.
b)Thisdoesnotmakebiologicalsensesincetheresourceshouldhavepositivevalue.

c)AmixedESSisobtained:about62%Hawksand38%Doves.

11.a)i=..120.
b)Thisis
nebecausetheinjurycostshouldbeanegativevalue.

c)BourgeoisisapureESS.

12.BourgeoisisstillapureESS.

13.a)MixedESS:0.66Hawksand0.34Bourgeois.

b)MixedESS:0.50Hawksand0.50Bourgeois.

c)MixedESS:0.60Hawksand0.40Bourgeois.

d)Becausethepayostothetwostrategiesarethesame,anymixtureofthetwoispossible.
Note:TheHawksaremoreeectiveagainsttheDovesthentheBourgeoisare,sotheyend
upreplacinganyDovesintheinitialpopulation.

14.a)Ifv+i>0,thenE(H;H)=v+i>v+i=E(B;H).
24
3vi2vv+i2vv
b)Ifv+i>0,thenE(H;B)=
+
=
+>
=
=E(B;B).
444442
c)HawkisapureESSin35generations.
d)HawkisapureESSin124generations.IttakessolongbecauseHawkhasonlyasmall
advantageoverBourgeois.
Chapter12.Appendix:DiscussionandSelectedAnswers
109

15.a)ThereisamixedESS.
b)Yes,itlookslikethe0:583Hawk,0.417DovemixedESSpredictedbytheequation.This
makessense,afterall,becausethegamehasbeenreducedtoaHawk--Dovecontest.

12.6AnswersforChapter9

1.ItmakesnodierencewhatthevalueofVisinthiscase.Asxbecomesin
nitelylarge,sodoes

x=V..x=V
eandconsequentlyitsinverse,ebecomeszero.Thatis,inthelanguageoflimits,

(1=ex=V
limP(x)=lim1..)=1..0=1:

x!1
x!1

ThereforeP(x)=1:0inallcases.

..0:6..0:6=51..0:03
2.ForV=1:P(x)=1..e1..0:55=0:45.ForV=5:P(x)=1..e=1..e.

0:6=0:5..1:2
1..0:89=:11.ForV=0:5:P(x)=1..e=1..e1..0:30=0:7.

4.Q(x)andP(x),respectively.

5.1..e..1=V
.Recallthatthechanceofquitting,(9.13),isnothingmorethan1..Q(x).Nowsince
(9.14)isessentiallythesameas(9.16),then1..e..1=Vgivesusthechanceofquitting.So,fo
r
example,ifV=1,chanceofquittingperunitcostis0.632.

6.No,theyareequivalent.Inbothcases,thecontestanthasnoideawhichmaximumcostitis
facing(providedthatencounterswithdierent
x(x)supportingstrategiesarerandominthe
mixedpopulationandthatinneithercasethemaximumcostistippedbeforebeingreached).

7.Onewaywouldbetosaythatinanycontestwithmembersofthispopulation,thereisaconstant
chanceperincrementofcostthatone'sopponentwillquit.Thiscorrespondstotheideathat
one'schanceofopposingagiventypeofsupportingstrategist(maximumx)wouldbeequalto
itsfrequencyinthepopulation(asdeterminedbyintegrating(9.12)).Supportingstrategie
swith
lowmaximumxvalueswouldbemorecommonsoyouwouldbemorelikelytofacethem.

8.Iftheopponenthassomereasontoknowvar'sintentions,therewillbestrongselectivepres
sure
forittoactinawaythatthwartsvarandservesitsownbestinterests.Forinstance,ifitis
certainthatvarwillnotquitbeforereachingtheopponent'smaxcost,itwillpaytheopponent
toquitimmediatelyandcutitslosses.Likewise,ifvariscertaintoquitonthenextmoveor
overthenextbitofcost,itwillpaytheopponenttowaitvaroutandgaintheresource(as
comparedtovarwhointhiscasegainsnothing).

9.ThisisequaltoQ(x)sinceQ(x)givesthechancethatvarhasnotquitasofcostx.

10.ThisisequaltoP(x)sinceP(x)givesthecumulativechancethatvarhasalreadyquitasof
somecostx.
Chapter12.Appendix:DiscussionandSelectedAnswers
110

11.ThisisequaltoP(x)sinceP(x)givesthechancethatvarhasenduredtocostxwithout
quittingbutwillquitbeforepayingcostx+xwherexissomeadditionalcost.Itshouldbe
lessforthesmallerrangeofcosts|i.e.,lessin0.60to0.61thanin0.60to0.62.Inthiscase,a
ll
wehavedoneismakeacostintervallargerby0.01.So,therearemorequittingtimesinthis
largerintervalandthereforeagreatertotalprobabilitythatanindividualvarwillquitwit
hin
thisinterval.

..0:60
..0:61..0:60=0:5
12.a)ForV=1:P(x)=e..e0:00546.ForV=0:5:P(x)=e,

..0:61=0:5
e0:00596.

..1:0
..1:01..1:0=0:5
..1:01=0:5
b)ForV=1:P(x)=e..e0:00366.ForV=0:5:P(x)=e..e.
0:00268.

c)Noticethatthechanceofquittingwithinaspeci
ccostintervalP(x)ofaconstantrange
(0.01)decreasesastheaveragecostoftheintervalincreases.Thisisnotbecausethechance
ofquittingper0.01incrementincosthaschanged.Indeed,itisalwaysproportionalto1=V,
ectsthelowerchance
regardlessoftheinterval.Sowhythedierence?Thedierencere
thatanindividualwillactuallyhaveplayedtothehighercost.Thus,thechanceofactually
havingplayedtox=0:60isP(0:6)=0:549butthechanceofplayingallthewaytox=1
isP(1)=0:368.Ifyouapplyaconstantchanceofremainingoverthenext0:01xtoeach
ofthesenumbers(ifV=1:0,itis0:99)youwillseethatfeweractuallyquitinthesecond
interval(becausetherearefewertheretoquit!).

13.Thisisgivenbyp(x)dxanditisaverysmallnumber.

14.varlosesanycontestthatcostslessthanm.Therearelotsofwaysthiscanhappen|eachlosi
ng
costhasauniqueprobabilityofoccurrencebasedonvar'sprobabilitydensityfunction.Thus

Z
m

CosttovarLosingto
x(x=m)=xp(x)dx:

15.Followingourusualrule,eachsidewins50%ofthetime.Sincethereisa100%chancethatvar
willplayattime0andthecostequals0,then

E(var;
x(0))=0:5[(V..m)..x]=0:5[(V..0)..0]=0:5V:

16.Allofthesechancesareindependent.Inthesecases,thereisamoreorlessconstantprobab
ility
ightofadisaster(thismightbetheworstexampleofthethreesinceclearlyapoorpilot,
per
badweather,poormaintenanceorwhatevercouldchangeyourodds).Whathappensonother
ightsdoesnotaectthenextoneyougeton.Thesamewithasteroidsandlotterytickets.As
withvar,aconstantprobabilitymeansthatitcanhappenanytimeormaybeevennotatall.
Themaindierencebetweentheseexamplesandthewarofattritionisthatinthe\war"weare
concernedwiththedistributionofquittingcostswhileintheotherexamplestheemphasisiso
n
theconstantprobabilityofsomeevent.