A Role for Children in Hominid Evolution

Author(s): A. C. Zeller
Source: Man, New Series, Vol. 22, No. 3 (Sep., 1987), pp. 528-557
Published by: Royal Anthropological Institute of Great Britain and Ireland
Stable URL: http://www.jstor.org/stable/2802504 .
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 A ROLE FOR WOMEN IN HOMINID EVOLUTION

A. C. ZELLER
ofWaterloo
University

The proposal thatan increasein hominidfertility could resultfromcontributions made by

offspring to theirmother'ssubsistenceis examined,usingthreelinesof evidence.The firstpoint
contraststhe low rateof reproductionin the closelyrelatedgreatape taxa with the situation
existingin modernhumansand with the availableevidenceforearlierhominids.The second
sourceofdatais an examinationofthefactorswhichaffectmodernhumanfertility and how they
mighthave been influencedby the level of food supplyin the past. The thirdapproachis an
assessmentof the types of contributionsactuallymade by childrenin hunter/gatherer and
hunter/horticulturalgroupsat thepresenttime.A surveyof fourteengroupsat thislevelreveals
thatin twelvecases childrencontribute to a wide rangeof activitiesthatreducesubsistencestress
on theirmothers.In thetwo groupsin whichthelevel of contribution is low, thereproductive
ratesare also decreased.Evidencefromthesethreesourcessuggeststhatbothdirectand indirect
contributions by childrento theirown and theirmother'ssubsistencebase mayhave contributed
to thesuccessofhumansin populatingtheworld.

Introduction
In thehistoryof researchon humanevolutiona numberof factorshave been
proposedas thecausalmechanismsunderlying thehominisationprocess.Since
tool use was long consideredto be a 'marker'of humanity,themanufacture of
tools was an earlycandidateas theinstigating development(WashburnI959).
This materialevidencewas quicklylinked with the patternof huntingas a
subsistencebase whichwould differentiate hominidsfromthe otherprimates
(Washburn& LancasterI968; LaughlinI968a; WashburnI98I). The develop-
mentoflinguisticbehaviourwas seenas a corollaryoftheco-operationrequired
forhuntingin socialgroups(Hockett& AscherI964).
More recentthoughtsarebased on thereinterpretation ofdataobtainedfrom
huntingand gatheringmodelswhichsuggestthatgatheringis themajorsource
of dailyfoodin thissubsistencepatternat thepresentand mayeasilyhave been
so duringearlystagesofthissubsistencestrategy(Lee I972; Kolata I974; Tanaka
I976; Tanner I98I). This suggestionshiftsthefocusof attention frommalesas
theprimesubjectsofevolutionarypressuresto femalesas at leastequal subjects
and sourcesofadaptivedevelopments(IronsI 979). The potentialimportanceof
women in theevolutionaryprocesswas recognisedby Darwin (I872) butonly
recentlyhas thisrecognitionbeen utilisedin constructing models of hominid
development(Lancaster & Lancaster I 98 3; Lancaster
& King I 985; TannerI 98 I;
Zihlman I98I).
in proposing
oftheavailabledatagoes further
Lovejoy's (I980) interpretation
Mat, (N.S.) 22, 528-57
 A. C. ZELLER 529

thattheproductionof childrenmayalso have had a roleto playin thecourseof

human development.He argues (Johanson& Edey I98i) thatthe birthrate
increasedas early prehominidsshiftedtheirreproductivestrategyto a less
extremeversionof the 'K' selectionthatcharacterises pongids. In Lovejoy's
presentation, a morefrequent productionofoffspring increasedtherequirement
fora concentration of attentionand energyon motheringby thefemales.The
males then developed a gatheringas opposed to foragingadaptationwith
subsequenteffectson matingpracticesand the developmentof the bipedal
locomotorpatternsincethemaleneededto carrysufficient foodforhismateand
offspring. Lovejoy's focus,however,is on thebehaviouraland structural effects
on theparentsofbearingandnurturing an increasednumberofoffspring, rather
thanon how largernumbersof childrenmightactuallyaffectthe subsistence
base.
In order to begin the divergencefrom an ape to a human patternof
exploitation,some change,which may have been minorin itself,must have
occurred.Lovejoy's proposal of an increasedbirthrateis the type of minor
changewhichcould have majorramifications in termsofthefuturesuccessof a
species. An increasedbirthrate will reduce the age at which the young are
supplantedas a primaryfocusofmaternalcare.This difference is emphasisedby
a comparisonofape andhumanchildrearingpatterns.One notabledifference is
the scope of activityundertakenby young offspring.Ape infantsare totally
dependentphysicallyandemotionallyforfiveto six years,whereasbyage 3 to 4
human childrenare beginningsome typesof usefulactivity(WhitingI963;
Munroeetal. I984). Itis thepurposeofthisarticleto substantiate thisstatement
and to argue thatthe typesof activitythathuman childrenundertakecould
significantlyincreasetheirmother'sreproductivepotential.Thus, not only
could an increasednumberof childrenaffectthepatternsof adultbehaviouras
Lovejoy suggests,butitcouldalso actin a positivefeedbackcycleto increasethe
numberof offspring per female,and eventuallythe overallspecies numbers.
This would occurifchildrenwere able to contribute enough,eitherdirectlyor
indirectly,to thesubsistencebase to affectmaternalfertility
ratesby shortening
theinterbirth interval,and increasingoffspringsurvivability.
The interactionofmanyfactorsis subsumedin thisexplanatoryfocuson one
aspectofthewhole. Evolutionworksin sucha way thateach changemustbe an
integrative phenomenonforsuccessfuladaptationto occur. Changes towards
furlessness,bipedalism,language use, tool fabrication,and social grouping
were all partof thepatternleadingto hominisationJohnsonI978). Neverthe-
less it seems reasonableto propose that the continuedfeedbackinteraction
betweenbiologicaland culturalfactorswould increasingly promotethesuccess
of vital hominidadaptations.Increasednumbersof offspring, coupled with
improvingculturaladaptations,would allow hominidsto expandtheirranges
intonew regionsand eventuallypopulatetheworld.
Three typesof data will be utilisedto examinethisproposal. The firstis a
comparativestudyofreportedape andhumaninterbirth intervalsandreproduc-
tive success,in orderto substantiatethe claim thathumansreproducemore
effectively.Extrapolationsto the situationamong Australopithecines can be
made from primate models (Tanner I98I; Teleki et al. I976) and from
530 A. C. ZELLER

archaeologicaland palaeo-demographic evidence(Mann I975; Howell 1976a).

The second point to be discussedis a briefsurveyof the factorsinfluencing
currentnaturalhuman fertility and how these factorsmight have affected
ratesin the past. The thirdsource of data concernsthe contributive
fertility
activitiesof childrenin modern hunter/gatherer and limited horticultural
societies. The publishedreferencesto childhood contributiveactivityhave
primarilydealtwithpeasantand pastoralsocieties(AinsworthI985; Dumond
I975; Munroe etal. I984; Nag etal. I978; Hassan I980; White I975; Whiting
I963). However, publisheddata on economic contributions by childrenare
relativelyrare for horticultural and hunting-gathering societies. McDowell
(I98I) compiled materialon the value of childrenin Papua New Guinea
horticultural societies,while childhoodactivitieshave been noted among the
hunter-gatherer !Kung althoughtheircontributions to thesubsistencebase are
not emphasised,for reasons which will be discussed below (Draper I976;
ShostakI976).
The quantityand scope ofthedatacollectedforthisstudywereonlyintended
to providean initialidea of the potentialforthisresearchapproach. A much
largerand moresystematicsurveywould be necessaryto confirmor denythe
trendswhich are suggestedhere. I conductedinterviewswith eleven field
researchers studyingthirteen hunter/gathererand limitedhorticulturalgroups
fromnorthAmerica,Africa,and Australia/PapuaNew Guinea. They were
asked to provideinformation about children'scontributionsto eightareas of
potentialactivity,and theages at whichtheseactivitiesbegan. The materialon
the!Kung as a fourteenth groupwas takenfrompublicationsby Draper (I976)
and Shostak(I976). Information derivedfrombirthratedata, fertility factors
and childhoodactivitiesis used to examinetheargumentthattheactivitiesof
Australopithecine childrencould have been a major contributing factorto the
success of the hominidadaptationby increasingtheirmothers'reproductive
potential.This capacityforpopulationgrowthmaynot have been of a scale to
innumbers
causea rapidincrease (BirdsellI968; DeeveyI968; DumondI975;
Kennedy1975; Washburn I98I) butit wouldallowgroupsto recoverquickly
frompopulationbottleneckscaused by environmentalhazards,and eventually
to increasetherangeof hominidoccupation(Lovejoy I980; Johanson& Edey
I98I). This would contrastwithape populationgrowthwhichis estimatedto
have remainedstablethroughoutthePleistocene(Telekietal. I976).

