The Evolution of Reciprocal Sharing
Jim Moore
GenetIcal models of the e\olutlon of reciprocal altrmsm
was distinct from cooperation. mutuahsm. or nepotismI
have ddllcultv explammg the mitral estabhshment of an
altrmst gene in a selfish deme Though potential mech-
anisms have been suggested. there IS an alternatl\e
much “altrmstlc” behatlor ma\ m fact be pureI\ selfish
m orlgm and consequentI) reclprocit\ need not be m-
*oked to protide a selectwe benefit to the actor Shanng
and hr$mg are fundamentalI\ different behavior cate-
genes and should not be confused Patterns of resource
shanng m chimpanzees correspond to predictions made
b\ a sellish model but not to those of a reciprocal altru-
1sm model, and man) observations of human gift ex- change are consistent with the selfish. but not the al-
truistic. model This suggests that presumed hominid
meat exchange mav have been the result of competition
not altnusm or eben cooperation. and that e\olutlonar\
models of “altruistic” behavior should be treated with
caution
hey ifords 4ltrwm Reclprocu) Chlmpanree Hom-
mid
INTRODl’CTION
43 emphavzed b\ M llson I 1975 p 3) true al-
trul>m 15 blologtcall~ parado\lcal m that dn) ge-
nettc InchnatIon towards \ell-sacnlice mu5t cl151
m >p~ttt of that \er\ bacrlfice Trl\er\ addre:s\ed
the problem of Jtrul5m hetbern unrelated In-
dlvtdual5 In ;\ landmark paper I 1971) He rrrgurd
that \uch 4trwm 15 nd\antageou, to 11% prx-
tlttoner If directed onI1 toHard those Indl\~duA\
it ho UIII reclprocdte gt\en the right co\t-ben-
efit ratto over ttmr each member of a reclp-
rocal relatlonshtp will do better than b) acting
alone 45 ConcelLed bv Tnbers reClprOCdl al-
truism 15 mo51 Itkel\ to evolve tn species \ilth
long It\e> good memorte5 and ~IKOUS popu-
lations tthosr tn H htch tno Indt\tduals ~111 meet
many times and remember each other 3 pa51 be-
ha\ Ior 1 There are at lea51 1v.o dtfficultles mtth Trek-
er5 formulatton First. the mttlal altruist In a
populatton 15 a lone altrul>t \ttthout rectprocltl
the tndl\ldudl IS at a dt>ad\antage 50 the trait
n111 not spread I Wd5on 1975 pp IX-121 I Thts
mtttal dtstdkantage applw equalI> to all altruts-
IIC beha\ Ior regardless of \pecte\ and ha> been
addressed a5 a populatton genetics problem tv
vseral author5 tWll5on 1977 Boorman and
Lebttt I980 Fagen 1980 Auelrod and Hamilton
1981 I all of Hhom hake relied on arguments that
\ome form of drift hm or group Aectton 15 nec-
e55ar) to establish the altru151 gene tn a popu-
lation For example failure of ktn-recognttlon
>\ 51ems due IO genetic or demographlc change\
could lead to km-selected dltrul5m becoming ac-
cldentall) generaltrcld to nonkln a 5e:cond se1 ol
adaptations then arlstng IO enlorce dtscnmln+
[Ion agaln5t nonkln nonrecIprocator I Alelrod
and Hamtlton 19811 Second among human3 It
13 b\ no mean5 clear that the rectprocal ex-
changes to which Trt\er, refers are altruistic
in an) cen5e of the word tSahltn\ 19761 Contraq
IO what 14 often assumed much human rectp-
rocal gift-gll mg IS charactertred not bq altrutsm
