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T F W F O U F F O

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)VNBO4PDJBM*OTUJODUT
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UMANSOCIETIESAREEXTRAORDINARILYCOOPERATIVECOMPAREDWITHTHOSE
OFMOSTOTHERANIMALS)NTHEVASTMAJORITYOFSPECIES INDIVIDUALS
LIVESOLITARYLIVES MEETINGTOONLYTOMATEAND SOMETIMES RAISETHEIR
YOUNG)NSOCIALSPECIES COOPERATIONISLIMITEDTORELATIVESANDMAYBE
SMALLGROUPSOFRECIPROCATORS!FTERABRIEFPERIODOFMATERNALSUPPORT
INDIVIDUALSACQUIREVIRTUALLYALLOFTHEFOODTHATTHEYEAT4HEREISLITTLE
DIVISIONOFLABOR NOTRADE ANDNOLARGESCALECONmICT#OMMUNICATION
ISLIMITEDTOASMALLREPERTOIREOFSELF VERIFYINGSIGNALS.OONECARESFOR
THESICK ORFEEDSTHEHUNGRYORDISABLED4HESTRONGTAKEFROMTHEWEAK
WITHOUTFEAROFSANCTIONSBYTHIRDPARTIES!MEND(OBBESTOACCOUNT
FORNEPOTISM ANDHISPICTUREOFTHESTATEOFNATUREISNOTSOFAROFFFOR
MOSTOTHERANIMALS)NCONTRAST PEOPLEINEVENTHESIMPLESTHUMAN
SOCIETIESREGULARLYCOOPERATEWITHMANYUNRELATEDINDIVIDUALS(UMAN
LANGUAGEALLOWSLOW COSTHONESTCOMMUNICATIONOFVIRTUALLYUNLIMITED
COMPLEXITY4HESICKARECAREDFOR ANDSHARINGLEADSTOSUBSTANTIALmOWS
OFFOODFROMTHEMIDDLEAGEDTOTHEYOUNGANDOLD$IVISIONOFLABOR
ANDTRADEAREPROMINENTFEATURESOFEVERYHISTORICALLYKNOWNHUMAN
SOCIETY ANDARCHAEOLOGYINDICATESTHATTHEYHAVEALONGHISTORY6IOLENT
CONmICT AMONG SIZABLE GROUPS IS COMMON )N EVERY HUMAN SOCIETY
SOCIALLIFEISREGULATEDBYCOMMONLYHELDMORALSYSTEMSTHATSPECIFY
THE RIGHTS AND DUTIES OF INDIVIDUALS ENFORCED ALBEIT IMPERFECTLY BY
THIRDPARTYSANCTIONS
4HUS WE HAVE AN EVOLUTIONARY PUZZLE $OUBTLESS THE SOCIETIES
OF OUR 0LIO 0LEISTOCENE HOMININ ANCESTORS WERE MUCH LIKE THOSE OF
OTHERPRIMATES SMALL WITHOUTMUCHDIVISIONOFLABORORCOOPERATION





%VOLUTIONARY 0ERSPECTIVES

3OMETIMEOVERTHELASTlVEMILLIONYEARS IMPORTANTCHANGESOCCURRED
INHUMANPSYCHOLOGYTHATGAVERISETOLARGERMORECOOPERATIVESOCIETIES
'IVENTHEMAGNITUDEANDCOMPLEXITYOFTHECHANGES THEYWEREPROBABLY
THEPRODUCTOFNATURALSELECTION(OWEVER THESTANDARDTHEORYOFTHE
EVOLUTIONOFSOCIALBEHAVIORISCONSISTENTWITH(OBBES NOTOBSERVED
HUMANBEHAVIOR!PESlTTHEBILL NOTHUMANS
3OMETHINGMAKESOURSPECIESDIFFERENT ANDINTHISCHAPTERWEARGUE
THATSOMETHINGISCULTURALADAPTATION/VERTHELASTMILLIONYEARSOR
SO HUMANSEVOLVEDTHEABILITYTOLEARNFROMOTHERHUMANS CREATING
THEPOSSIBILITYOFCUMULATIVE NONGENETICEVOLUTION4HESECAPACITIES
WERE STRONGLY BENElCIAL IN THE CHAOTIC CLIMATES OF THE 0LEISTOCENE
ALLOWING HUMANS TO CULTURALLY EVOLVE HIGHLY RElNED ADAPTATIONS TO
RAPIDLYVARYINGENVIRONMENTS(OWEVER CULTURALADAPTATIONALSOVASTLY
INCREASEDHERITABLEVARIATIONAMONGGROUPS ANDTHISGAVERISETOTHE
EVOLUTIONOFGROUPBENElCIALCULTURALNORMSANDVALUES4HEN INSUCH
CULTURALLYEVOLVEDCOOPERATIVESOCIALENVIRONMENTS GENETICEVOLUTION
CREATEDNEW MOREPROSOCIALMOTIVES
7EBEGINBYREVIEWINGTHEEVOLUTIONARYTHEORYOFSOCIALBEHAVIOR
EXPLAININGWHYNATURALSELECTIONDOESNOTNORMALLYFAVORLARGE SCALE
COOPERATION 4HEN WE ARGUE THAT CUMULATIVE CULTURAL ADAPTATION
GENERATES BETWEEN GROUP VARIATION WHICH POTENTIATES THE EVOLUTION
OFCOOPERATION.EXT WESUGGESTTHATSUCHCHANGESWOULDLEADTOTHE
EVOLUTIONOFGENETICALLYTRANSMITTEDSOCIALINSTINCTSFAVORINGTRIBALSCALE
COOPERATION ANDSUMMARIZESOMEOFTHEEVIDENCECONSISTENTWITHTHIS
HYPOTHESIS &INALLY WE BRIEmY DISCUSS HOW THESE IDEAS RELATE TO THE
THEMEOFTHISVOLUME THENATUREOFEVERYDAYHUMANINTERACTIONS

#OOPERATIONIS$ElNEDAS#OSTLY 'ROUP BENElCIAL


"EHAVIOR
)NTHISCHAPTER WEUSETHEWORDCOOPERATIONTOMEANCOSTLYBEHAVIOR
PERFORMEDBYONEINDIVIDUALTHATINCREASESTHEPAYOFFOFOTHERS4HIS
USAGEISTYPICALINGAMETHEORY ANDCOMMON BUTBYNOMEANSUNIVERSAL
IN EVOLUTIONARY BIOLOGY )T CONTRASTS WITH ORDINARY USAGE IN WHICH
COOPERATIONREFERSTOANYCOORDINATED MUTUALLYBENElCIALBEHAVIOR)T
ISIMPORTANTTODISTINGUISHBETWEENCOOPERATIONINNARROW TECHNICAL
SENSEUSEDHEREANDOTHERFORMSOFCOOPERATIONBECAUSETHEYHAVEVERY
DIFFERENTEVOLUTIONARYPROPERTIES
4OSEEWHY CONSIDERAGAMECALLEDTHEhSTAGHUNTvSEE&IG SO
NAMEDBECAUSEITISTHOUGHTTOCAPTURETHESTATEOFNATUREASDESCRIBED
2OUSSEAUINHIS$ISCOURSEON)NEQUALITY!SSUMETHEREISAPOPULATIONIN