Ape reproductive
patterns
Due to the long reproductivespan in apes, longitudinalinformationon the
interbirthintervaland completedfertility is stillbeingcollected.Thereforethe
relativelengthsof human and ape interbirth intervalsare stillunderdispute.
Lovejoy (I980) contendsthathumansin a stateof 'naturalfertility' produce
more offspring and have a shorterinterbirth intervalthanapes. Short (I976;
I984) suggeststhathumansare characterised by thelongestmammalianinter-
birthintervals.Othersconsiderthattheinterbirth intervalinhumansis equival-
entto thatin apes (Lancaster& LancasterI983; Lancaster& King I985; Sussman
 A. C. ZELLER 531

I972; Tanner I98I; Washburn I98I). However, Teleki et al. (I976), Tutin
(I980), Tutin & McGuinis (I98I), Harcourt et al (I980; i98ia; I98Ib) and
Galdikas(I978; I979; I98I), amongothers,have presenteddata whichindicate
thatthebirthintervalin all threeape speciesis longerthantheforty-four months
fornomadic !Kung (Howell I979; Lee I979) who are reportedas havingthe
longest natural human interbirthinterval (Howell I976a;b; I979; Huss-
AshmoreI980; Lee I979). Partofthisdifference ofopinionmaybe basedon the
use of data fromcaptiveanimalswho have a markedlymodifiedreproductive
patternwhichis influenced by factorswhichwill be discussedbelow.
From her studiesof wild apes, Galdikas (I978; I979; I98I) commentsthat
birthintervalsprobablyexceed fiveyearsin orangutans(Pongo). Mackinnon,
who also studiedfree-ranging orangutans,statesthatthemeanbirthintervalis
5.5 yearsor 66 months(I979). These are minimumperiods with surviving
infantsandmaybe lengthened to sevenor eightyearsifecologicaldisruptions or
non-viablebirthsintervene(Galdikas,personalcommunication).Age at first
parturition is about twelveyears,althoughfemalesoftenundergoa periodof
adolescentsterilityforseveralyearsbeforetheygivebirth.The totalnumberof
viableoffspring possiblein a forty-year lifespanwithtwenty-eight fertileyears
and a 5.5 year interbirth intervalwould be about four with two possible
non-viablebirths.This would be a veryhighrateofsuccessbutactualcomplete
fertilities
arenotknownforfreerangingorangutans(tablei).
More extensiveinformation is availableon gorillassincetheylive in groups
and can be studiedin largernumbers.Fossey (I979) has publishedknown
interbirthintervalsforeightcases,whichrangefromfortyto fifty monthsin the
mountaingorilla. However, the numbersof infantsper femalesurvivingto
threeyearsin herfourstudygroupsrangedfrom.7 to I. 57 infants/female per
ten-yearperiod (Fossey I983). Therefore,loss and replacementof unweaned
infantswould shortenthe intervalwhen comparedto contiguoussurviving
offspring.
Anothersurvey,by Harcourtetal. (I98 Ia), reportedthatFossey'sstudysite,
theKarisokeresearchareaof theVirungahabitat,was probablyone of thebest
areasforgorillasurvivalin termsoffoodand huntingpressure,at leastup until
I978. Theircollaborative studyforthetwelve-year periodI967-I979 foundthat
the averageproductionof survivingoffspring over thewhole Virungaregion
was about one infantper eightyearsfor a total of threeoffspringraised to
reproductivematurityper female(Harcourtetal. I980). Female gorillasbegin
to breed at age io-ii, a littleearlierthanotherapes, but thisis offsetby a
TABLE I. Reproductive
patterns
ofhigher
primates.
Inter
First Life Fertile birth Potential Number
Genus birth span span (months) fertility raised

Pongo I2 40 28 66 5?
Gorilla II 35 24 53 S? 3
Pan I3 38 25 68 5 2
Australopithecus
(reconstructed) I5 40 25 48? 6? 4?
Homo (Dobe!Kung) I9. 5 6o+ 25 44 7 5
532 A. C. ZELLER

shorterlifespanaveragingthirty-five years, which provides approximately

twenty-four yearsofpotentialreproduction (tablei). Sincegorillasdo notshow
obvious sexual swellings,thelengthof gestation,whichaverages225 days, or
81/2 months(Tutin& McGuinisI98 I), mustbe usedto calculateprobabletimeof
conception.It appearsthatabout 2-4 sexual cyclesoccur betweenthe end of
lactationalamenorrheaand refertilisation (Harcourtetal. I980). Their repro-
ductivetimespanis maximisedby rapidreplacementof infantswho die and a
somewhatshorterinterbirth intervalthanoccursin chimpanzees.The median
timespansinvolvedare twelvemonthsto replacea dead infantand fifty-three
monthsinterbirth intervalforviableoffspring (Harcourtet al. I980; i98 ib). In
orderforpopulationto maintaina stablelevelatleast40 percent.oftheanimals
should be immatures(Harcourtet al. i9i8a). Male infanticideand deathsof
motherscontribute to infantmortality, thuskeepingtheoverallnetpopulation
increasedown to .02 to .6 per cent.per yearin thispopulation,in spiteof an
adequatebirth-rate (see table2, Karisoke).
Whenthegorillaandwildchimpanzee(Pan) situationsarecomparedthereare
variationsin thetimespansinvolvedforreproductive activitiesbutverylittlein
the overall rate of populationgrowth. The twentyyear period surveyedat
Gombe (Goodall I979) indicatesthatinterbirth intervalsfor survivingcon-
tiguousoffspring rangefromfiveto ten years.Tutin (I980) and Teleki et al.
(I976) have examinedthereproductive patternsofknownfemalesat Gombe in
an effortto assess total fertility.Wild chimpanzeesbegin to show sexual
swellingsataboutages 8-9 withmenarcheoccurringI-3 yearslater.A periodof
adolescentsterility followsduringwhichyoungfemalesmatefrequently butdo
not conceive. Thus most femalesdo not begin to reproduceuntil they are
thirteenyearsold, and nearlyhalflose theirfirstinfant(Tutin I980). The time
requiredto replacethelost infantis slightlylongerthanthetwelvemonthsin
gorillas,averagingI2 to i6 months.Between I964 and I978 themeaninter-sib
intervalforsurvivingyoungwas 68 monthswitha rangefrom53 to 79 months,
althoughin two cases therewereintervening pregnancylosses. The numberof
cycles betweenresumption of sexual activityand the subsequentpregnancy
rangesfrom to cycles
i i i with a of
mean 4.5. Itwasnoted,however, thatittook
longer(6.3 cyclesmean) forfemalesto become pregnantthethirdtime.Tutin
suggeststhatprobablyearlypostpartumcyclesareanovulatoryas is thecase in
humans(ShortI984). Therefore,witha probablelifespanof thirty-eight years
(Telekietal. I976; Washburn I98I) and menarche at ageI3, chimpanzee females
would have a reproductivelifespanof about twenty-fiveyears. With an
TABLE 2. Infant perI000 females
production peryear.
no. of Statusof
Genus Location infants population Source

Gorilla Calculated 125-148 Maintain Harcourt et al. Ig8ia

requirement
Gorilla Karisoke 286.5 Increase Harcourtet al. I98 ia
Pan Gombe I64.3 Maintain Goodall I979
Pan Kibale 123-I66 Maintain Goodall I979
Homo Dobe Kung 128.o6 Increase Howell 1976b
(6 yearmean)
 A. C. ZELLER 533

interbirth intervalof 68 months(Teleki et al. I976; Tutin ig80) this would

providetimefor4-5 birthsto surviveto weaning(tablei). However, foreight
femalesobservedoverthelong term,theestimatednumberofoffspring ranged
fromi to 5 witha mediannumberof3 (TutinI 980; Tutin& McGuinisI 98 I). Of
thosefemaleswithknownoffspring, themaximumnumberto reachreproduc-
tiveage is three(Tutin& McGuinis i98i), and themediannumberis 2 (Tutin
ig80), althoughit is possiblethatolderfemaleoffspring mayhave migratedto
other populations.Tutin (ig80) concludes that the replacementrate in the
Gombe populationonlymatchesthedeathrate,and does notexceedit.
The situationis very similarin Nishida's chimpanzeepopulation in the
Mahale mountains.Over fourteen yearsofdatacollectiontheinterbirth interval
and speedofoffspring replacement areaboutthesame as thosedocumentedfor
Gombe (Tutin ig80). Two otherchimpanzeepopulationsstudiedby Ghiglieri
(i984) in theKibale foresthad crudebirthratesequivalentto the estimatesof
Harcourtetal. (i98 ia) forpopulationreplacement(table2).
Thus it is clearthatproductionof reproductively viableoffspring by female
apes is a veryslow undertaking (tablei). Althoughin eachspeciestheinterbirth
intervalssuggestthatabout fiveviableoffspring shouldbe produced,in actual
countsthemaximumactuallyraisedis usuallythree,witha medianoftwo. This
low rateimpedespopulationgrowthbothintrinsically and also becausetruerate
ofgrowthis based on theproductionofdaughtersto continuethespecies(gross
rateof reproduction,Howell I979). Producingtwo or threeviable sons will
keep up an individual'srepresentation in thegene pool of thenextgeneration,
but will not ensurethattheyreproduceor thatthe populationwill continue.
Anotherconstraining factorin ape reproduction is theobservationthatinfants
bornwithinsixyearsoftheendofthemother'slifewillprobablynotsurviveher
(Goodall I973; Telekietal. I976). Orphanedapes mustbe atleastthreeyearsold
to survivetheloss of theirmother'smilk,butin mostcases,in spiteof careby
others,themother'sdeathwill usuallydoom offspring undersix to deathfrom
depressionand neglect(Goodall I973; Teleki etal. I976). On a populationlevel
althoughsome individualfemalesmayhave verysuccessfulreproductive lives,
some othersmaybe sterileor nevermanageto raisean infant(Goodall I979).