a5 ~\ct commonI\ use the term but lx prev
tatton \\hrch are In theor) Loluntari disin-
teresled and spontaneous hut are in tact oblig-
ator) and interested The form u\uallq Liken IS
that ol the gift grnerou~l\ ottered bur the A-
cornpan) tng beha\ tor I\ formal pretence and \o-
CIA dcceptton Hhlle the transxtlon ttwlf 15
6
DEFINITIONS
Author\ dl\cu\\lnp human .tl1rul\m h.l\e tended
to Jdopt term\ Iron1 ctjmmcw u\Jge \\ rthout
cle.~rl\ dellrnng them \~orJ~ \uch J\ nltru- i\m helping cooperJtion .md \hJr-
w JII hJ\e \llghtl\ dlfterent cc~nnotJtton\ Jnd
II IX Important to dl5tlngul\h Jmong them For
e\Jmplt: Jlthough Trl\w ~\~IIcII~\ e\cludd
cooperJtwn lco~qwrJtl\c huntIngI In hi\ trtt.lt-
men1 ot reciproL,ll ~ltruiwl \wlrnd and HJm- 11ton t~tld J recent pdprr The e\alutlon ()I CO- 0perJtion Jnd cnnslder ~onperJtlon Jltruww
Jnd re\rr.nnt during competition 10 be reIdted
phenonww e~pl~c‘.~ble by either bin wlc‘cllon or
reclprl~cJtlLm theor\ f \wl~od Jnd HJmllton
19x1 I In the preceding p~“JgrJph\ I hJ\e uwd
thr: term Jllrul\m a\ Trl\er\ Jnd other, hJ\e ~14 J 5rngle conceptual tr,ut For m.ln\ purpo\e\
ho\\e\el II I\ ImportAnt IO m.thr &I dl\tlnc1lon
het\rern helping and 5harlng
ing penrrdll) cannot le g J baboon eliciting help In 3 fight Gould onI1 double he problem\
h\ JttJching a noncomplidnl potential all) and
threat5 Irom a drwnnlng ptrwn mai not lx \er\
etfecll\el
-\/frrr~c/rc bt#rcr~ w I\ her? &lined a) behJ\- ior th.11 benttlit~ an UnrelJted indt\ 1dual1~1 \\ hlle
being detrtmrnlsl Ico\t benelltt to the actor
In the short term Art\ Jddtttonnl long-ttirm ben-
t‘ltt\ to the actor ~\thtch might rr\s,tl the C-B
tnequ.lltt\ I are conttnpent on bubwquttnt beha\ - wr tw the rttctptrnt tar other41 wc‘r \shtch the
JCtl)r has no cixItrol coot .HId hen~til .1re
Sl/~Vf I( 11)1 Jnd L,vrc lClll/ hlutusl bent IIt .m J c’\trrme wit-intri~~t can t-v combined to produ,e niutualiwi Ic‘ g inseLI pollination 0I
Ilo\rer\r In \I hlch both pJrtw berefit nsJrl\ \I-
mJltJnWU5l\ tram <aCh tntera~tton M hen the
re\rdrdb Jre tmmttdto tar both JchWl ch<dter\
LJn bc punl5hed \impl\ b\ termtndting 1he in-
terJctwn rln the case nl ~lmultrrneou~ benelit 1 or
b\ ph\\~cal attaLL II the che.ltrr trw IL) Ie~\e
\\lthout rtclprocstlng IL< hen ben&it~ Jr-s w-
qurntlJl irrth ;I short Intc‘r\Jll Hwe\rr ,I\ the
intsr\al bet\iwn rs\s.Ird\ lncreJ\c‘b 11 lwomr\ eawr lor one tndt\ I~~UJI 11) c hrdl at the e\ptlne
ol the other cla>vL;Ll \&Ltlon (a\ortng sAfi\h-
net\ should thpn decouple mUtUJl hen&t Irnm wll-interr~t II the Intel\dl het\\een InlerdLllun~
IS \er\ long-e g It J IrlL’nd \J\e\ nie tram
drwning Jnd no1 unlil ren Iedr\ lJter JrC the
t~hlc\ turned-there I\ I~ttl~: to \IO~ the beneti- LlJr\---me-~rolh \\Jlking aiiJ\ Jnd It I\ ditfi,ull
10 blOlofl~Jll\ e\plain m\ LkLlw)n to rE’CIpr1)
<are It on the other h.lnd m\ tlwnd \r~wld l\lbrnehw I benefit ImmellJ1el\ tram m\ >JI\J
IilvI there 15 no allrui~m and nc) pJrJdO\ L~rp- ICJII\ there must be wntt‘ length ot time helo\\
\rhlch benefits Are prompt Jnd mutualiw Jde