#ULTUREANDTHE%VOLUTIONOFTHE(UMAN3OCIAL)NSTINCTS 

&IGURE .OWSUPPOSETHATTHEREWEREAPOPULATIONOFPEOPLEWHOARE
PAIREDATRANDOMANDPLAYTHESTAGHUNT4HEAVERAGEPAYOFFOFEACHSTRATEGY
ASAFUNCTIONOFTHEFRACTIONOFPLAYERSWHOCHOOSETOHUNTSTAG!SSUMING
THATSTRATEGIESWITHHIGHERPAYOFFSINCREASEINFREQUENCY THEREARETWOSTABLE
EQUILIBRIAEVERYBODYCHOOSESSTAGOREVERYBODYCHOOSESHARE.OWHOWEVER
THEAVERAGEPAYOFFOFTHEWHOLEPOPULATIONISMAXIMIZEDONLYATTHEALLSTAG
EQUILIBRIUM(OWEVER UNLESSSTAGHUNTINGHASAMUCHLARGERPAYOFFTHANHUNTING
HARESHS THEBASINOFATTRACTIONOFTHESTAGEQUILIBRIUMISSMALLERTHANTHAT
OFTHELOWERPAYOFFHAREEQUILIBRIUM

WHICHPAIRSOFINDIVIDUALSHAVETWOOPTIONS4HEYCANHUNTFORhASTAGv
ORFORhHAREv(UNTINGHAREISASOLITARYACTIVITYANDANINDIVIDUALWHO
CHOOSESTOHUNTHAREGETSASMALLPAYOFF H NOMATTERWHATTHEOTHER
INDIVIDUALDOES3TAGHUNTING HOWEVER REQUIRESCOORDINATEDACTION
)FBOTHPLAYERSHUNTFORTHESTAG THEYUSUALLYSUCCEEDANDEACHGETSA
LARGEPAYOFF S(OWEVER ASINGLEINDIVIDUALHUNTINGSTAGALWAYSFAILS
ANDGETSAPAYOFFOFSEE4ABLE 
4HE BEST THING FOR THE POPULATION IS IF EVERYBODY HUNTS STAGS SO
STAGHUNTINGIShCOOPERATIVEvINSENSEOFAMUTUALLYBENElCIALACTIVITY
(OWEVER ITISNOTCOOPERATIVEINTHETECHNICALSENSEBECAUSEINDIVIDUALS
DO NOT EXPERIENCE A COST TO PROVIDE A BENElT 7HEN MOST OF THE



%VOLUTIONARY 0ERSPECTIVES

4ABLE 4HE3TAG(UNT)N2OSSEAUSPARABLE HUNTERSCANEITHER


HUNTSTAGORHARE(UNTINGTOGETHERDOESNOTAFFECTTHESUCCESSOFHARE
HUNTERSTHEYALWAYSGETASMALLPAYOFF H)FTHEYHUNTSTAGTOGETHER
THEYARELIKELYTOSUCCEEDANDACHIEVEAHIGHPAYOFFS BUTASINGLESTAG
HUNTERFAILSANDRECEIVESAPAYOFFOFZERO

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3TAG
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 H
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POPULATIONHUNTSSTAG SWITCHINGTOHUNTINGHARELOWERSANINDIVIDUALS
PAYOFF ANDTHEREFOREONCEITISCOMMON STAGHUNTINGISNOTCOSTLYIT
ISINDIVIDUALLYBENElCIAL!SSUMINGTHATSTRATEGIESWITHHIGHERPAYOFF
SPREADBECAUSEOFNATURALSELECTIONIFTHEYAREGENETICALLYTRANSMITTED
ORBECAUSESUCCESSFULBEHAVIORSAREIMITATEDIFTHEYARECULTURALLYTRANS
MITTED THENITFOLLOWSTHATBOTHBEHAVIORSAREEVOLUTIONARILYSTABLE
MEANINGTHATONCECOMMONTHEYCANRESISTRAREINVADERS)NTHEJARGONOF
GAMETHEORY THESTAGHUNTISAGAMEOFCOORDINATIONBECAUSEPLAYERSDO
BETTERIFTHEYCOORDINATETHEIRBEHAVIORWITHTHEBEHAVIOROFOTHERS
.OWCONTRASTTHESTAGHUNTWITHITSMOREFAMOUSCOUSIN THEPRISONERS
DILEMMASEE4ABLE /NCEAGAINCONSIDERAPOPULATIONOFPLAYERS
WHOINTERACTINPAIRS%ACHINDIVIDUALHASANOPPORTUNITYTOHELPHIS
PARTNER)FHEDOES THEPARTNERSPAYOFFISINCREASEDANAMOUNTBBUTTHE
HELPERSPAYOFFISDECREASEDANAMOUNTCTHISISCLEARLYCOOPERATIVEIN
THENARROWSENSE!SLONGASHELPINGPROVIDESMOREBENElTTHANITCOSTS
BC EVERYBODYISBETTEROFFIFEVERYBODYHELPS(OWEVER UNLIKETHE
STAGHUNT THEGROUPBENElCIALBEHAVIORISNOTEVOLUTIONARILYSTABLE!S
SHOWNIN&IG NONHELPERSCONVENTIONALLYLABELEDhDEFECTORSv HAVE
AHIGHERPAYOFFNOMATTERWHATTHEFREQUENCYOFHELPERSCONVENTIONALLY
CALLED COOPERATORS  4HIS MEANS THAT DEFECTORS ALWAYS INCREASE AND
EVENTHOUGHEVERYONEISBETTEROFFIFEVERYONECOOPERATES COOPERATION
CANNOTEVOLVE

4HE0OTENTIALFOR#OOPERATIONIS%VERYWHEREIN
.ATURE
/PPORTUNITIESFORCOOPERATIONAREOMNIPRESENTINSOCIALLIFE%XCHANGE
ANDDIVISIONOFLABORINCREASETHEEFlCIENCYOFPRODUCTIVEPROCESSESFOR

#ULTUREANDTHE%VOLUTIONOFTHE(UMAN3OCIAL)NSTINCTS 

4ABLE 4HE0RISONERS$ILEMMA%ACHINDIVIDUALHASTHEOPPOR
TUNITYTOCOOPERATEBYHELPINGTHEOTHERINDIVIDUAL(ELPINGINCREASED
THEPAYOFFOFTHERECEIVERUNITSANDCOSTSTHEHELPERUNIT

,EFT

2IGHT
#OOPERATE

$EFECT

#OOPERATE

BC BC

C B

$EFECT

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&IGURE &INALLYSUPPOSETHATTHEREWEREAPOPULATIONOFPEOPLEWHOARE
PAIREDATRANDOMANDPLAYTHEPRISONERSDILEMMA4HEAVERAGEPAYOFFOFEACH
STRATEGYASAFUNCTIONOFTHEFRACTIONOFPLAYERSWHOCHOOSETOCOOPERATE.OW
THEREISONLYONESTABLEEQUILIBRIA EVERYBODYDEFECTSATWHICHTHEAVERAGEPAYOFF
OFTHEWHOLEPOPULATIONISMINIMIZED4HEPAYOFFMAXIMIZINGEQUILIBRIUM
EVERYBODYCOOPERATES ISUNSTABLEBECAUSEDEFECTORSHAVEAHIGHERPAYOFFTHAN
COOPERATORS