toAustralopithecines
Extrapolation
When comparingmodernapes to earlyhominidsit is clearthatsome typesof
information will be unobtainable,some will be based on inferences
and only a
small proportioncan be foundedon actual data. The lengthof the interbirth
intervalforAustralopithecines is probablyundiscoverablealthoughit is esti-
mated by McKinley (in Dumond I975) as 3-5 years. The length of the
reproductivelifespancan onlybe assessedby drawingon modernhumansand
apes as models. We do, however,have some data on overalllifespanwhichis
anotherlimitingfactorin determining overallfertility.
Howell (I976a) com-
mentsthatprehistoric huntingand gatheringpopulationsprobablyfellintothe
rangeofa 20- to 30-yearlifespan.Data fromNeandertalburialssuggestthat5o
percent.ofthepopulationdied as infantsorjuveniles,butthatover30 percent.
534 A. C. ZELLER

reachedtheage of 3o; althoughonly a few (3 per cent.) were consideredto be

over 40 at death (Kennedy I976; WashburnI98I). Allan Mann argues that
Australopithecine lifespansmay have been shorterthanthis and suggestsan
averagelifeexpectancyofeighteenyears(Mann I975). This figurecan be putin
perspective,however,by realisingthatthe averagelifeexpectancyof Gombe
chimpanzeesis I3.5 years(Telekietal. I976). Once theyhavesurvivedto age I5
both chimpanzeesand Australopithecines are assesseda lifeexpectancyof 27.7
years (Hassan I980; Teleki et al. I976). This is supportedby Mann's (I975)
figuresfor a combinedsample of thirtygracileAustralopithecines in which
sevenwere underage i8, and twenty-three over age i8, withthreebeingover
age 35. He assesses the maximumlifespanas 40 years ?5, which is very
comparableto chimpanzees.
Although!Kungsubsistencepatterns haveoftenbeenusedas a modernhunter
and gatherermodelon whichto base interpretations ofearlyhominidlifestyles,
itis notthecase thattheyhaveadaptedto theircurrent habitatoverthousandsof
years(Gordon I984; Ripley I980). Nonethelessextrapolationsare oftenmade
from!Kung to Australopithecine data. The medianage at last birthfor!Kung
womenis about32.5 yearsbuttheaverageage atfirstbirthis I9.5 years(Howell
I979). Thus, proportionately, thetimeintervalbetweenestimatedage at first
birthin Australopithecus (age I5) and probablelifeexpectancy(aboutage 28) is
quite similarto thefertileintervalof thirteenyearsthatoccursstatistically for
!Kungfemales.These figuresarebasedon therelativefertile lifeexpectancyafter
theattainment of reproductiveage, as suggestedby Lancaster& King (I985).
The nomadic !Kung have an averageof 4.7 to 5 live birthsper woman witha
modal 44 monthinterbirth intervalmaintainedbylactationand postpartumsex
taboos (HarpendingI976; Howell I976b; I979; SaucierI972). Witha pre-
reproductivemortalityrate of 34 per cent. (Howell I976b) to 40 per cent.
(HarpendingI976) thisreproductivepatternprovidesa growthrateof 0. 5 per
cent./yearforthepopulation.
Anotherfactorwhich may account forsome of the populationgrowthis
increasedsurvivability. Teleki etal. (I976) estimatetheprobabilitiesof survi-
in
vorship chimpanzeesas .364 and in Australopithecines as .65o. Lancaster&
Lancaster(I983) accountforthisdifference by foodstressat theweaningstage.
Only one in threechimpanzeessurvive thejuvenile stage, but the number
doubleswhen adequateprovisioningoccurs.Thus, insteadof raisinga median
numberof two offspring to reproductivematurityout of 5-6 as chimpanzee
mothersdo, Australopithecines mayhavebeenable to raise4 out of6 produced.
Mann (I975) and Washburn(I98I) statethattheAustralopithecine patternof
tootheruptionand enamelformationindicatea slowerdevelopmentalprocess
than occurs for the chimpanzee.Argumentsby Bromage and Dean (I985)
suggest,however, thatAustralopithecus and earlyHomo demonstrate'bio-
logicalequivalenceto modernmanatroughlytwo-thirds thechronologicalage'
(Bromage& Dean I985). Mann's argumentproposesthatsexual maturity may
have been delayeduntilage I5 in Australopithecus, whereastheBromage and
Dean approachsuggestsa moreape-likepatternof I2 to I3 yearsfortheadvent
of reproductivebehaviourin females.Actualage at firstbirthwould have less
influenceon number of offspringproduced than would interbirth interval
 A. C. ZELLER 535

(Wood et al. I985). If Australopithecines had the same birthintervalsas

chimpanzees,approximatelythe same age at menarcheand an equivalent
lifespan,populationgrowthwould havebeenat aboutthesame rateas occursin
chimpanzees.If insteadof a 5-6 yearinterbirth interval,however, the birth
spacingoccurredon a more modernhumanpatternof 3 to 4 yearsas Mann
(I975) suggests,therateofoffspringproductionwould increaseby 50to ioo per
cent. Even if Pleistocenepopulationsonly increasedat the rate of 0.0007 to
o.ooi5 per cent. per annum (Dumond I975) froma startingpopulation of
IOO,OOO individuals,
thenumbers
woulddoublein IO,OOO years.Cohen(I98o)
and Hassan (I980) estimatetherateofPleistocenepopulationgrowthat .OOI to
.003 per cent. per annumwhichwould providean even fastergrowthrate.It
seems doubtfulthatearlyhominidpopulationsgrew even thatquicklybut it
seemsclearthata constant,althoughslow, rateofgrowthdistinguishes theearly
humandemographicpatternfromtheape pattern(BirdsellI968; Dumond I975;
Teleki etal. I976). This trendtowardsgrowthis important
in allowinghominid
populationsto recoverfromperiodicdisastersand to spreadinto unoccupied
areas.

Factors
affectingfertility

Captivityand theruralurbandichotomy. Captivity,with the resultantincreased

foodsupplyandlessenergeticactivity appearstodecreasetheinterbirth
patterns,
in
intervalquite markedly apes. Supportiveevidence for this includes the
numberof monthselapsingbetweenparturition and resumptionof maximum
sexual swelling.In a (small) captivesample of chimpanzeesthistime period
rangedfrom3 to 26 months,whereasin wild chimpanzeesit rangedfromi i to
8i monthswitha meanof 42 months(Tutin I980). Tutincommentsthatwild
femaleswhose infantsdie usuallyconceive again fromfourto eightmonths
later, which gives a strongindicationthat lactationand the mechanicsof
sucklingaremajorsuppressivefactorsin theresumptionoffertilecycling(Tutin
I980). Anotherfactoraffecting thetotalnumbersofoffspring producedis age at
menarche.Captive chimpanzeesreach menarcheat a mean age of 7 years,
whereasforfreeranginganimalsfirstsmall swellingsappearedat ages 8 to 9
yearsin femaleswith menarcheoccurringfromI-3 yearslater(Tutin I980).
This disparityin age may be closelyrelatedto the greaterweightsforage in
captiveanimals(Telekiet al. I976). In thissamplethemeanweightforcaptive
non-obeseadultfemaleswas 56.8 ? 2 kg., whereaswild females(usuallywith
clingingoffspring) rangedfrom32.3 to 37 kg. (Tutin I980). Since the mean
weightat menarcheforcaptivefemaleswas 29. I ? 4.2 kg. it is clearthatwild
femalesdo notgainsubstantially moreweightthanthelevelrequiredto achieve
It is quite possible thatwild chimpanzeesmust weigh the same as
fertility.
captiveones to reachmenarche,buttakelongerto gaintheweight(Teleki etal.
I976). All thesefactorssuggestthatsituationsprovidingimproveddietin terms
ofquantityand consistency, possiblycoupledwithless activelives,can have an
effecton thepotentialnumbersofoffspring born.
Among humansthereis a dichotomyin thetimingof reproductiveactivity
536 A. C. ZELLER

between rural and urban women even when nutritionlevels are presumed
adequatein bothcases. Urban girlsbeginto cycleearlierthanrurallivingones
(ShortI976). A Polishstudycomparingtheage at menarcheofruralgirlswith
thosewho livedin Warsawindicatedthatthecitygirlsbeganto cycleI.03 years
earlieron averagethanthose fromruralregions(Laska-MierzejewskaI975).
Age at menarchehas also declinedduringthiscentury.In a Frenchstudythe
meanageofvillagegirlswas I5.7 yearsin i900, andI3.4 yearsin I946. These
ages were stillin excess of themean ages at menarchein Pariswhichfellfrom
I3.5 to I2.9 yearsover thesame timespan(LeridonI977).
Among Rwandan women, urbanlivingis correlatedwith a twelve-month
reductionin theinterbirth intervalfornursingmothers,when comparedwith
rurallivingwomen(Bonteetal. I974; seealsoBongaarts I980; BuchananI975;
Van GinnekenI977; Huss-Ashmore
I980; ShortI976). This difference
may
have manyunderlying
influences
suchas socioeconomicfactors,activitylevels
andbreastfeeding BothLee (I979;
patterns. I980) andShort(I976) notethatin
thecasesof!Kung who havebecomemoresedentary andhaveaccessto milkand
grain as dietarysupplements,periods of amenorrheaand birthintervalsare
reduced 30 to 50 per cent. Modern westernwomen have mean interbirth
intervalsof 24 months, while rural Punjab women average 30.5 months
interbirthinterval(Leridon I977). Thus thereis a slight,thoughmeasurable,
differencebetweentheproportionsoftheirlivesthatwomenin ruraland urban
situationsare at riskof becomingpregnant.For cases in whichdietarydiffer-
encesarenegligible,atleastpartofthisdifference
probablyarisesfromincreased
levels of manual labour in ruralwomen (Laska-MierzejewskaI975). These
differencesofreducedage atmenarcheand shorterbirthintervalsarein thesame
directionas thedifferencesbetweenwildandcaptiveapes. Itseemsreasonableto
suggestthatbothdietand activitypatternsarefactorswhichcould affect therate
of offspring production.