quote lo explain brhJ\ kw (the >hort Isrm 1 .md aho\ c: u hlc h the potenttJl lor cheating I\ w greJt
The EL olut~on OI Reciprocal Sharing 7
19821 No global rank >\,tern 13 lmphcd b\
the finding of a po\ltl\e correl~tlon bet\\een bo-
clal rank or prestige and RS among monkc\s
apes and humans of both se\e\ I Hau5ldter 1975
Russ .md Goodall 1977 Gomlla 1975 Hat-tune
1976 Chagnon 1979 Iron\ lY7Y Deu sbur\
IVXZI
FOOD SHARING B\ CHlhlP4NZEES AND Hl’hlANS
Chlmpdnrees u ho hake meat \!III share It filth
others u ho beg for it I Lam Ick-Goodall I968 Te-
Iekl lY73L albeit sometIme\ reluctantl) lNl\hldd
et al 19791 The >harlng ot plant foods I\ Ie\f
common and occurs pnmarll\ \\lthln lamI-
Iw-of 457 transfer5 ol banana\ ob\er\ed A
Gombr 391 18hCc1 \\ere bet\\ern mother and otf-
spring and 47 ( IO?) \\ere tram adult male:\ to
unrelated adult females I hlcGre\\ 1975) Food
shanng among relatr\e> clearI\ doe5 not require
a rrclproctt\ theor) the dlscu\\lon here ii111
therefore be restrtctcd to con\umptlon ol \cr-
tebrste pre\ (often dssumed to be a ke\ tdctor
tn human e\ olutlon-e g SchAler and Len thcr
1969)
that selt-mtero\t should wm (the long
term I nece\sttatmg an r\plnnatlon tor reel-
proLIt\ \uch as TrlLrrs or I offer Rigorous Jet-
mltlons of short term and long term need
to be developed but are not nece\>ari for un-
derstsndmg or te,tmg the model presented here
Tnver3 listed fi\r t\pe> of human beha\lor
that he con4rred AtrulstlC I I I cdrmg lor sick
dlsdbled or otheru l>e Incompetent mdl\ Idual>
(21 \harmg kno\\lrdge 13) toad sharing (41 \har-
mg Implements and 15, helping In time, of dnn-
ger Caring for Incompetent Indl\ ~duals Include,
aspects ol shanng and of helpmg a> defined
abate Sharing of knom ledge doe5 not ed,rl) fit
either catrgor) Homeber mo%t practlLal knot{ I-
edge Ldnnot be Implemented \rlthout \harmp
it among all interested ob\er\ers c g among
the ‘Kung San of the Kalaharl there I\ ier\ little
actike mstructlon ot children-the) simpl\
watch adults and practice M hat the) \cte (Draper
1976 \ce al>o Whltmg and Whltmp I975 p lx01
If mdlvlduals Share knouledge pa>\l\cl\ II I\
unneces\ar\ to tnbokr altruism ol an) sort to
eiplam the phenomenon MLGren I 1977) make\
a slmllar dt\tmctlon betaren ob>er\atlonal
learning and Inbtructton m chlmpanrcrs
In this article I use the spccllic term, ~r&rrrp
and shrrnq \\ hen it IS neces+dr) to dlrtlngulrh
them othcr\%~se both are Included m the general
term nlrrrtrsm Cooperatlon doe\ not tit the ba\lc
condttlon of altruism m that all the actors deme
lmmedlate benefits from the dLt of cooperation
classlcal I mdtvldual I sclectlon I\ sufliclent to ac-
count for the e\olutlon of cooperattve behablor
tTmers 1971 I Except uhere specified coop-
eration IS not conslderrd m this paper
Donr~ntrnc (J and prrcrrgr are con\ldered to be
two aspects of the 5ame phenomenon analogous
to the dtstmctlon made b) Maslo\t 11917) be-
t\heen ’ face-to-face and cultural domi-
nance Both dominance and prestige are usualI\
associated wtth access to rr5ource5 that can af-
fect reproductive success tRS1 and mdt\ Idual<