ALLTHEREASONSGIVENBY!DAM3MITHIN4HE7EALTHOF.ATIONS(OWEVER
PARTICIPATINGINEXCHANGETYPICALLYREQUIRESCOOPERATION)NALLBUTTHE
SIMPLESTTRANSACTIONS INDIVIDUALSEXPERIENCEACOSTNOWINRETURNFOR
ABENElTLATERANDAREVULNERABLETODEFECTORSWHOTAKETHEBENElTBUT
DO NOT PRODUCE THE RETURN %XCHANGE AND DIVISION OF LABOR ALSO ARE



%VOLUTIONARY 0ERSPECTIVES

TYPICALLYCHARACTERIZEDBYIMPERFECTMONITORINGOFEFFORTANDQUALITY
THATGIVERISETOOPPORTUNITIESFORFREERIDING4HEPOTENTIALFORCONmICT
OVERLAND FOOD ANDOTHERRESOURCESISEVERYWHERE)NSUCHCONmICTS
LARGERMORECOOPERATIVEGROUPSDEFEATSMALLERLESSCOOPERATIVEGROUPS
(OWEVER EACHWARRIORSSACRIlCEBENElTSEVERYONEINTHEGROUPWHETHER
ORNOTTHEYTOOWENTTOWARANDTHUSDEFECTORSREAPTHEFRUITSOFVICTORY
WITHOUTRISKINGTHEIRSKINS(ONEST LOW COSTCOMMUNICATIONPROVIDES
MANY BENElTSCOORDINATION IS GREATLY FACILITATED RESOURCES CAN BE
USEDMOREEFlCIENTLY HAZARDSAVOIDEDTHELISTISLONG(OWEVER ONCE
INDIVIDUALSCOMETORELYONTHESIGNALSOFOTHERS THEDOORISOPENFOR
LIARS mIM mAMARTISTS ANDALLTHEREST7IDELYHELDSTABLEMORALSYSTEMS
ENFORCEDBYSTERNSANCTIONSCANSOLVEMOSTOFTHESEPROBLEMSCHEATS
COWARDS ANDLIARSCANBEPUNISHED4HEPROBLEMISTHATPUNISHMENTIS
TYPICALLYCOSTLY ANDDEFECTORSCANREAPTHEBENElTSOFTHEMORALORDER
WITHOUTPAYINGTHECOSTSOFPUNISHMENT
(OWEVER ASIDEFROMHUMANS ONLYAFEWOTHERTAXA MOSTNOTABLY
SOCIALINSECTS COOPERATEVERYMUCH)NTERESTINGLY THOSETHATHAVEARE
LIKEHUMANS SPECTACULAREVOLUTIONARYSUCCESSES)THASBEENESTIMATED
FOREXAMPLE THATTERMITESACCOUNTFORHALFOFTHEANIMALBIOMASSIN
THETROPICS3O IFCOOPERATIONPRODUCESSUCHSPECTACULARBENElTS WHY
ISITSORARE

4HE'ENETIC%VOLUTIONOF#OOPERATION2EQUIRES
!SSORTMENT
4HEANSWERISSIMPLECOOPERATIONBENElTSGROUPS SOMETIMESLARGE
SOMETIMES SMALL AND AS WE HAVE SEEN GROUP BENElTS ARE USUALLY
IRRELEVANTTOCOURSEOFORGANICEVOLUTION3ELECTIONUSUALLYFAVORSTRAITS
THAT INCREASE THE REPRODUCTIVE SUCCESS OF INDIVIDUALS OR SOMETIMES
INDIVIDUALGENES ANDWHENTHEREISACONmICTBETWEENWHATISGOODFOR
THEINDIVIDUALANDWHATISGOODFORTHEGROUP SELECTIONUSUALLYLEADS
TOTHEEVOLUTIONOFTHETRAITTHATBENElTSTHEINDIVIDUAL
3ELECTIONFAVORSCOSTLYGROUPBENElCIALBEHAVIORONLYIFTHEBENElTS
mOWDISPROPORTIONATELYTOINDIVIDUALSWHOAREGENETICALLYSIMILARTOTHE
ACTORWHOPERFORMSTHEBEHAVIOR4OSEEWHY SUPPOSETHATGROUPSARE
FORMEDATRANDOM4HENEACHPROSOCIALACTHASTHESAMEAVERAGEEFFECT
ONTHElTNESSOFHELPERSANDEGOISTS4HISMEANSTHATPROSOCIALBEHAVIOR
HASNOEFFECTONTHERELATIVElTNESSOFHELPERSANDSELlSHTYPES SOTHERE
WILLBENOCHANGEINTHEFREQUENCYOFTHESETWOTYPESINTHEPOPULATION
4HEGROUPBENElTSOFTHETRAITAREIRRELEVANTTOITSEVOLUTION!TTHE
SAMETIME ITISIMPORTANTTOSEETHATTHECOSTSOFPERFORMINGPROSOCIAL

#ULTUREANDTHE%VOLUTIONOFTHE(UMAN3OCIAL)NSTINCTS 

BEHAVIORSOLELYFALLONHELPERS ANDTHUSDECREASETHEIRlTNESSRELATIVE
TO EGOISTS 4HUS THE GROUP BENElCIAL BEHAVIORS DO NOT EVOLVE .OW
SUPPOSEINSTEADTHATGROUPSAREMADEUPOFCLOSERELATIVES3ELECTIONCAN
FAVORTHEGENESTHATGIVERISETOPROSOCIALBEHAVIORBECAUSETHEBENElTS
OFPROSOCIALACTSARENONRANDOMLYDIRECTEDTOWARDOTHERSWHOCARRY
THESAMEGENES4HUS THEBENElTSOFTHEACTRAISETHEAVERAGElTNESS
OFTHEGENESLEADINGTOTHEPROSOCIALBEHAVIOR ANDIFTHISEFFECTISBIG
ENOUGHTOCOMPENSATEFORTHECOST SELECTIONWILLLEADTOTHEEVOLUTION
OFTHEBEHAVIOR
4HISSIMPLEEXAMPLEILLUSTRATESAFUNDAMENTALEVOLUTIONARYPRINCIPLE
COSTLY GROUP BENElCIAL BEHAVIOR CANNOT EVOLVE UNLESS THE BENElTS
OF GROUP BENElCIAL BEHAVIOR mOW NONRANDOMLY TO INDIVIDUALS WHO
CARRY THE GENES THAT GIVE RISE TO THE BEHAVIOR !LTRUISM TOWARD KIN
CANBEFAVOREDBYSELECTIONBECAUSEKINARESIMILARGENETICALLY7$
(AMILTON WORKEDOUTTHEBASICCALCULUSOFKINSELECTIONIN
ANDDEDUCEDMANYOFITSMOSTIMPORTANTEFFECTSONSOCIALEVOLUTION
&ULLSIBLINGSCANCOUNTONSHARINGHALFOFTHEIRGENESTHROUGHCOMMON
DESCENT ANDCANTHEREFOREAFFORDTOHELPASIBLINGREPRODUCESOLONGAS
THElTNESSPAYOFFSARETWICETHECOSTS-OREDISTANTRELATIVESREQUIREA
HIGHERBENElTCOSTRATIO4HISPRINCIPLE OFTENCALLED(AMILTONSRULE
SUCCESSFULLY EXPLAINS A VAST RANGE OF BEHAVIOR AND MORPHOLOGY IN A
VERYWIDERANGEOFORGANISMSEG +ELLERAND#HAPUISAT1UELLER
1UELLERAND3TRASSMANN 