Diet. Iftheinterbirth intervalsin Australopithecines

werefiveor moreyearsthe
reproductivespan would only allow the same numberof birthsas occur in
moderngorillasand chimpanzees.This is barelysufficient to maintaina stable
population,as indicatedabove. The questionthenbecomes,whatfactorscould
affectthe fertility
ratesof earlyhominids,and possiblyalso reducemortality
levels. GenerallyspeakingBronson'sstatement'Diet is thecoreenvironmental
constraint on a mammal'sreproduction'(I980: i i6) seemsto be thefoundation
offertility The caloriesconsumedmustmeettherequirements
effects. forbody
maintenance,thermoregulation, foraginglocomotion,growthand reproduc-
tion.In theabsenceofsufficient calories,growthand reproductiveeffectiveness
arethefirstprocessesadverselyaffected (BronsonI980).
The abilitya woman has to build up the nutritionalreservesnecessaryto
conceive,carrya pregnancyto termand lactateuntilthechildcan eat sufficient
amounts of food to maintainitselfis based on threefactors.These are the
availabilityof food, theenergya femalespendsacquiringit and caringforher
offspring, and thespeed withwhich theyoung can transfer theirdependence
away fromthemotheras theirmainsourceof food.
The availabilityof foodin termsof calorielevelis largelya factorof external
 A. C. ZELLER 537

conditionsand subsistencetechnique.The typeof food may also influencethe

outcomeofa pregnancy,sincelackofproteinand mineralsin thedietmaydraw
uponthemother'sbodyintheformation ofthefoetuswithresulting poor health
formothers
andnewborns
(BuchananI975; CohenI980; Huss-Ashmore
I980;
Katz I972; ScrimshawI983). If the level of proteinintakeis low the age of
pubertymay be retarded(Short I976). Body weightdoes not appearto affect
fertility as long as thecriticalmass(seebelow) is attained,butlower
significantly
maternalweightincreasesboththelengthofthelactationperiodand thelength
of lactationalamenorrhea(Premaetal. I979 in Knauer I984; BongaartsI980;
FrischI977; van GinnekenI972; Huss-Ashmore
I980; WrayI977). Frisch
(I977) also commentsthatlow maternalweighttendsto resultin a low birth
weightfortheoffspringwhichreducesitssurvivability.
The numberofavailablekcalrequiredto sustainpregnancyand threemonths
lactationareestimated
by FrischandMcAndrew(I974; FrischI975; I977) as
I44,000. Ofthisamount,Buchanansuggests
(I975) that
55,oookcalarerequired
to maintainthepregnancy.Althoughlactatingmammalsareextrememodelsof
in theuse of nutrients(Misra I979), thelevel of caloricsupportfor
efficiency
humanlactationis estimatedat I,000 kcal/dayor go,oookcal forthreemonths
(CohenI980; FrischI975). Insituations
ofrestricted
diet,calorieaccumulation
can be slow and fertilityratesareaffected. Lee's (I980) input-outputanalysisof
!Kung dietindicatesthatadultsconsume2I40 kcal/daywhileexpendingI,975
kcalto acquirefoodandmaintainthemselves.At thislevel,calorieaccumulation
to thelevelthatFrischsuggestswould takeseveralyears.Steinand Susser(I977)
investigatedthe reproductiveresultsof a five-monthfamineduringwhich
caloricintakedroppedfromI 500 to 700 kcal/dayand discoveredthatoffspring
productionwas severelycurtailed.Undertheseconditionsitwas mainlyyoung
mothersbearingtheirfirstchild who had the necessaryreservesto continue
procreation.In light of the calorie levels considerednecessaryBongaarts's
(I980) studyon thedistinction between'low' (I300 kcal/day)and 'high' (I630
kcal/day)levelsofmaternalcalorieintakeprobablyonlysampledwomen at the
low endofthescale.Thus hisresultthatcalorieintakeonlyaffected thelengthof
lactationby one month,may not be valid forhigherlevels of nutritionsince
Short (I976) statedthateven 2I00 kcal/dayand 42 gms of proteinwere not
sufficientto maintaina positivenitrogenbalancein themother.
Frisch(I975; I977; I978; I982) appearsto have established(Cohen I980) that
thereis a criticalweightof44 to 48 kilos,and proportionofbody fat(I7-22 per
cent.) whichis necessaryto allow a developingfemaleto undergomenarche.
She has also arguedthatthisstagecoincideswitha changein metabolicratecost
from35 cal/kg/day to 28 cal/kg/day and a changein proportionof lean body
weightto fatfrom5:I to 3: I (I97 5). Althoughcriticalweightseemsestablished,
thereis some argumentconcerningherassertionthatthelean:fatratiomustbe
regainedbeforereconception canoccur(Bongaartsig80;Jain& Sun I972;Jain&
BongaartsI98I; Ripley I980; ShortI984; Lee I972; I980). Frisch(I975; I978)
has suggestedthatthemechanismby whichfataffectsmenstrualcyclicityand
theabilityto conceivemaylie in theinteraction ofgonadalactivityand estrogen
levelswiththelevel of fatin thebody. Protein,calorieand vitamindeficitsare
correlatedwithgonadal hypofunction (Katz I972). Estrogenmetabolismmay
538 A. C. ZELLER

be influencedby factorsinvolvedin fatmetabolismand thus more abundant

whena highfatdietis consumed(FrishI975; I978). Therefore thana
rather
simple storage of calories, fat may provide the basis for varyinglevels of
reproductive hormones.
Cohen (I98o) hassuggestedthatthelean:fatratiomaybe irrelevant in modern
dietsbecausethelevelofcaloricintakeand thusthelean:fatratiomayno longer
provide a reliableguide to the body, indicatingan environmentsuitablefor
reproduction.He arguesthatearlyin hominiddevelopmentthelevel of caloric
intake,and thusthebody's level of storedfat,was also an indicatorof protein
and othernecessarynutrients in thediet.Consumingcurrentadultdietsloaded
with non-nutritive caloriesmay allow modernwomen to conceive and bear
youngin environments whichdo not reallyhave enoughproteinresourcesto
sustainthem. One example of this is the increasein the fertility rate to 8
offspring (Neel I983) in theYanomamo since theybegan the cultivationand
consumptionof bananas which have only I.2 gms of proteinper ioo gms
(Cohen I980). Thus caloriclevelsmayallow reproduction to occurin situations
in whichproteinresourcesareinsufficient to maintainthehealthofthechild.On
theotherhand,therearealso somepopulationsinwhichthewomenweigh35 to
40 kg. and are able to maintainpregnancyand lactationon a dietof I500-I 6oo
calories/day, with35 to 40 gm. ofprotein.Adaptationmechanismsin pregnant
women seem to allow an extremelyefficient utilisationofprotein,calciumand
iron (Beaton I96I in Misra I979). During pregnancy some nutrients appearto
be storedin thebody and releasedduringlactation(Misra I979). These factors
may contributeto the occurrenceof pregnanciesand continuedlactationin
women who do notappearto have an adequatecalorieintake.However, under
these conditionsstillbirths and low birthweightchildrenoccur 'with much
greaterfrequency'(p. I83) thanin well-nourished women (Misra I979).
If infantsare born to a poorly nourishedmothermany proteinand fat
resourcesare drawn fromher body. In situationsin which maternalcaloric
intakeis below 2,000 cal/day,thevolume of milkdropsto levelsof400 to 600
c.c. perdayratherthantheusual8oo C.c. (WrayI 977). In one study,Edozian (in
WrayI977) supplementedmaternaldietswithI00 gramsof proteinwhichwas
controlledso as notto changethecalorieintake,and milkproductionincreased
by up to 200 c. c. perday. Thus proteinlevelsin bothmother'sand infant'sdiet
area majorfactorin determining how well theinfantwill thrive.
Diet also influenceslevel of prolactionwhichcan inhibitmenstrualcycling.
This effectwas demonstrated(Lunn et al. I984) in a studyof pregnantand
lactatingGambianvillagewomen. They gave a highenergyproteinandvitamin
supplementthatraised calorie levels to 229I kcal. eitherduringlactationor
duringpregnancyand lactation.As a pregnancysupplement,it improvedthe
birthweightofchildren,and in bothcasesitloweredprolactinlevelsduringthe
lactationperiod. The women supplementedduringpregnancyand lactation
resumed cycling and ovulating in one-halfthe time that unsupplemented
women did (43 V. 79 weeks for resumptionin 50 per cent. of the women).
Improvingthedietduringpregnancyand lactationratherthanmerelyduring
lactationwas more effective in shorteningtheinterbirth interval(37 V. I9 per
cent.)whencomparedto theunsupplemented condition.
 A. C. ZELLER 539