are expected to behave so as to maxlmlze their
status on one or both of these axes The cor-
relation between dommance tor pre:stlge) and RS
need not be perfect tar selectton to favor attn-
butes that contrtbute to dominant Status hke-
UIS~ domtnancc rankings constructed nlth ref-
erence to dtfferent behavioral domains te g
acces\ to plant food and access to mate\) need
not correlate perfectI\ lor domtnancr IO be a
meaningful concept Isee. for example Smuts
Chimpanzee predation and meat \hartng ha\
been dlrcussed extensl\el) and de\cnptlon\ can
be tound m Lawlck-Goodall I 19681 Telekl
(19731 Nlshtda et al I 19791 Kananaka tlYX21
and elsewhere The \ahent points for thl, JIS-
cushion are as foIloHs
SPA Ajjrrencrs-Hunting and meat Len-
sumptlon are pnmartlq male actnltles (above au-
thors also Wrangham 1975 McGrcn 1979 but
note exceptions do occur-e g Moms and
Goodall 19771 Telekl found that female\ recel\e
meat roughlq as often as do males but the) tend
to get much smaller porttons after most of the
ktll has been consumed Famlllal and perhaps
hormonal status pIa\ a part m females’ meat con-
sumption Just IUO females. Flo and her daugh-
ter F~fi account for oker half of Telekl’s obser-
bations of female partlctpatlon tn takmg
requesting mteractlons and the onI1 female he
sah receivrng a fair portion was GI~I who
was m r\trous at the time GI~I IS an Infertile
posslblv adrogemzed female tR13s and Goodall
1977) so II IS not clear whether her masculmtt)
or her estrous contnbuted to thts success Be-
cause meat shanng IS a prcdommantlq male ac-
ttbtty m chimpanzees. the model presented here
could be aoohed to ’ innate sex dtfferences m . .
human reciprocal e:uchange or rn the dI\Irlhutlon
of hln-dlreLrrd bersu\ nonkln dltrul\rn IF
dt~\ 510 /t cltll~w th es t’\f\f Comempornr) \ie~I-
ern cuhure rend> 10 dl~cnmlna~e agam\r \\om-
en 5 particlpalion in pubilL --clfecti\el~
nonhm-affair, and con\rquemlI drmon~trd-
tlon of an) hiporhrllcal Innare \ZY dlttrrences
Ias dislmcl from cultural t 13 probdbl\ impo5,lblr
and 14 not attempred here The model presented
I\ based on genellc kelfishness and Jn J\socla-
tion bet\tern >talus dnd reproduLf1on in pnn-
ctple it applier equal11 uell 10 fem.rle\ and male5
Igee Hrdb 1981 tor a re\reu of Lompetltron
among temale pnmatesl
Prrdclrrott cud ~lrc c otrrrol rrh~ -During
rhe tirsl mornen tollo\rmg a capture an\ In-
d~~ldual pre\rnt can grab a piece In fact \orne
preb are killed IncIdentall) b\ dlsmrmbrrmenl
ITrlekl 1973, \er\ rhortt\ though the holder
of meaf becomes 11s po,sc,sor and \natchlng
pieces 15 more or lrcs replaced h\ \ariou3 torm5
of begging Dominance <1;11115 IS nor totall\ QI)-
pendrd and \er\ IOH-ranbmg Indl\lduat\ do tend
1o to\e IheIr hrll\ bur rn general the holder a\-
surncb m haI Telrkl terms the control role and
other\\ use domlnonr rndl\ ~dunls u rtl respect Ihe
holder s o~nsrshrp The basic plcrure I\ one ot
clusters ot three 10 four adult, Around each
holder ot a major share begging or tr) lng ho run-
dggres\lvel) I take piece> of the prel Fallen