3ELECTIONCAN&AVOR#OOPERATION!MONG3MALL'ROUPS
OF2ECIPROCATORS
7HENANIMALSINTERACTREPEATEDLY PASTBEHAVIORALSOPROVIDESACUE
THAT ALLOWS NONRANDOM SOCIAL INTERACTION 4O SEE WHY SUPPOSE THAT
ANIMALSLIVEINSOCIALGROUPSANDTHESAMEPAIROFINDIVIDUALSINTERACTS
REPEATEDLY$URINGEACHINTERACTIONONEMEMBEROFTHEPAIRHASTHE
OPPORTUNITYTOHELPTHEOTHER ATSOMECOSTTOITSELF3UPPOSETHATTHERE
ARETWOTYPESDEFECTORSWHODONOTHELPANDRECIPROCATORSWHOUSE
THESTRATEGYHELPONTHElRSTINTERACTION!FTERTHAT HELPYOURPARTNERS
ASLONGASTHEYKEEPHELPINGYOU BUTIFTHEYDONOTHELP DONOTHELP
THEMANYMORE)NITIALLY PARTNERSARECHOSENATRANDOMSOTHATDURING
THElRSTINTERACTIONRECIPROCATORSARENOMORELIKELYTOBEHELPEDTHAN
DEFECTORS(OWEVER AFTERTHElRSTINTERACTION ONLYRECIPROCATORSRECEIVE
ANYHELP ANDIFINTERACTIONSCONTINUELONGENOUGH THEHIGHlTNESS
OFRECIPROCATORSINSUCHPAIRINGSWILLBEENOUGHTOCAUSETHEAVERAGE
lTNESSOFRECIPROCATORSTOEXCEEDTHATOFDEFECTORS



%VOLUTIONARY 0ERSPECTIVES

"EYOND THIS BASIC STORY THERE IS LITTLE AGREEMENT AMONG SCIENTISTS
ABOUTHOWRECIPROCITYWORKS4HECONTRASTWITHKINSELECTIONTHEORYIS
INSTRUCTIVE4HESIMPLEPRINCIPLEEMBODIEDBY(AMILTONSRULEALLOWS
BIOLOGISTSTOEXPLAINAWIDERANGEOFPHENOMENA$ESPITEMUCHWORK
EVOLUTIONARYTHEORISTSHAVENOTMANAGEDTODERIVEANYWIDELYAPPLICABLE
GENERALPRINCIPLESDESCRIBINGTHEEVOLUTIONOFRECIPROCITY7ORSE THERE
ISLITTLEEVIDENCETHATRECIPROCITYISIMPORTANTINNATURE4HEREAREONLY
AHANDFULOFSTUDIESTHATPROVIDEANYEVIDENCEFORRECIPROCITY ANDNONE
OFTHEMAREDElNITIVE(AMMERSTEIN 

2ECIPROCITYIN,ARGE'ROUPSIS5NLIKELYTO%VOLVE
$ESPITEITSMANYPROBLEMS THEORETICALWORKDOESMAKEONEFAIRLYCLEAR
PREDICTIONTHATISRELEVANTHERERECIPROCITYCANSUPPORTCOOPERATIONIN
SMALLGROUPS BUTNOTINLARGERONES!XELRODAND$ION"OYDAND
2ICHERSON.OWAKAND3IGMUND )NSTEADOFASSUMINGTHAT
INDIVIDUALSINTERACTINPAIRS SUPPOSETHATINDIVIDUALSLIVEINGROUPS
AND EACH HELPING ACT BENElTS ALL GROUP MEMBERS &OR EXAMPLE THE
HELPINGBEHAVIORCOULDBEANALARMCRYTHATWARNSGROUPMEMBERSOF
ANAPPROACHINGPREDATOR BUTMAKESTHECALLERSCONSPICUOUSANDTHEREBY
INCREASESTHEIRRISKOFBEINGEATEN3UPPOSETHEREISADEFECTORINTHE
GROUPWHONEVERCALLS)FRECIPROCATORSUSETHERULE ONLYCOOPERATEIF
ALLOTHERSCOOPERATE THISDEFECTORINDUCESOTHERRECIPROCATORSTOSTOP
COOPERATING4HESEDEFECTIONSINDUCESTILLMOREDEFECTIONS)NNOCENT
COOPERATORSSUFFERASMUCHASGUILTYDEFECTORSWHENTHEONLYRECOURSE
TODEFECTIONISTOSTOPCOOPERATING(OWEVER IFRECIPROCATORSTOLERATE
DEFECTORS THENDEFECTORSCANBENElTINTHELONGRUN
3OMEAUTHORSHAVEEMPHASIZEDTHATPUNISHMENTTAKESOTHERFORMS
NONCOOPERATORSAREPUNISHEDBYREDUCEDSTATUS FEWERFRIENDS ANDFEWER
MATINGOPPORTUNITIESEG "INMORE &OLLOWING4RIVERS WE
WILLCALLTHIShMORALISTICPUNISHMENTv!LTHOUGHMORALISTICPUNISHMENT
AND RECIPROCITY ARE OFTEN LUMPED TOGETHER THEY HAVE VERY DIFFERENT
EVOLUTIONARY PROPERTIES -ORALISTIC PUNISHMENT IS MORE EFFECTIVE IN
SUPPORTING LARGE SCALE COOPERATION THAN RECIPROCITY FOR TWO REASONS
&IRST PUNISHMENTCANBETARGETEDSOTHATONLYDEFECTORSAREAFFECTED
4HIS MEANS THAT DEFECTORS CAN BE PENALIZED WITHOUT GENERATING THE
CASCADEOFDEFECTIONTHATFOLLOWSWHENRECIPROCATORSREFUSETOCOOPERATE
WITHDEFECTORS3ECOND WITHRECIPROCITY THESEVERITYOFTHESANCTIONIS
LIMITEDBYTHEEFFECTOFASINGLEINDIVIDUALSCOOPERATIONONEACHOTHER
GROUPMEMBER ANEFFECTTHATBECOMESSMALLASGROUPSIZEINCREASES
-ORALISTICSANCTIONSCANBEMUCHMORECOSTLYTODEFECTORS MAKINGIT