Activity levelsandnursing patterns.The effectof maternalactivitypatternsmay

occur throughseveralchannels.In a predominantly gatheringsituation,the
quantityof food may be adequatebutthecaloricenergyexpendedin amassing
and processingit may be a significantly limitatingfactoron the woman's
capacityto buildup thenecessaryfatlevelsto conceiveand bringthepregnancy
to term(Buchanan I975; Dumond I972; Howell I976b; I979; LaughlinI968b;
Lee I972; I979; I980). Lee statesthat'any changein thesubsistenceeconomy
thatallows reducedmobilitymaybe sufficient to increasefertility'
(I979: 3I9).
This observationappearstobe substantiated bythedeclineininterbirth intervals
from44 to 36 monthsamong !Kungwomen who have becomemoresedentary
(Lee I979). Even in industrialised westernsocietywomen who engagein high
levels of activity,such as ballet dancersor marathonrunners,may undergo
primaryor secondaryamenorrhea(WarrenI979 inHuss-AshmoreI980). Frisch
statesthatthe loss of 30 per cent. of body fatreserveswill triggersecondary
amenorrhea(I977).
Anotherpossibleeffect ofmaternalactivitypatternmayresultfromtheeffect
of dailyscheduleson thefrequencyand patternsof nursing.When maximum
nursingintervalsoccurbeforesix monthshavepassed,or whennightnursingis
discontinued,cyclingis resumed more quickly even though lactationmay
continueup untilI8-20 months(Knauer I984). Women who nurseless than
fourtimesa day after6 monthsshow a drop in prolactinlevels which will
usuallyresultin theresumptionof cycling(Jain& BongaartsI98I). The use of
waterand supplementary foodsbefore6 monthswill also reducetheperiodof
lactationalamenorrhea.Full breastfeedingon demandwithoutsupplementary
foodor liquidsforsix monthsis stronglycorrelatedwithlengthenedperiodsof
amenorrhearangingfromI I to 36 months(Knauer I984). The resumptionof
cyclingmaynotbe evidenceof fertility sincethefirstfewreturning cyclesmay
be anovulatory(Short I984). On the otherhand, ovulationcan occur before
cyclingis evidentand severalstudiesindicatethata smallpercentageofwomen
becomepregnantagainwithoutmenstruating (5.4 percent.Bonte& Van Balen
I969; 6 per cent.Jain& Sun I972). Thus in spiteof folkwisdom, menstrual
cyclingcan resumewhile lactationis stillin progress(Leridon I977). This is
relevantbecause long lactationperiods have frequentlybeen suggestedas a
major cause of long interbirth intervalsin non-contracepting populations.
Infantswho geta smallervolumeof milkmaynursemorefrequently and with
greatervigour. Both frequencyand intensity of sucklingare reportedto have
markedeffectson the hormonecycle which controlsthe onset of ovulation
(Bongaarts I980; Frisch1975; Knauer I984; Konner & WorthmanI980; Lee
I972; I979; ShortI976; I984).
Thus women who leave theirinfantswitha caretakerforsubstantialpartsof
the day, even thoughtheyare stillnursing,will tendto resumecyclingmore
quicklythanthosewho keeptheinfantwiththemand feedit frequently.
Lee in particular(I972; I979; I980) statesthatcontinueddemandnursingup
to age 3 maybe a majorcause of thelong birthspacingamong the!Kung. This
argumentis based moreon thestimulusof sucklingon themother'sneuroen-
docrinesystemthanon theeffect ofcontinuednursingon themother'sfatlevels
(Konner& WorthmanI980; Lee I980). The importanceofthesucklingstimulus
540 A. C. ZELLER

is confirmed
by experimental
denervationofmammaryglandsin animals,who
thenresumedreproductivecyclingwhilelactating(ShortI976).

Weaning foods.In additionto diet, maternalactivityand patternsof nursing,

cycleresumptionis also affected by thetypesof weaningfoodsavailable.Most
societiesbegin supplementary feedingbetween 6 monthsand i year of the
infant'slife. Until 6 months,lactationcan supply all of the infant'sdietary
requirements butpastthatpointitis notsufficient, althoughfrequently itis the
infant'sonly source of high qualityprotein(Frischi982). Between 6 and I2
monthsan adequatelynourishedmothercanprovideaboutthree-quarters ofthe
infant'sproteinrequirements (BuchananI975). Scrimshaw(i983) suggeststhat
.75 gm/kghighqualitymilk,meator egg proteinis requiredto establishand
maintainadequategrowthratesand protectionfrominfection.Ifthemotheris
poorlynourishedmanyoftheseproteinandfatresourceswillbe drawnfromher
body to supplytheinfant.
Continued nursingamong the !Kung is considerednecessaryto provide
adequateproteinforinfantsbecauseof thelackofsuitableweaningfoodsin the
nomadichuntingand gatheringdiet.This problemofproteinsupplycontinues
intothesecondand thirdyearoflifeamongthe Kungand is emphasisedby the
severe growth retardationobserved in post-weaningchildren(Truswell &
Hansen I976). !Kung infantsgrow at ratescomparableto westernchildrenfor
thefirstsix months,nourishedon breastmilk,but afterthisdrop to below the
3rd percentileof the height/weight standardfor age of Americanchildren
(Truswell& Hansen I976; Lee ig80). Between6 and 12 yearsof age theselow
weight tendenciesare more pronounced,with !Kung childrenattainingan
average weight of only 63 per cent. of the soth percentileaccordingto the
Harvard standard(Truswell & Hansen I976). This resultsin adults who are
shortand light,with male and femalefatlevels only 43 and 6i per cent. of
Westernstandardsrespectively (Lee I979). The adultweightoffemalesaverages
about 42 kg. (Lee I979) which is close to the low end of the range Frisch
postulatesas necessaryto achievereproductivestatus(FrischI975). Children
who live at thecattlestationswithpotentialweaningsupplementsaretallerand
heavierfor theirage to adulthood (Lee I979). Thereforeit seems clear that
periodsof proteinor proteincalorieshortageare presentduringthe!Kung life
cycleand may have an effecton theexceptionallylong birthintervalnotedfor
thisgroup.
Overall, then, the literatureon modern humans suggests that interbirth
intervalsaregovernedbya complexsetoffactors,butseldomextendbeyond36
months.The exceptionsnotedincludejainand Bongaarts(i98i) studyindicat-
ing 38.2 monthsin India,38.4 monthsin Indonesia,37.9 monthsin Sri Lanka,
and Lee's (I972) (Howell I976b; I979) reportsuggesting44 monthsfor the
!Kung in theirnomadicphase. These maximuminterbirth periodsarestillonly
two thirdsthelengthofthosereportedforwild apes.
To summarise,the factorsaffecting the lengthof the interbirth intervalin
humansincludelevel of body fat,maternaldiet,maternalactivity(subsistence
patterns),lengthof lactation,patternof nursing,intensityof sucklingand
presenceof weaningfoodsin theenvironment. There are otherfactorssuch as
 A. C. ZELLER 54'

maternalage whichaccountforintra-individualvariance,butarenotconsidered
here.Level offood supplyand itseffecton maternalactivityare startingpoints
forexaminingthevalue ofchildren'scontributionsto theirparents.

Contributions
ofchildren

Pastoralandpeasantcultures. Data on theeconomiccontribution of childrenare

includedin some studiesof peasantand pastoralsocieties.Care of infantsand
livestockis a frequentformofcontributive Althoughtheseactivitiesmay
effort.
notprovidedirectsubsistencegain theyfreeadultsforsuch activityand can be
classedas a valuableindirectcontribution (WhiteI975). Borgerhoff-Mulder &
Milton(I985) examinedpatternsofinfantcare,and therefore patternsofsibling
involvementin caretakingamong theAfricanKipsigis.They notedthatfairly
youngchildren(medianage 6.3 years)assumedhighlevelsofresponsibility for
infants.In Munroe etal.'s studyof fourcultures(I984), childrenspent23 per
cent. of theirtimeworking.This figurewould have been only 7 per cent. if
infantcareand domesticchoreshad notbeenincludedas contributions (Munroe
et al. I984). Indeed, child care is mentionedspecificallyas a contributionby
childrenamongtheGusiiofKenya(LeVine& LeVine I966), theRajputsofIndia
(Minturn& HitchcockI966), the Mixtecansof Mexico (Romney& Romney
I966) andtheTarongofthePhilippines(Nydegger& NydeggerI966) whichare
all peasant level societies. Other typesof helpfulactivitiessuch as running
errands,caring for livestock and domestic chores are also mentionedfor
childrenfrom3 to IO yearsof age. Child care and animalcare are particularly
notedas economiccontributions byJavaneseand Nepalese childrenstudiedby
Nag etal. (I978). Six- to eight-year old childrenspentan averageof3.5 hrs.per
day in work activitiesin Javanesevillages and 4.3 hrs. per day in Nepalese
villages.Nine- to eleven-year-old Javaneseand Nepalese childrenspentfrom
3. I to 6.5 hrs.perdayin contributive effort.The hoursspentperday continued
to riseas theages increasedup to adulthood(Nag et al. I978). These studiesall
indicatethat childrenin agriculturally orientedsocietieshave an economic
importanceto theirparents,whetheror nottheparentsperceivetheactivitiesas
beingveryuseful(McDowell 198I).