scraps dre eaten b\ mdlvlduals too loam-ranhlng
10 Jam the clulrers ipnmarll) \ubadult\ and
home female\, dnd rhe entire kill IS consumed In
from I S KI 9 hours (Telekl 19721
While this beha\~or has been Interpreted as
,m example of reciprocal altruism ie g Wilson
1975 p 128 McGren 19791 branghdm 119751
has suggcbted that rhe pnmark morr\atlon of rhe
nmmal posscssmg the meal rndl slmpl\ he relf-
protectIon Because chimps habe a relatIveI)
open dominance hlerarch) (ban Law Ich-Goodall
196X. Tetekl 1973, rhe po\sosor of the meal
I Ego) 15 potentIalI\ \ utnerablr ho artack b\ Oth-
ers H ho desire a share Meal IS an rasd\ carned
(and shielded) resource so rhe prob.ibilrt) ot
Other gaming possessron of the meat uuh a cas-
ual threat IS IOU whereas the cost ofa real altack
IS potentIalI) high te g energ) loss rlrk of in-
Jur)) Note that if Other Judges this potential
cost 10 be less than the benetir Ego IS slltt at
nsk furthermore Just the need to guard against
a possible attack by Other IS a problem for Ego
the resource ma) be detendabte bur there I\ no
opporrunui lo eat If It ma\ therefore be In Ego \
Immediate mleresl IO g~\e up [u/r/ ot the hill
trading II tor the opportunui 10 eal \thdt 13 letI
m rrlatrle peace rtor J IheorctlLdl di>cus5ran ot
Ihe\e I\QI~S see hla~nard Smith and Pdrher
IY7hl
Human >hannp and ekchanpe ImerJclron\
occur m three more or Ies> dlstlncl cla\\e\--re-
clprocal e\chdnge Ithe focu\ of hlau>s c anall\l\
and Triter\ \ and rn\ model\, bdrler or >ale dnd
simple nonreciprocal pi\ rng Barter need5 no t?\ - otutiondr\ explandrion except perhapc 11)
note rhac b\ ,ub>tltutlng ,hon-term tor long-
term balancing ot interactron, 11 3~1s 10 reduce
the opportunlt) for \elti<h Lheatmg on reclprocdl
exchanges Human toad >harmg turrl\ the eJr-
Iie5t form ot nonreciprocal 1d5 \!elln\ reciprocal1
grump ha\ been e\len\l\eI\ re\ie\red b\ Feln-
man I lY791 who conclude, Ihat the \ ~51 rndJorl1)
of \uch Interactions occur bet\\een relarcd In-
di\idudl\ and so fit the predlcrlon\ ot hln \etec-
lion The remdmlnp mterdctlon\-gl\ rng ~a non-
relall\e, \\rth benstir\ 10 Ihe lmtlnl aItruI\t
dpparentll depending k>,n long-rerm rrclprocd-
IIOn-dre the focus of throne\ ot rrciprocdt AI-
lrul\rn In Lonfrast IO Intratamdlal \harlng ldnd
aI50 10 Ihe balls of Tn\cr\ 5 mode11 \uc h Inter-
actions are often hostlIe and charscterlzed hi
lensIon and deception tMau>\ 1967 Sahlln\
tY6S 19761 Moreover as I) apparenlti the c;l\e
ullh chimpanzees among human3 the \hanng of
resources is socialI\ enforced For a Hadza
hunter 10 fall 10 3hare a large ammat ulth other
members of the camp m which he 13 II\ Ing I\ to
m\ltr vIolen retnbutlon I Woodburn t 972
see also Sahhns 1965 p 1661 Lee I 19731 poinr\
auf that homicide rates among the ‘Kung in-
crea5r \slth group \lze larger groups breaking
apan over d~spules about the dl<tnbutlon ot
tood Woodbum (19721 also ,tafe> thar Incred\rs
in group size can cause trouble The more SUL-
cessful hunters In partlculdr UIII be ternpled 10
depart presumabl\ belcausr the\ feel exploIted