#ULTUREANDTHE%VOLUTIONOFTHE(UMAN3OCIAL)NSTINCTS 

POSSIBLEFORCOOPERATORSTOINDUCEOTHERSTOCOOPERATEINLARGEGROUPS
EVEN WHEN THEY ARE RARE #OWARDS DESERTERS AND CHEATERS MAY BE
ATTACKEDBYTHEIRERSTWHILECOMPATRIOTSANDSHUNNEDBYTHEIRSOCIETY
MADETHETARGETSOFGOSSIP ORDENIEDACCESSTOTERRITORIESORMATES4HUS
MORALISTICPUNISHMENTPROVIDESAMUCHMOREPLAUSIBLEMECHANISMFOR
THEMAINTENANCEOFLARGE SCALECOOPERATIONTHANRECIPROCITY
4HERE ARE TWO PROBLEMS WITH MORALISTIC PUNISHMENT THAT REMAIN
TOBEEXPLAINED&IRST WHYSHOULDINDIVIDUALSPUNISH)FPUNISHINGIS
COSTLYANDTHEBENElTSOFCOOPERATIONmOWTOTHEGROUPASAWHOLE
ADMINISTERINGPUNISHMENTISACOSTLYGROUPBENElCIALACT ANDTHEREFORE
SELlSHINDIVIDUALSWILLCOOPERATEBUTNOTPUNISH3ECOND MORALISTIC
PUNISHMENTCANSTABILIZEANYARBITRARYBEHAVIORWEARINGATIE BEING
KINDTOANIMALS OREATINGTHEBRAINSOFDEADRELATIVES)TDOESNOTMATTER
WHETHERORNOTTHEBEHAVIORPRODUCESGROUPBENElTS!LLTHATMATTERSIS
THAT WHENMORALISTICPUNISHERSARECOMMON BEINGPUNISHEDISMORE
COSTLYTHANPERFORMINGTHESANCTIONEDBEHAVIOR WHATEVERITMIGHTBE
7HENANYBEHAVIORCANPERSISTATASTABLEEQUILIBRIUM THENTHEFACT
THATCOOPERATIONISASTABLEEQUILIBRIUMDOESNOTTELLUSWHETHERITISA
LIKELYOUTCOMEORNOT"OYDAND2ICHERSON 
!LTHOUGH MUCH OF THE DEBATE ABOUT MORALISTIC PUNISHMENT HAS
FOCUSEDONTHElRSTPROBLEM WETHINKTHESECONDPRESENTSAMUCHBIGGER
OBSTACLETOTHEEVOLUTIONOFCOOPERATIONINLARGEGROUPS%XPLAININGTHE
PERSISTENCEOFMORALISTICPUNISHMENTISMUCHEASIERTHANEXPLAINING
WHY MORALISTIC PUNISHMENT WOULD BE USED TO MAINTAIN COOPERATION
RATHER THAN SOME OTHER FORM OF BEHAVIOR )F MORALISTIC PUNISHMENT
IS COMMON AND PUNISHMENTS SUFlCIENTLY SEVERE THEN COOPERATING
WILLPAY!SARESULT MOSTPEOPLEMAYGOTHROUGHLIFEWITHOUTHAVING
TO PUNISH VERY MUCH 4HIS IN TURN MEANS THAT ON AVERAGE HAVING A
PREDISPOSITIONTOPUNISHMAYBECHEAPCOMPAREDWITHADISPOSITIONTO
COOPERATEINTHEABSENCEOFPUNISHMENT 4HISMEANSTHATRELATIVELY
WEAKEVOLUTIONARYFORCESCANMAINTAINAMORALISTICPREDISPOSITION AND
THEN PUNISHMENT CAN MAINTAIN GROUP BENElCIAL BEHAVIOR (OWEVER
GETTINGAROUNDTHESECONDPROBLEMISMOREDIFlCULT)FEVOLUTIONARY
CHANGEISDRIVENONLYBYINDIVIDUALCOSTSANDBENElTS THENMORALISTIC
PUNISHMENT CAN STABILIZE COOPERATION BUT IT CAN STABILIZE ANYTHING
ELSETOO"ECAUSECOOPERATIVEBEHAVIORSAREATINYSUBSETOFALLPOSSIBLE
BEHAVIORS PUNISHMENTDOESNOTEXPLAINWHYLARGE SCALECOOPERATION
ISSOWIDELYOBSERVED)NOTHERWORDS MORALISTICPUNISHMENTMAYBE
NECESSARYTOSUSTAINLARGE SCALECOOPERATION BUTITISNOTSUFlCIENTTO
EXPLAINWHYLARGESCALECOOPERATIONEVOLVESINTHElRSTPLACE



%VOLUTIONARY 0ERSPECTIVES

3ELECTION!MONG,ARGE 0ARTIALLY)SOLATED'ROUPSISNOT%FFECTIVE
'ROUP SELECTION MAY BE THE NUMBER ONE HOT BUTTON TOPIC AMONG
EVOLUTIONARY BIOLOGISTS AND AS WITH MANY HEATED CONTROVERSIES IT IS
MOREABOUTHOWTOUSEWORDSTHANABOUTWHATTHEWORLDISLIKE4HE
CONTROVERSY BEGAN IN THE EARLY S WHEN 6 # 7YNNE %DWARDS
A "RITISH BIRD BIOLOGIST PUBLISHED A BOOK THAT EXPLAINED A NUMBER
INTERESTINGBIRDBEHAVIORSINTERMSOFTHEBENElTTOTHEGROUP7YNNE
%DWARDS   !LTHOUGH THIS KIND OF EXPLANATION WAS COMMON IN
THOSEDAYS 7YNNE %DWARDSWASMUCHCLEARERTHANHISCONTEMPORARIES
ABOUTTHEPROCESSTHATGAVERISETOSUCHGROUPLEVELADAPTATIONS'ROUPS
THATHADTHEDISPLAYSURVIVEDANDPROSPERED ALTHOUGHTHOSETHATDID
NOTOVEREXPLOITEDTHEIRFOODSUPPLYANDPERISHED4HEBOOKGENERATED
A STORM OF CONTROVERSY WITH BIOLOGICAL LUMINARIES SUCH AS 'EORGE
7ILLIAMS  AND *OHN -AYNARD 3MITH  PENNING CRITIQUES
EXPLAINING WHY THIS MECHANISM THEN CALLED GROUP SELECTION COULD
NOT WORK !T THE SAME TIME (AMILTONS NEWLY MINTED THEORY OF KIN
SELECTIONPROVIDEDANALTERNATIVEEXPLANATIONFORCOOPERATION4HERESULT
WASTHEBEGINNINGOFANONGOING ANDHIGHLYSUCCESSFULREVOLUTIONIN
OURUNDERSTANDINGOFTHEEVOLUTIONOFANIMALBEHAVIOR AREVOLUTION
THATISROOTEDINCAREFULLYTHINKINGABOUTTHEINDIVIDUALANDNEPOTISTIC
FUNCTIONOFBEHAVIORS
)N THE EARLY S A RETIRED ENGINEER NAMED 'EORGE 0RICE 
 PUBLISHEDTWOARTICLESTHATPRESENTEDANEWWAYTOTHINKABOUT
EVOLUTION 5P UNTIL THAT TIME MOST EVOLUTIONARY THEORY KEPT TRACK
OF THE AVERAGE lTNESS OF ALTERNATIVE GENES JUST AS WE DID ABOVE IN
EXPLAININGKINSELECTIONANDRECIPROCITY 0RICEARGUEDTHATITWASALSO
FRUITFULTOTHINKABOUTSELECTIONGOINGONINASERIESOFNESTEDLEVELS
AMONGGENESWITHINANINDIVIDUAL AMONGINDIVIDUALSWITHINGROUPS
ANDAMONGGROUPS ANDHEDISCOVEREDAVERYPOWERFULMATHEMATICAL
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SELECTIONISCONCEPTUALIZEDASOCCURRINGATTWOLEVELSSELECTIONWITHIN
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GROUPSFAVORSGROUPSWITHMOREHELPERSBECAUSEEACHHELPERINCREASES
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MULTILEVELSELECTIONAPPROACH ANDTHEOLDERGENECENTEREDAPPROACHES
AREMATHEMATICALLYEQUIVALENT ANDIFYOUDOYOURSUMSPROPERLY YOU
WILLCOMEUPWITHTHESAMEANSWEREITHERWAY