Hunter-gathererandhorticultural The materialfora surveyof children's

societies.
contributions inhunter-gathererandhorticultural societiesis notreportedin the
literatureforverymanysocieties.Irons(I983) commentsthatchildrenin these
subsistencepatternsarean economicburdenuntilaboutage I2 to I 5 whenthey
begin to become economic producers.Since I agree with White (I975) (also
Munroe etal. I985; Nag etal. I978) thatthecontributive activitiesof children
could bothdirectlyand indirectlyreducetheeconomicdemandsmade on their
parents,I interviewedelevenfieldresearchers who providedfirsthandinforma-
tionon thirteen cultureswithhunting-gathering and limitedhorticultural bases
(table3). The !Kung areoftenconsidereda prototypical hunter-gatherer group
and no first-hand interviewswere available,so I used 2 publishedsources,one
542 A. C. ZELLER

fromthe fieldworker'sperspective(Draper I976) and the othera reportof

childhood memories(Shostak I976), bringingthe total to I4 cultures.The
surveydealtwitheightmajorclassifications ofbehaviour,each subdividedinto
variousactivitiesconstituting eitherdirector indirectcontributions.The activi-
tieslistedarethosebywhichchildrencouldcontribute to theirown maintenance
(direct)or assisttheirmothers(indirect)to work moreefficiently. Table 4 lists
theactivitieswithparticipation dividedbyage intothreestages,3-5 years(stage
i), 6-Io years(stage2) and I I-I 5 years(stage3). The materialis notdividedby
gendersincein mostcases bothsexes contributeassistance,althoughin several
casesitwas notedthatgirlshelpedtheirmothersmorethanboys. The age offirst
contributionis indicatedbut activitiesnoted at one age are usuallyfoundin
subsequentage categories.
Childrenof theyoungeststage (age 3-5) made theirmajor contributions at
theindirectlevelofchildcare,carrying objectsand runningerrands(table5). In
many cases theyused tools, althoughat a fairlylow level; nevertheless, they
were being given a veryearlyopportunityto familiarisethemselveswith the
technologicalaspectsof theirculture.Knives, axes, fire,and carrynetswere
used by childrenat ages whichprovokedcommentby theinformants. 'Whena
motherneededherknifeshetookitfromtheyoungchild,who was playingwith
it,used it,and gave it backto him' (Rodman,personalcommunication).Snares
and digging stickswere put to use by four-yearolds, who rapidlygained
proficiency withthem.In manycasestheyareconsideredreasonablycompetent
aroundfire,and among theCree are sometimesdetailedto watchtheroasting
meatand turnit as necessary.Contributions to gardeningwerenotveryhighat
thisstage,probablydue to lack of strength,but the conceptsof plantingand
harvestingwere introducedat an early age in those cultureswhich utilised
gardens.
The majorityof contributionsin both directand indirectactivitieswere

TABLE 3. tosurvey
Contributors ofchildhood
activity
patterns.
Informant Location Tribename Subsistence

ElspethYoung P.N.G. Highland Garden

ElspethYoung CentralAustralia Wabwi H&G
Nancy McDowell P.N.G. Bun Fish/Sago
Pat Townsend P.N.G. Saniyo-Hiyowe H & G, Sago
JaneGoodale N. Australia Tlwi H& G
JaneGoodale S.W. New Britain Kaulong H & G, Garden
MargaretRodman Vanuatu Vanuatu Garden
Naomi Scaletta W. New Britain Kabana Garden,Fish
DorothyCounts W. New Britain Kaliai Garden,H & G
LeslieMarshall Truk Namaluk Garden,Fish
Sara Preston N. Ontario Cree H& G
JackieSolway Kalahari Bakgalagadi Pastoral,H & G
Mike Lambeck Comoro Islands Mayotte Swidden,Fish
P. Draper* Kalahari !Kung H& G
M. Shostak** Kalahari !Kung H&G

*Draper,I976
** Shostak,
1976
I wouldliketoextend
mythanks
totheeleveninformants
whoprovided onwhichthissurvey
thematerial was
based.
 A. C. ZELLER 543

TABLE 4. Childhood activitypatterns-subsistence related number of groups.

3-5 years 6-1o years 11-15 years Total

Indirect
contribution
Childcare-start6-io months
carry I 9 IO
watch 5 9 I4
protect 4 8 I2
feed I 4 I 6
cook for I 5 I 7
Carry
tools 6 2 I 9
water I 7 I 9
firewood 2 7 I IO
messages(fire) 5 I 6
runerrands 6 I 7
food (produce) 3 4 7
buildingmaterial 2 2 4
walk long distance(miles) 3 3
Householdhelp
sweep 6 6
wash 8 8
Garden
weed I 3 2 6
fixfences 2 2
plant I 3 4
harvest I 3 4
prepareground 2 2
own I 2 3
Carefor
grandparent 5 I 6
new mother 2 2
livestock 4 4
chickenls I 2 3

Directcontribution
Use tools
fire 3 5 8
knives 4 4 8
axes 3 4 7
diggingsticks 2 3 5
carrynets 3 2 5
slingshots I 2 3
gun,bow, spear I 3 4
netfish 2 2 4
snare I 3 4
canoe 2 3 5
Catchandgather
birdsand eggs 2 7 9
smallanimals 3 5 8
grubs 2 5 7
shellfish 4 4
plants 3 6 9
fish(spear,hook) 2 2 4
hunt-spear, gun I 3 4
netfish 3 3
own protein I 4 5
Preparefood
helpa little 3 2 I 6
wash, grate,pound,peel I 3 2 6
cook forfamily 3 2 5
huskand gratecoconut I 3 2 6
 544 A. C. ZELLER

initiatedbetween6 and io yearsof age (Stage 2, table4). On a proportional

basis, childcare was themostimportantactivitycarriedout by children,and
rangedfromplayingwiththebabyto caringforhimor heroverthecourseofa
dayor evenovernight(LambeckI985). This would relievethemothersofmuch
of thestrainof continuouslycaringfora one- to three-year-old sinceshe could
forageor garden,undistracted by a clinginginfantwhom shewould nothaveto
carryand nurseall day. Otherindirectcontributions includerunningerrands
and carrying firewood,water,produceand messages.Veryyoungchildren(age
3) maystartwithone or two sticksofwood, or yamsin a carrynet,butby age 8
theyarecarrying fivelitresofwaterwhiletheirmotherscarrytwenty(Solway).
Constantpracticefroma youngage buildsup boththestrength and techniques
necessaryto transportvarioustypesof loads. At thispoint, carryingone- to
two-year-oldsiblingsmay occur forsubstantialportionsof the day. Among
CentralAustralians,femalesunderage I 5 performed38 per cent.of all infant
transport(Denham I974). In addition,help with household chores such as
washingclothesand dishes,sweepingand mending,is frequently expected,as
can be seenfromtable4.
One specialisedaspectof domesticwork, which can be regardedas both a
directand indirectcontribution, is thetimeand energyconsumingpreparation
ofcoconuts,taro,seedsandnuts.By six to tenyearsofage childrenwash,grate,
pound,peel and cook foodon a moreor lessregularbasis. In some casesyoung
childrenare lefthome in the chargeof a ten-yearold who caresforthemand
preparesthefamilymeal. In thissituationtherearealmostalwaysotheradultsin
thevicinityin case ofemergency.Even unweanedchildrenareleftsincetheycan
be nursedmorningand night,and fedmashedbananasor theequivalentduring
theday.
The abilityto undertakehouseholdmaintenanceactivitiesalso made children
usefulto adultsotherthantheirparents.In factit was specificallymentionedby
informants thatchildrenarefrequently adoptedbyindividualswho do nothave
them,such as grandparents, forboth theirlabour and companionship.These
children,who rangein age from5 to IO, contribute to theproductiveprocessby
gardening,carryingwater and wood, preparingfood and washing clothes.
LambecknotesthatamongtheMayotteone oftheprimaryreasonsfortakingin
childrenis to have a helpinghand(LambeckI985: 9). Young mothersmaytake

TABLE 5. indescending
Categories orderofproportional
contribution.
Categories Stage1 Stage2 Stage3 Total *Proportion

Child care 12 35 2 49 9.8

Carry 23 25 7 55 6.8
Household help 0 14 o 14 7.0
Catch and gather II 34 8 53 5.8
Preparefood 5 II 7 23 5.7
Use tools 17 28 5 50 5.0
Garden 4 15 2 21 4. 2
Carefor I 13 I 15 4.2