b\ rncreasmg numhrrs of stachers Man\ gath-
erer-hunter soclellcs have elaborate rule\ go\-
ermng the dlstnbutlon of meat honeler Ihese
rules are often onI1 poorI\ self-enforced and
when opportumtles to conceal meat trom people
oulslde the farnil) an5e the\ are often taken
it: g Turnbutl 1961 pp 94-108 DeVore per-
sonal communication)
The Evolution of Reciprocal Shanng
THE EVOLUTION OF RECIPROCAL SHARING
If Ego (an mtelllgent chimp) risks a se\err bent-
mg whenever he or she catches somethmg edible
and tnes to keep It he or she u III soon learn not
to be stmgy Ego should learn qulcklv and 11111
be better off learnmg not to be stmgk rather than
not to hunt It IS these qualmes-a propenslt\
to learn the right sort of thmg and to learn It
quIckI) -that ma) have been favored b\ natural
selection (Hamburg 19631 Because of the threat
of enforcement. the propenslt\ to learn to share
with non-relatives may have evolved among
chimps and humans pure11 through mdlvldual
selection The chimp evidence IS especIalI) ,lg-
mficant because chimps possess all SIY of the
charactenstlc5 Tnvers con5lder5 favorable lor
the evolution of reciprocal altrulsm and \et meat
sharmg IS the onI1 altruism 5een outside the
contexts of
of their
et
of human
15 based on the
a chimplike social orgamzatlon rcf Revnolds
1966. McGrew 1979) uho were capable of
( polItIcal I stra-
tegies to achieve their goals (Walker Leonard
1979. Curtm 1981) Based on studies of chlm-
panzees. three propositions about these ances-
tor\ form the basl5 of the model
I They had a relatlbely open dominance hoer-
archy. bet male5 at lea\t mere hIghI mo-
111 ated to achleb e high rank I see van LaH Ick-
Goodall 1968, pp 212-214 R15s and Goodall
1977) When rank IS correlated NIth RS m-
dwlduals are expected to behave so as to
maximize rank eben uhen competltlon for
tmmedtate resources 15 not an Issue le de
Waal Hoekstra 19801 In long
positIon reward but rn 5hot-t
run. position Itself IS the reuard 1 Wash-
and 1968 p 4731 a
for high It5elf ma) exist to for-
relatlonshlps so that serious fighting
over resource5 tmcludmg matr5) IS abolded
l5ee Wilson 1975 chap 131 high rank ma\
also confer relative reproductive adiantage5
9
IIU spiteful behavior ISIlk 1980) and endo-
crmological 5treS5 effect5 on subordmstes
[Dunbar and Dunbar 1977 Kelerne 19811
The) exhIbIted begg:ng beha\lor that na5
probabl) as m chimps performed mo5t often
b) an Infant toHard IIS mother tMcGret\r
1975 Silk 1978) Such beggmg wal b\ nature
submlssl\e m that It signaled I Uant tihal
1 ou ha\ e but am l no% unable lo tahr II
lou’
I \~ould
be
thdt
m transltlon from adulthood m particular stwe to achwe status These In- do\ [duals te\t more
cost
to t
Other
(o&SF)
/ /
/ /
/ /
/
t Time I,
The two ordinate scales
Benefit
t to
Ego (EBs)
are not equivalent.
The E\olutwn ol Rrclprocal Shartng II
ttng the nxldent pa>\ due\ not Iowe the IYWC islth higher-ranking Other5 H ho LouId cheat by
but In ellect turther relnlorce> Ego \ temporJr\ euploltlnp Ego The present model on the other
gJm m >LIIU> Other rndl\ duals ha\e >ren Other hand hold, that II ,harmg I\ a mean5 ot avordlng