#ULTUREANDTHE%VOLUTIONOFTHE(UMAN3OCIAL)NSTINCTS 

4HEMULTILEVELSELECTIONAPPROACHHASLEDTOARENAISSANCEINGROUP
SELECTIONINRECENTYEARS ANDTHISHASLEDTONEWWRANGLINGBETWEEN
THOSEWHOTHOUGHTTHATTHEYHADKILLEDGROUPSELECTION ANDTHOSEWHO
THINKINGINMULTILEVELTERMS SEENOTHINGWRONGWITHITEG 3OBERAND
7ILSON 4HISARGUMENTISMAINLYABOUTWHATKINDSOFEVOLUTIONARY
PROCESSESSHOULDBECALLEDhGROUPSELECTIONv3OMEPEOPLEUSEGROUP
SELECTION TO MEAN THE PROCESS THAT 7YNNE %DWARDS ENVISIONED
SELECTIONBETWEENLARGEGROUPSMADEUPOFMOSTLYGENETICALLYUNRELATED
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INVOLVINGANYKINDOFGROUPINAMULTILEVELSELECTIONANALYSIS INCLUDING
GROUPSMADEUPOFCLOSEKIN
4HE REAL SCIENTIlC QUESTION IS WHAT KINDS OF POPULATION STRUCTURE
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ISFAIRLYSTRAIGHTFORWARD3ELECTIONBETWEENLARGEGROUPSOFUNRELATED
INDIVIDUALSISNOTUSUALLYANIMPORTANTFORCEINORGANICEVOLUTION%VEN
VERYSMALLAMOUNTSOFMIGRATIONARESUFlCIENTTOREDUCETHEGENETIC
VARIATIONBETWEENGROUPSTOSUCHALOWLEVELTHATGROUPSELECTIONISNOT
IMPORTANT!OKI2OGERS (OWEVER ASWEWILLSEEBELOW THE
SAMECONCLUSIONDOESNOTHOLDFORCULTURALVARIATION

!MONG0RIMATES #OOPERATIONIS,IMITEDTO3MALL'ROUPS
4HEPUNCHLINEISTHATEVOLUTIONARYTHEORYPREDICTSTHATCOOPERATION
INPRIMATESANDOTHERSPECIESTHATHAVESMALLFAMILIESWILLBELIMITED
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WHENTHEREARELARGENUMBERSOFCLOSELYRELATEDINDIVIDUALS4HESOCIAL
INSECTS WHERE A FEW FEMALES PRODUCE A MASS OF STERILE WORKERS AND
MULTICELLULAR INVERTEBRATES ARE EXAMPLES OF SUCH EXCEPTIONS 0RIMATE
SOCIETIESARENEPOTISTIC BUTCOOPERATIONISMAINLYRESTRICTEDTORELATIVELY
SMALLKINGROUPS4HEORYSUGGESTSTHATRECIPROCITYCANBEEFFECTIVEIN
SMALLGROUPS BUTNOTINLARGERONES2ECIPROCITYMAYPLAYSOMEROLE
INNATUREALTHOUGHMANYEXPERTSAREUNCONVINCED BUTTHEREISNO
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SOCIALITY!LLWOULDBEWELLIFHUMANSDIDNOTEXIST BECAUSEHUMAN
SOCIETIES EVENTHOSEOFHUNTER GATHERERS AREBASEDONGROUPSOFPEOPLE
LINKEDTOGETHERINTOMUCHLARGERHIGHLYCOOPERATIVESOCIALSYSTEMS

2APID#ULTURAL!DAPTATION0OTENTIATES'ROUP3ELECTION
3OWHYARENOTHUMANSOCIETIESVERYSMALLINSCALE LIKETHOSEOFOTHER
PRIMATES &OR US THE MOST LIKELY EXPLANATION IS THAT RAPID CULTURAL



%VOLUTIONARY 0ERSPECTIVES

ADAPTATIONLEDTOAHUGEINCREASEINTHEAMOUNTOFBEHAVIORALVARIATION
AMONGGROUPS)NOTHERPRIMATESPECIES THEREISLITTLEHERITABLEVARIATION
AMONGGROUPSBECAUSENATURALSELECTIONISWEAKCOMPAREDWITHMIGRATION
4HISISWHYGROUPSELECTIONATTHELEVELOFWHOLEPRIMATEGROUPSISNOT
ANIMPORTANTEVOLUTIONARYFORCE)NCONTRAST THEREISAGREATDEALOF
BEHAVIORALVARIATIONAMONGHUMANGROUPS3UCHVARIATIONISTHEREASON
WHYWEHAVECULTURETOALLOWDIFFERENTGROUPSTOACCUMULATEDIFFERENT
ADAPTATIONSTOAWIDERANGEOFENVIRONMENTS
)NTHE/RIGINOF3PECIES $ARWINFAMOUSLYARGUEDTHATTHREECONDITIONS
ARENECESSARYFORADAPTATIONBYNATURALSELECTION&IRST THEREMUSTBEA
hSTRUGGLEFOREXISTENCEvSOTHATNOTALLINDIVIDUALSSURVIVEANDREPRODUCE
3ECOND THEREMUSTBEVARIATIONSOTHATSOMETYPESAREMORELIKELYTO
SURVIVEANDREPRODUCETHANOTHERS ANDlNALLY VARIATIONMUSTBEHERITABLE
SO THAT THE OFFSPRING OF SURVIVORS RESEMBLE THEIR PARENTS !LTHOUGH
$ARWINUSUALLYFOCUSEDONINDIVIDUALS THESAMETHREEPOSTULATESAPPLY
TOANYREPRODUCINGENTITYMOLECULES GENES ANDCULTURALGROUPS/NLY
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GROUPS&OREXAMPLE VERVETMONKEYGROUPSCOMPETEWITHONEANOTHER
ANDGROUPSVARYINTHEIRABILITYTOSURVIVEANDGROW BUT ANDTHISIS
THEBIGBUT THECAUSESOFGROUP LEVELVARIATIONINCOMPETITIVEABILITY
ARE NOT HERITABLE SO THERE IS NO CUMULATIVE ADAPTATION /NCE RAPID
CULTURAL ADAPTATION IN HUMAN SOCIETIES GAVE RISE TO STABLE BETWEEN
GROUPDIFFERENCES THESTAGEWASSETFORAVARIETYOFSELECTIVEPROCESSES
TOGENERATEADAPTATIONSATTHEGROUPLEVEL
4HESIMPLESTMECHANISMISINTERGROUPCOMPETITION4HESPREADOFTHE
.UERATTHEEXPENSEOFTHE$INKAINTHETH CENTURY3UDANPROVIDESA
GOODEXAMPLE$URINGTHETHCENTURYEACHCONSISTEDOFANUMBEROF
POLITICALLYINDEPENDENTGROUPS#ULTURALDIFFERENCESINNORMSBETWEEN
THETWOGROUPSMEANTTHATTHE.UERWEREABLETOCOOPERATEINLARGER
GROUPSTHANTHE$INKA4HE.UER WHOWEREDRIVENBYTHEDESIREFORMORE
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THEIRTERRITORIES ANDASSIMILATEDTENSOFTHOUSANDSOF$INKAINTOTHEIR
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EG 0ALMER ET AL  THERE IS NO NEED FOR GROUPS TO BE STRONGLY
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AREPERSISTENTCULTURALDIFFERENCESBETWEENGROUPS ANDTHESEDIFFERENCES
MUST AFFECT THE GROUPS COMPETITIVE ABILITY ,OSING GROUPS MUST BE
REPLACEDBYTHEWINNINGGROUPS)NTERESTINGLY THELOSERSDONOTHAVE
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#ULTUREANDTHE%VOLUTIONOFTHE(UMAN3OCIAL)NSTINCTS 