* Proportion
refers
tothetotalnumber ina category
ofcontributions divided
bythenumber
ofactivities
inthat
category.
 A. C. ZELLER 545

in a nieceor youngsiblingto assistthemwiththeworkinvolvedin caringfora

new baby. AmongtheCree, olderpeoplewho stillwishto live alone areable to
managetheirhouseholdchoreswiththehelpofa five-or six-yearold, who can
cook, carrywood and clean fish (Preston). It is quite possible that having
childrenwho arecapableofundertaking theseactivitiesforolderkinwillrelieve
mothersof tasks which they would otherwisebe socially constrainedto
perform.Animalcareis includedin thiscategorybecause it involvesa certain
amountofresponsibility, althoughofa different type.Since,however,mostof
thesecultureswere surveyedon thebasis of a hunter/gatherer or horticultural
subsistencebase, onlya fewgroupshad livestockor poultryto be caredfor.As
mentionedabove animal care is oftena major contributionof childrenin
pastoral(AinsworthI985; Hassan I980) or farmingcommunities(White1975).
This age categoryhas also begundirectsubsistencecontribution by collecting
birdsand eggs, smallanimals,grubs,shellfish, plantfoodsand by nettingfish
whereavailable.They may not accompanytheirmothersto do this,but may
formpartiesofpeersandforageon theirown. Young AustralianAboriginegirls
begindiggingforlizardsataboutage 4 and areroutinely successfulat thisskilled
and energeticactivityby age 8 to io (weightoflizard:2 to 4 lb, ElspethYoung).
Childrendo not alwayssharefoodwithadults.Frequentlytheymakefiresand
cook it in the bush duringthe day (Rodman). In some cases thisprovidesa
substantialportion of their proteinintake for the day (Chowning I972).
Laughlin commentsthatAlaskan Eskimo '(c)hildren. . . actuallyproduce a
greatpercentageof theirown food supplyin the subarcticzone-including,
especially,shellfish'(i968b: 241-2). There are a numberof casual reportson
childrencatchingbirds,rats,grubsandinsects,whichtheyeithereatthemselves
or sharewiththeirmothersand siblings(Dwyer I974). The nutritional value of
thispatternof searchingforand eatingfood duringthe course of the day is
confirmed byRobson and Yen's (I980) reportfortheTasaday. They discovered
that, based on the amount of food broughtback to the home cave, daily
on-the-spotconsumptionofsmallitemsin theforestmustprovidealmost2/3of
theTasaday's dailyintakeof energyand protein.Childrenmay also undertake
time-consuming taskssuch as berrypicking,grassseed collecting,or digging
forgrubs-which theydo bringback to thefamilyunit.Girlsmayaccompany
theirmothersonto thereefto gathershellfishand boys will oftenaccompany
teenagerson a spearfishingexpedition(Scaletta,personalcommunication).
There are only a few culturesin which contributive effortis initiatedat the
eleven-to fifteen-year-age category.Thereare some typesof assistancesuchas
cookingforyoungersiblingsor huntingfairlydangerousanimalswithspears
and firearms whichare morecommonlyadded to thebehaviouralrepertoire at
thisage. Heavy gardenworksuchas fencing,preparinggroundand harvesting
sago may be leftuntilthisage also, althoughweedingand harvestingsmaller
food may begin earlier.By age I2 some Huntingand Gatheringchildrenare
potentiallyself-supporting in a subsistencesense (Tiwi and Cree) whereas
others,suchas the!Kung,arenotexpectedto be. Goodale mentionedthatsome
Tiwi childrensubsistedforseveralmonthsin thebush afterthedeathof all the
adultsin theirgroup(personalcommunication).Solway saysoftheBakgalagadi
thatby the time a girl is I2 she can run a household,and beforethis she is
546 A. C. ZELLER

expectedto putin a fulldayofwork.AmongtheKaliai girlsage I 3 canmanagea

gardenand are capable of makingcopra forsale. Thus it is not so much the
in subsistencebase thataccountforvariationsin childproductivity,
differences
in whatis expectedin theculturalpatternand whatis expectedof
as differences
children.This pointis supportedbyBorgerhoff-Mulder & Milton'sresearchon
thesocialandeconomicfactorswhichappearto be correlatedwithhighlevelsof
siblinginvolvementin childcare. They foundthatfamilystructure, patternof
residenceand divisionof labour,contributeto differencesbetweenculturesin
thepatternsofinfantcaretaking(i985).

Levelsofcontribution.The fourteengroupssurveyedshow greatvariationin the

numbersof activitiesby which childrencontributeto theirmaintenance.The
eightcategoriesof contribution are subdividedintoforty-eight areasin which
thereis potentialforchildhoodinvolvement.In twelveof thegroupsassistance
occurs in fromten to thirty-eight activities,while in the othertwo cultures
minimalamountsofhelpinthreeareasarenoted(table6). In theI 2 cultureswith
moderateto highlevelsof childhoodcontributions, a totalof 270 activitiesare
undertaken witha meanof22.5 activitiespercultureout ofa possibletotalof48.
The averageparticipation is actuallymuchhigherthanit appearsbecause some
of these activitiesare not presentin all cultures.For example, the Central
Australians,Kung and Bakgalagadido nothavefishingtoolsor activityas part
oftheirculture,andin some groupsgardeningis notpartofthesubsistencebase
(e.g. Tiwi, CentralAustralia,Cree). Thus ofthe48 potentialcategories,thefour
groupsin whichchildrenundertake30 or moreactivitiesshow a veryhighlevel
of childhoodinvolvementin theoverallculturalpattern.
At theotherend of thescale are thetwo culturesin thissurveywhichshow
negligiblelevelsof inputfromchildren.These are theSaniyo-Hiyoweand the
Kaulong, bothofPapua New Guinea.In bothcasestheinformants commented
thatlevels of contributionby childrenwere verylow and thiscorrelatedwith
low levelsofcompletedfertility. Among theSaniyo-Hiyowe,sago provides85
per centof thecaloriesutilisedwhile meatsauces, made fromferalpig, small
game, fishand insectlarvaaccountforio percent.The remaining5 percent.is

TABLE6. Contribution
levelsofchildren
indescending
order.
Stagei Stage2 Stage3 Total

Kabana 5 22 II 38
Kaliai I0 22 2 34
Vanuatu 5 26 I 32
Cree 10 17 3 30
Mayotte I 19 I 21
Numluk 8 8 3 19
Bakgalagadi 2 15 2 I9
CentralAustralian I5 3 0 I8
PNG 13 4 0 17
Tiwi 0 15 2 17
!Kung o 8 7 15
Bun 2 7 0 9
Sanlyo-Hiyowe 2 I 0 3
Kaulong 0 I 2 3
 A. C. ZELLER 547

composedofotherplantfoods.Enoughsago canbe gatheredquiteefficiently by

one woman to lastforseveralweeks;itis, however,a veryhighstarchdiet,and
people mustsupplementitwiththeotherdietarynecessitieswhichare gathered
from the forest.This is especiallyimportantfor young childrenwho are
undergoingthetransition fromnursingto adultdiet.Sago is noteasilydigested
and a numberof one- to three-year old childrenshow symptomssuch as soft,
discolouredhair and listlessness,which are associatedwith proteinshortage
(Townsend I980). This problemof feedingyoungweanlingchildrentendsto
prolong the nursingperiod, and as a result post partum sex taboos and
infanticide areused to maintainbirthspacingat about threeyears.Townsend's
(I980) studyindicatesthatthemeannumberof live birthswas 5.3, withinfant
andearlychildhoodmortality of43.2 percent.Itis an interesting
pointthat57 of
I32 childrendied beforeweaning,and only fivedied after.This may relateto
dietaryproblemsin theprolongednursingperiodas wellas thehighincidenceof
diseaseslethalto thesechildren.In mostcultureswithhighinfantmortality the
greateststressoccursin thepost weaningperiod (Lancaster& LancasterI983;
Lancaster& King I985; Lee I972; I980; Ripley I980). Eleven per cent. of all
childrenbornaresubjectto infanticide whichis thecauseof 23 percent.ofinfant
and earlychildhooddeaths. In almost all cases the explicitreason given for
infanticide is birthspacing.It seemsto weigh moreheavilyon girlsthanboys,
sinceeightof elevenvictimswere female(Townsend I980). The demographic
consequencesof thispracticeis a populationwhichis barelyreplacingitselfin
termsoffemales(grossreproductive rate,Howell I 979). The forty-ninewomen
ofcompletedfertility in thisgroupproducedforty-nine daughterswho maynot
all surviveand reproduce(Townsend I985). Thus, althoughthesewomenhad a
mean birthrateof 5.3, only two or threesurvivedto reproductiveage, witha
bias in favourof males.This is about thesame levelof successas ape reproduc-
tion.The correlationbetweenlevelsof contribution by childrenand reproduc-
tive success are maintainedin thisculturesince offspring engage in veryfew
contributive activities.Girlsdo nothelpwithchildcarebeyondplayingwiththe
infantfora few hours,althoughtheydo help theirmothersgatherfirewood.
These activitiesare the only ones mentionedduring the data collection
survey.
Another group with low levels of population replacement(about four
offspring) and minimalhelpfromchildrenaretheKaulong of SW New Britain
(Goodale). These people acquireapproximately 6o per cent.of theirfood from
thebush and about 40 per cent.fromgardening.Birthspacingof fouryearsis
maintainedbyprolongednursingandsexualabstinence.Fouryearsofnursingis
consideredessentialforthecontinuedlifeandhealthofyoungchildrenand poor
healthis a majorcauseofinfantmortality. The oldersiblingmustbe able to walk
independently to the gardenarea beforea new infantwill be allowed to live.
Mothershave totalcareoftheinfants,whichincludesholdingthemat all times,
even when the infantsare asleep. Afterage 3 infantcare is sharedby older
siblings,buttheyarenotleftin a highlyresponsiblesituation.Whenthefamily
goes to thegardenchildrenpickup snailsand nibbleon variousfoodsduringthe
course of the day. Girlsbegin helpingin the gardenearlierthanboys but all
childrenare stillquite dependentuntiltheyare in theirmid-teens.Mothersdo
548 A. C. ZELLER

mostof thewood, waterand foodcarryingas well as cookingforchildren,but

menand teenagersmustcook forthemselves.
These two cases of low levels of populationreplacementcoupled withlow
childhood productivityare in contrastto the other culturessurveyed.The
!Kung appearto be an anomalouscasein thatfieldresearchers reporta verylow
level of childinvolvementin contributiveactivitiesalthoughfifteenareas of
activityarementionedin theirreports.These oftenoccur,however,underadult
supervisionand at an olderstageof childhood(stage3) thanis thecase in many
other groups. Anotherfactoraccountingfor this discrepancymay be the
amountof timeinvolved. Solway (personalcommunication)commentsthat
Bakgalagadichildrenarecontributing severalhoursperdaybythetimetheyare
6 to 8 yearsold and mostof theday by age io. Draper's (I976) studyof !Kung
childrenfindsthattheycontributedforonly a few (I-5) minutesin a sample
hour. Lee (I979) mentionsthat!Kung childrenarenotpressedintothesubsist-
ence questuntiltheyare teenagers,althoughtheyrunoccasionalerrandsat an
earlierage. The correlationbetweenlow levelsof contribution by childrenand
low birthrateshowever,agreeswiththehypothesis proposedin thisarticle.The
!Kung have an averageof4.7 to 5 childrenwitha loss rateof about 34 percent.
(Howell I976b). Thus theirrateof populationgrowthin nomadic groups is
quitelow. However, the !Kung livingsituationis undergoingrapidchangeas
andthebirth
theybecomesedentary interval
decreases
(Lee I972; I979; I980).