beg from Ego to mJmtdln po5ltwn In the hwr- dttxk and ot gdlnlng SOLIdI >talu> 11 should be
.Irch\ Other should .r\oij the AppeJrJnLe 01 prelcrentl~ll~ dlrccted tward those \i ho are
Lieakenlng Thl\ line 01 re~wninp 15 demon- mo5l Ilkrl~ lo attack and tho5e M ho hJ\e higher
>tratrd In the phenomenon ot redu-ected &?grr5- 5tdtu, to trade
\lon In M hlch the lowr of a light attack\ a lwer- Ewmtntnp tndt\tduJ wcce>\ rate5 ol eight
ranhlnp b\>t,mdttr dsmon>trdttng 10 all other m- Jult mdle Gombe chtmpanzce\ I+ rnngham
JI\IJuAI> that hi4 or her competltl\e ab111t\ ,nd tl97Ct Iound that Jllhough there Hc‘re no \I\-
rJnb ha\e no1 in tact Lhdnped (Popp and tematic dtf~ercncr~ among male> in r-ale, nl prei
De\nre 19’1 p 611 capture reldttke 5uco35 of ohtaininf meat and
Other ma\ hwe\er choo,c to rectpro- reldtt\e age \\ere po\ili~el) correlated ISpear-
fate-Jt some time in the nex future Other ma\ mdn t = 0x1 p 0 01 I N’ranghnm 5ugge515 obtain J slmlldr valuable rewurc~ and IJsWm- that tht\ correlatton 13 due to the louer rrpro-
mg Other can no more monopoltze II than could
Ego) ii111 \\ant Ego to knw about II Jnd lo beg lor II Ego \\III be on11 tw hJpp\ lo do w JY
long ~4 EBn EC\ Predtctrd then tn chtnt-
panZee\ 15 pO5ltl\e WhXtlOn lgenetlL w~cIJ~ Or ontogenctl~ I lor IrJlt) ~14w~i~te~ \\ lth I J?$!re\ \i\el >harlng JnJ ri%lprlxJtlng ld\W?l\t?~\ 10
rw4tablt\h \I~IU\I
In humrw the ho\ttltt\ of potlatch-t\pe r+
ctproctt\ can he wen tn thl> larger conte\t Ego
recJllmg Other > eJrller wbrnl\>l\e heh.l\ lor
\r~ll be motl\a~ed to rectprocate .md w the t-e-
Iattotwhtp IS begun Ego 5 dwre to rrclproc~~te
\i III be matchrtd b\ Other 5 remlndmg Ego ot the
debt to publ~~li empha\lze the current >LIIU~
relatlonshlp the demand lor repa)mcnt mJ\
ortglnall\ hake been Implwtl\ or r\plrc~tl\ a5-
wLlated with the hope that the debt \~oulJ re-
m,un If the debt relatton>htp ttwlt hcLomet\ a
ductlve value of old Indl\lduals the) nsk less.
In term\ ol potentlnl offsprmg lor rxh unit of
rewurce gained and nre therelore more ~\~lllnp
to at~d~h In tact the t\\o olde5t male\ iMIke
Jnd Hugo1 \!ere ln\ol\ed in YU’; ot obwr\ed
JttAhs wer meat 1 v = 201 Because of 5oc1rll rank change5 .lmong the
male\ dunng the study period Wrangham \+a5
unable to ~tatlstlcall\ analvze the relatton5hlp
hetheen ~LCCI\ at obtalnlng meat and social
ranh The three male5 \i ho held alpha ranh dur-
mg L+‘mnphnm 5 \rud) rhllhe Flgan and Hum-
phrr\ I ranhrd 2 1 and 5 ln wxe51 nt getting mrdt the fourth-ranked NCI> Fnben brother ot
Ftgan and the mo\t su~~e~~ful male Hugo \{a>
old and lo\\ rJnktng SImIlarI) Nlshlda I 10701
Telrhl I 19731 Jnd Kawanaka II9811 all conclude
that high-ranking males are more Itkrl\ to obtain
meat than are lwer ranking one\ though hter-
mator Impetus Iward> \harlng then II lollw\ arch14 relatton,htp\ are great11 relawd rel.ttt\c
that Jttlicultl 01 repJ\ ment not UIIIIIL \~III be 11~ other \ltuatwn4 Thu5 among chimpanzee
3 primat-\ factor in wlecltng a gill tone 4u5pect5 male\ older and/or higher-ranking tndt\ 1dual5