CONFORMITYORPUNISHMENT SOEVENVERYHIGHRATESOFPHYSICALMIGRATION
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DATACOMEFROM.EW'UINEA WHICHPROVIDESTHEONLYLARGESAMPLE
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THEYEXPERIENCEDMAJORCHANGESBECAUSEOFCONTACTWITH%UROPEANS
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IN .EW 'UINEA 3OLTIS ET AL   -ANY STUDIES REPORT APPRECIABLE
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THE RATES OF EXTINCTION OF NEIGHBORING GROUPS SEE 4ABLE   4HE
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BYCONmICTWITHNEIGHBORSANDlNALLYTOSUFFERASHARPDEFEAT7HEN
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FURTHERATTACK MEMBERSTAKESHELTERWITHFRIENDSANDRELATIVESINOTHER
GROUPS ANDTHEGROUPBECOMESSOCIALLYEXTINCT!TTHESERATESOFGROUP
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TO YEARS FORANINNOVATIONTOSPREADFROMONEGROUPTOMOSTOF
THEOTHERLOCALGROUPSBYCULTURALGROUPSELECTION
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.UMBER
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%VOLUTIONARY 0ERSPECTIVES

EVOLUTIONBYCULTURALGROUPSELECTIONMAYEXPLAINTHE YEARLAG
BETWEENTHEBEGINNINGSOFAGRICULTUREANDTHElRSTPRIMITIVECITY STATES
ANDTHElVEMILLENNIATHATTRANSPIREDBETWEENTHEORIGINSOFSIMPLE
STATESANDMODERNCOMPLEXSOCIETIES
! PROPENSITY TO IMITATE THE SUCCESSFUL CAN ALSO LEAD TO THE SPREAD
OFGROUPBENElCIALVARIANTS0EOPLEOFTENKNOWABOUTTHENORMSTHAT
REGULATEBEHAVIORINNEIGHBORINGGROUPS4HEYKNOWTHATWECANMARRY
OURCOUSINSHERE BUTOVERTHERETHEYCANNOTORANYONEISFREETOPICK
FRUITHERE ALTHOUGHINDIVIDUALSOWNFRUITTREESTHERE3UPPOSEDIFFERENT
NORMSARECOMMONINNEIGHBORINGGROUPS ANDTHATONESETOFNORMS
CAUSESPEOPLETOBEMORESUCCESSFUL"OTHTHEORYANDEMPIRICALEVIDENCE
SUGGESTTHATPEOPLEHAVEASTRONGTENDENCYTOIMITATETHESUCCESSFUL
(ENRICH AND 'IL 7HITE   #ONSEQUENTLY BEHAVIORS CAN SPREAD
FROMGROUPSATHIGHPAYOFFEQUILIBRIATONEIGHBORINGGROUPSATLOWER
PAYOFFEQUILIBRIABECAUSEPEOPLEIMITATETHEIRMORESUCCESSFULNEIGHBORS
!MATHEMATICALMODELSUGGESTSTHATTHISPROCESSWILLSPREADOFGROUP
BENElCIALBELIEFSOVERINAWIDERANGEOFCONDITIONS"OYDAND2ICHERSON
 4HEMODELALSOSUGGESTSTHATSUCHSPREADCANBERAPID2OUGHLY
SPEAKING ITTAKESABOUTTWICEASLONGFORAGROUPBENElCIALTRAITTOSPREAD
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TOSPREADWITHINAGROUP
4HERAPIDSPREADOF#HRISTIANITYINTHE2OMAN%MPIREMAYPROVIDE
ANEXAMPLEOFTHISPROCESS"ETWEENTHEDEATHOF#HRISTANDTHERULE
OF#ONSTANTINE APERIODOFABOUTYEARS THENUMBEROF#HRISTIANS
INCREASEDFROMAONLYAHANDFULTOSOMEWHEREBETWEENANDMILLION
PEOPLEDEPENDINGONWHOSEESTIMATEYOUACCEPT 4HISSOUNDSLIKEA
HUGEINCREASE BUTITTURNSOUTTHATITISEQUIVALENTTOAnPERCENT
ANNUALRATEOFINCREASE ABOUTGROWTHRATEOFTHE-ORMON#HURCHOVER
THELASTCENTURY!CCORDINGTOTHESOCIOLOGIST2ODNEY3TARKMANY2OMANS
CONVERTEDTO#HRISTIANITYBECAUSETHEYWEREATTRACTEDTOWHATTHEYSAW
ASABETTERQUALITYOFLIFEINTHEEARLY#HRISTIANCOMMUNITY0AGANSOCIETY
HADWEAKTRADITIONSOFMUTUALAID ANDTHEPOORANDSICKOFTENWENT
WITHOUTANYHELPATALL)NCONTRAST INTHE#HRISTIANCOMMUNITYNORMS
OFCHARITYANDMUTUALAIDCREATEDhAMINIATUREWELFARESTATEINANEMPIRE
WHICHFORTHEMOSTPARTLACKEDSOCIALSERVICESv*OHNSON QUOTED
IN3TARK 3UCHMUTUALAIDWASPARTICULARLYIMPORTANTDURINGTHE
SEVERALSEVEREEPIDEMICSTHATSTRUCKTHE2OMAN%MPIREDURINGTHELATE
)MPERIALPERIOD5NAFmICTEDPAGAN2OMANSREFUSEDTOHELPTHESICKOR
BURYTHEDEAD!SARESULT SOMECITIESDEVOLVEDINTOANARCHY)N#HRISTIAN
COMMUNITIES STRONGNORMSOFMUTUALAIDPRODUCEDSOLICITOUSCAREOFTHE
SICK ANDREDUCEDMORTALITY"OTH#HRISTIANANDPAGANCOMMENTATORS

#ULTUREANDTHE%VOLUTIONOFTHE(UMAN3OCIAL)NSTINCTS 

ATTRIBUTE MANY CONVERSIONS TO THE APPEAL OF SUCH AID &OR EXAMPLE
THEEMPEROR*ULIANWHODETESTED#HRISTIANS WROTEINALETTERTOONE
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THE#HRISTIANSIFTHEYWANTEDTOCOMPETEFORTHEIRSOULS CITINGhTHEIR
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STATUS AND GREATER MARITAL SECURITY WITHIN THE #HRISTIAN COMMUNITY
2OMANNORMSALLOWEDPOLYGYNY ANDMARRIEDMENHADGREATFREEDOM
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FAITHFULMONOGAMY0AGANWIDOWSWEREREQUIREDTOREMARRY ANDWHEN
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OF #HRISTIANITY -UTUAL AID LED TO SUBSTANTIALLY LOWER MORTALITY RATES
DURINGEPIDEMICS ANDANORMAGAINSTINFANTICIDELEDTOSUBSTANTIALLY
HIGHERFERTILITYAMONG#HRISTIANS

4HE#REDULITY2EQUIREDFORTHE#ULTURAL%VOLUTIONOF.OVEL&ORMS
OF#OOPERATIONIS#ONSISTENTWITHAN%VOLVED 'ENETICALLY!DAPTIVE
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'IVENTHATSUCHBEHAVIORSWERENOTFAVOREDBYNATURALSELECTION WHY
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NOVEL BEHAVIORS BY GENERATING THE CUES THAT ACTIVATE THESE MODULES



%VOLUTIONARY 0ERSPECTIVES

IN NOVEL COMBINATIONS &OR EXAMPLE COOPERATION AMONG RELATIVES


REQUIRESAMONGOTHERTHINGS AMEANSOFASSESSINGCOSTSANDBENElTS
ANDOFIDENTIFYINGRELATIVESANDASSESSINGTHEIRDEGREEOFRELATEDNESS
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)NDIVIDUALSHAVETOLEARNTHECOSTSANDBENElTSOFDIFFERENTBEHAVIORS
INTHEIRPARTICULARENVIRONMENT4HUS PEOPLEWHOLEARNTHATSINNERS
SUFFERANETERNITYOFPUNISHMENTMAYBEMORELIKELYTOBEHAVEMORALLY
THAN THOSE WHO ONLY FEAR THE REPRISALS OF THEIR VICTIMS )NDIVIDUALS
HAVETOLEARNWHOTHEIRRELATIVESAREINDIFFERENTENVIRONMENTS3OTHE
INDIVIDUALWHOLEARNSTHATMEMBERSOFHISPATRICLANAREBROTHERSMAY
BEHAVEQUITEDIFFERENTLYTHANONEWHOLEARNSTHATHEOWESLOYALTYTO
THEBANDOFBROTHERSINHISPLATOON/NCEACTIVATED SUCHCOMPUTATIONAL
SYSTEMS PROVIDE INPUT TO EXISTING MOTIVATIONAL SYSTEMS THAT IN TURN
GENERATEBEHAVIOR
4HISACCOUNTRAISESANOBVIOUSQUESTION)FCULTURALINPUTSREGULARLY
LEADTOWHATIS FROMTHEGENESPOINTOFVIEW MALADAPTIVEBEHAVIOR
WHYHASSELECTIONNOTMODIlEDOURPSYCHOLOGYSOTHATITISIMMUNE
TOSUCHMALADAPTIVEINPUTS4HISISACRUCIALQUESTION ANDWEHAVE
DEALTWITHITATLENGTHELSEWHERE2ICHERSONAND"OYDCH )N
BRIEF WE BELIEVE THAT CUMULATIVE CULTURAL EVOLUTION CREATES A NOVEL
EVOLUTIONARY TRADEOFF 3OCIAL LEARNING ALLOWS HUMAN POPULATIONS TO
ACCUMULATEADAPTIVEINFORMATIONOVERMANYGENERATIONS LEADINGTO
THE CULTURAL EVOLUTION OF HIGHLY ADAPTIVE BEHAVIORS AND TECHNOLOGY
"ECAUSE THIS PROCESS IS MUCH FASTER THAN GENETIC EVOLUTION HUMAN
POPULATIONSCANEVOLVECULTURALADAPTATIONSTOLOCALENVIRONMENTS AN
ESPECIALLYVALUABLEADAPTATIONTOTHECHAOTIC RAPIDLYCHANGINGWORLD
OFTHE0LEISTOCENE(OWEVER THESAMEPSYCHOLOGICALMECHANISMSTHAT
CREATETHISBENElTNECESSARILYCOMEWITHABUILTINCOST4OGETTHEBENElTS
OF SOCIAL LEARNING HUMANS HAVE TO BE CREDULOUS  FOR THE MOST PART
ACCEPTINGTHEWAYSTHATTHEYOBSERVEINTHEIRSOCIETYASSENSIBLEAND
PROPER AND SUCH CREDULITY OPENS UP HUMAN MINDS TO THE SPREAD OF
MALADAPTIVEBELIEFS4HISCOSTCANBESHAVEDBYTINKERINGWITHHUMAN
PSYCHOLOGY BUTITCANNOTBEELIMINATEDWITHOUTALSOLOSINGTHEADAPTIVE
BENElTSOFCUMULATIVECULTURALEVOLUTION

.ATURAL3ELECTIONIN#ULTURALLY%VOLVED3OCIAL%NVIRONMENTSMAY
HAVE&AVOREDNEW 'ENETICALLY4RANSMITTED0ROSOCIAL3OCIAL)NSTINCTS
7EHYPOTHESIZETHATTHISNEWSOCIALWORLD CREATEDBYRAPIDCULTURAL
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IN OUR LINEAGE #ULTURAL EVOLUTION CREATED COOPERATIVE GROUPS 3UCH

#ULTUREANDTHE%VOLUTIONOFTHE(UMAN3OCIAL)NSTINCTS 

ENVIRONMENTSFAVOREDTHEEVOLUTIONOFASUITEOFNEWSOCIALINSTINCTS
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ANDINTERNALIZESUCHNORMS.EWEMOTIONSEVOLVED LIKESHAMEAND
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IN GROUPCOOPERATION%VENTUALLY HUMANPOPULATIONSCAMETORESEMBLE
THEHUNTER GATHERINGSOCIETIESOFTHEETHNOGRAPHICRECORD7ETHINKTHAT
THEEVIDENCESUGGESTSTHATAFTERABOUT YEARSAGOMOSTPEOPLE
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FRIENDS4HETRIBALINSTINCTSTHATSUPPORTIDENTIlCATIONANDCOOPERATION
INLARGEGROUPS AREOFTENATODDSWITHSELlSHNESS NEPOTISM ANDFACE
TO FACE RECIPROCITY 0EOPLE FEEL DEEP LOYALTY TO THEIR KIN AND FRIENDS
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WORK4HELISTISENDLESS
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THATTHISSTORYISTOOCOMPLICATED7OULDITNOTBESIMPLERTOASSUMETHAT
CULTUREISSHAPEDBYAPSYCHOLOGYADAPTEDTOSMALLGROUPSOFRELATIVES
7ELL MAYBE"UTTHESAMEPEOPLEALMOSTUNIVERSALLYBELIEVEANEQUALLY
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ACQUISITIONDEVICEEG 0INKERn 3UCHINNATELANGUAGE
INSTINCTS MUST HAVE COEVOLVED WITH CULTURALLY TRANSMITTED LANGUAGES
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COEVOLVEDWITHCULTURALLYTRANSMITTEDSOCIALNORMS)NITIALLY LANGUAGES



%VOLUTIONARY 0ERSPECTIVES

MUSTHAVEBEENACQUIREDUSINGMECHANISMSNOTSPECIlCALLYADAPTEDFOR
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COMMUNICATION4HOSEINDIVIDUALSINNATELYPREPAREDTOLEARNALITTLE
MOREPROTOLANGUAGE ORLEARNITALITTLEFASTER WOULDHAVEARICHERAND
MOREUSEFULCOMMUNICATIONSYSTEMTHANOTHERSNOTSOWELLENDOWED
4HEN SELECTION COULD FAVOR STILL MORE SPECIALIZED LANGUAGE INSTINCTS
WHICH ALLOWED STILL RICHER AND MORE USEFUL COMMUNICATION AND SO
ON7ETHINKTHATHUMANSOCIALINSTINCTSCONSTRAINANDBIASTHEKIND
OFSOCIETIESTHATWECONSTRUCT BUTTHEDETAILSARElLLEDINBYTHELOCAL
CULTURALINPUT7HENCULTURALPARAMETERSARESET THECOMBINATIONOF
INSTINCTSANDCULTUREPRODUCESOPERATIONALSOCIALINSTITUTIONS

%XPERIMENTS)NDICATE0EOPLEHAVE0ROSOCIAL)NSTINCTS
,OTSOFCIRCUMSTANTIALEVIDENCESUGGESTSTHATPEOPLEAREMOTIVATEDBY
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-ANSBRIDGE 0EOPLEGIVETOCHARITY OFTENANONYMOUSLY0EOPLE
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