Chwtmas gills Jre wmrtlme> chown lor this are In fact more Itkel\ to retcet\c: \hare:s ol meat
redwnl -I\ Sahltn, ( 19761 hd5 potnled out po- de5pttr the lower probdblllt~ that the\ 1~111 re-
tlatch kula big men and other formal e\-
change \l >tem> are better undewtood In thl5 wl-
Vi\h light than the) are J*l e\antple> of Jltrui3nt
TEST OF THE MODEL
if ,hJnng 15 promoted b\ the e\pectJtlOn ot tu- turr rectprocatton and mutual henelit there it111
be a trndrnc\ 11) be \tlngi touard old mJI\Idual\
1s ho mai die More reslproc~tmp rTrl\er, IY’I I Regarding 51~1~3 Trl\er\ mahe< no prediction
J3 to chwce ot .i rectprocJl shartng partner bul
II wem\ Ithel\ that Ego >hould ~LOI~ shartng
ciprocate
The dtstnhutton 01 meat 3hartng the be\t-
knwn rumple ot reciprocal JltrUI,m in chimpanzee, I\ e\pldlneci b\ tht\ model hut not
b) that ol Trticrs
SH \RING AND RECIPROCAL ALTRl’lSM
HumJn \hanng helping and cooperating are be-
hai w,rA~ slmllar and are emottonall) tntercon-
netted II I\ pos\lble that cla551c 4fish wlec-
tton lor relctproctt\ JnJ ag!.gre\\l\t‘ \hartng itas
re\potwble tar e\Mll>hmg emotion\ that lorm
I2 Jim hloorr
SllhlhI-\R’I
thJl fiw JpeS mwing rnto .I hunting Iiled\lr
wch meJt status r\chsnge, pla\sd Jn Important
part In Jetermmmg an Indl\~duJl , ranh v.tthm
the wc1a1 hlerxLh\ and hence lnllucnced her or
hl\ RS
They model 01 the wgln\ ol rtxlprocal >harlnf
pro\ lde\ the loohed-for reproductl\e ~d\antJge
JccrUlng to an JltrUbt in J nonaltrulbl popu
IJtton Selertlnn tar tral1~ JYWCIJ~~~ \tlth thr:
Jblllt\ 10 optlmrlll\ IJk? ad\JntJ@ Ol wch rt’
ClproLJI ewhange I\ Ilhel~ to hJ\e Inllo\~rd Jnd
rhe C‘rlll~Jl mcredw ol these \rJltY in the pty?u
l.ltlon mJ\ h3\e alln\\ed true reclprocJl tr/fr/rr\~r
to become e>tahll\hed emorIon\ r\olied ln Lvw
cantext prcadnptlng lndl\~duA\ to t,lhe Jd
\Jntage l)f ne\! pl~~~lbi~i~it‘\ flw cwpe‘ratilm JnJ
helping as \rell a4 shJrlng c~w~clou~ manlpu-
lntlon 01 these emorIon\ awld then lorm the
bJ\I) ot J generJllZed Or \UclctJl rc’clprlxlt\
rthlL wch a> I\ nw Inund In human\ tTrl\rr4
1971 I and pos+l) odont0~ett‘ <&IceJIb IC on
nor and Nnrrlb 19821 \ltern.ltl\el\ II I\ irorth
notint: IhJt humJn> Jnd ~&wIncetr~ ma\ share
Jnother untque Ienturc-the ablllt\ 10 lnlllct ln
tur\ or &Jth on LonywclliL\ \iIthout \iJrnlng
Jnd \\ Ith 111tle IWM clrc,fc rl\h 1 human\ \i lth JI\-
[J&e \r~Jpon~ wch J\ by\\ \ er: blacDonJIJ JY-C Lee IY% LhJp I\ ~~dontocett‘\ b\ high-
lnten,lt! wnlc pul\rt\ Norrl\ Jnd hlohl IYXI I%? R C‘onnor pt‘rwnJl c~w~munl~dlwnl LII
tie InfOrmJtlOn I\ CI\ JIIJ~I~ I c‘t on blphln \ uw
01 high-lntcn>lt\ ,ound Jq .I tool l!eJpon Jnd
thl\ point mu51 be tre,ltCd JX sprculJti\e tot- no\r
lt trw it \\ould wgg~3 IhJt Jpparentlb .~~trui~ti~
helping In both group\ ma\ reprewnt J ~uIturC~ll\
enforced et\tension (>I nepotl\m \harlng nrl\lng
lrom \Jnctlon\ JgJln\l non,lllrul~tr rrlther th.ln
.ld\.mtJge> xauinp 1~ JItrui\ts Icl gewr.ll-
ized ,Iltrut~m Trl\<rl 19-I 1
The E\olutton of Reciprocal Sharing 13
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