byapeandAustralopithecine
Contributions children

Ape pattern.The major nonhumanmodel of Australopithecine behaviouris

based on the informationderivedfromstudyingchimpanzees,as a highly
intelligent,tool using, social, closelyrelatedprimate(Tanner I98I; Zihlman
I98I). Chimpanzee infantcare differsfrom the human patternin several
significantways. In contrastto the human situation,ape mother-offspring
relationsare muchmoreheavilyslantedtowardsthemotheras provider.After
chimpanzeefemalesbear theiryoung, theynurseand carrythemextensively
untiltheirthirdyear.Betweenage 4 and 5, finalweaningoccurs,but mothers
stillcarrytheiryoungiftheytravela long distance.The younglearnfoodhabits
by watchingtheirmotherspick fruit,or fishfor termites,and occasionally
motherssharefoodwiththem,or allow themto takeit(Goodall I 979). The next
siblingis usuallybornwhentheyoungster is initssixthyear,butin chimpanzees
and gorillasit is stilldependenton themotherforemotionalsecurityand some
care.Juvenilesfrequently sharethe mother'snightnestwithher and thenew
infant.Not untilthenew infantis about i yearold and thejuvenileabout7 will
the motherallow thejuvenile to carryit, and even thenonly forshorttime
periodsand undersupervision.Apes do not seem to utilisethemonkeypattern
of fairlyextensiveallomotheringand baby tendingwhich can serveto give a
monkey mothera respitefromher infants.Usually juvenile apes begin to
wanderfromtheirmothersforincreasedlengthsof timeat about age 7 or 8, as
theymatureinto adolescents.It is not untiltheyhave achieved thislevel of
 A. C. ZELLER 549

emotional independenceand knowledge of the environmentthat they are

capableof successfully survivingon theirown or adoptingan orphanedsibling
(Telekietal. 1976). Youngeranimalshavebeenobservedattempting to serveas
surrogatemothers,buttheyarerarelysuccessfulin keepingtheorphanedinfant
alive.
Motherchimpanzeesmustnursetheiryoung duringthe timethattheyare
becomingfamiliarwith, and capable of exploiting,the naturalfood sources.
The shiftto solid food takeslongerfor chimpanzeeyoung thanforhumans
becausetheymustbe ableto climbtreesto getatthefruit,crackthenuts,pullup
vegetation,and operatea termitestickthemselvesbeforetheyarenutritionally
independent.All these operationsrequire both learningand some level of
strength or dexterityto achieve.Theirmotherswillsharesome foodwiththem,
but not enoughto nourishthemiflactationwere to cease when theinfantwas
stillquiteyoung.Sincejuvenilesdo notcontribute to theirmother'sor younger
sibling'sdiet,and do notrelievethemotherofanysubstantial childcareburden,
theyareofverylittlehelpin raisingthenextsibling.In fact,theystilldependon
themotherfora considerableamountof social and psychologicalsupport,and
sometimesfortransportation.

patternand itsinfluence
Australopithecine on hominisation. Clearly, many of the
differencesbetweenAustralopithecine and chimpanzeelifestyle areresultsofthe
processofhominisation, butwhenand wherethesebehaviourswereinitiatedin
hominiddevelopmentis notyetevident.Fourmajorchangesin lifestyle would
contributeto thedevelopmentofthedifferences in childcarepatterns.The first
developmentconsideredis thetransition fromforagingto gathering,coupled
withthedevelopmentofsimpletoolsandcarrying aides. Secondlytheestablish-
ment of group life and the division of labour would be fosteredby the
developmentoflanguageoutofa moregeneralisedcommunication system.Itis
notknownwhenculturalpreparation offooddeveloped,butthischangemight
have markedeffectson the survivalrate of the young, especiallyin crisis
situations.Fourthly,a new locus of resourceexploitationutilisedby hominids
may have providedan advantageover thepreviousape-likepattern.There are
roleswhichimmatureoffspring could fillin all of thesedevelopments.

Gathering The habitof gatheringfoodas opposed to merelyforaging

vforaging.
foritis a majormilestoneinhumandevelopment.Itis highlycorrelatedwiththe
earlierweaningpracticesof humans(RipleyI980). All thegroupssurveyedin
thisstudyhad begun thepatternof supplementalfeedingat between6 and i 8
months,whenhumaninfantsarenotable to forageforthemselves.Young apes
beginto supplementtheirmilkdietat about I2 to i 8 monthsbutmustcontinue
nursingmuchlongerbecausetheymustrelyon theirown resourcesto achieve
thissolidfoodintake.The additionaladvantageforhumangatherers is thatthey
can leave theiroffspringin someone else's care, and returnto feed them (a
patternseen in birds and carnivores,but not in Anthropoidea).The use of
childrenage 6-Io to supplementthecareof youngerchildrenwould undoubt-
edly requirethepresenceof otheradultsin thenearvicinity,and thusa group
livingpattern.Young hominidscould also contributeby performingsimple
550 A. C. ZELLER

repetitivetaskssuch as berryor seed gathering.Small edibles at lake or rivcr

edge suchas crayfish,snailsor frogscould easilybe gatheredby theyoungand
consumedon thespot or carriedback to thegroupin a container.Data on the
effectsof proteinshortagespresentedearlierin this articleemphasise that
althoughnursingchildrencan surviveon milklow in protein,theyare much
moreviable when proteinresourcesare adequate. Thus small contributions of
proteinaceousfoodsmade by childrento theirmothersmay have a nutritional
impactfarexceedingtheirsize and caloriccontent.These 'snacks'would also be
verybeneficialiftheywereconsumedon thespotby thechildren,becausethey
would relievetheparentsofpartoftheirforagingresponsibilities.
Major advantageswould accrueto younghominidsfromtheuse oftoolsand
containers.The earlyinitiationof thisdevelopmentseems probable,based on
analogous ways which otherprimatesextend theirreach, open tough food
sourcesand deal withproblemsof procurement(Goodall 1973; Tanner I98I;
Zihlman I98I). If the earlieststone tools date from2.6 millionyearsago, it
seems possiblethatwood, shelland skinmightalso have been used at least as
early.The developmentof tools would allow childrento augmenttheirnatural
strength and capacityby carryingmaterialin skinbags or ostrichegg shells.A
diggingstickwould enhancetheirabilityto gathergrubs, small burrowing
animalsor tubers.These tools are technologically verysimpleand would not
leave much tracein the fossilrecord,but it seems quite possible based on a
chimpanzeemodel of tool use (TannerI98I) thattheymayhave been available
to earlyhominids.They arecertainly withinthecapacityof modernchildrento
manipulate.

Languageandgroupliving.The developmentof language skills sufficient to

enable humans to plan and co-ordinateactivities,instructthe young, and
arrangea rendezvousis one ofthemorevaluabledevelopmentsbecauseitallows
theorganisationofgroupactivities.Evidenceforthisdevelopmentis enigmatic,
sincethetracesofitspresenceleftin thefossilrecordareso slight.Thereis some
suggestionin endocranialcastsof AustralopithsthatBrocca's area on theleft
surfaceof the brainis beginningto show increaseddevelopment(Holloway
1976). Nevertheless,the developmentof language and its role in enabling
motherand offspring to communicateis undoubtedlya major contribution to
hominidlife. Language skills would enable a motherto instructand direct
offspringin thevarioustechniquesofgathering ratherthanrelyingon thechild's
interestto profitfromobservationallearning.The motherwould also be able to
directthechild'sactivityata distance,andto givethechildinstructionsto follow
in herabsence.Languagemightalso havefosteredchangethroughtheperiodof
innovationand play characteristic of youngprimates.It could have resultedin
an expansionof thediet,ifthechildgatheredsomethingthemotherwould not
have, and offeredto shareit. Language would allow mothersto teach their
younghow to careforinfants, ratherthanrelyingon observationallearning,and
to organisethebaby-sitting process.
Grouplifehas manyadvantagesin termsofpredatorprotectionand foraging
practices.A situationsuch as thatoccurringamong theAustralianAborigines
may have developed.Here mothersand childrentraveltogetherto a protected
 A. C. ZELLER 55I

place neara good gatheringarea. Motherswithbabiesstayhereand watchover

theyoungchildren(5-7 yearolds) who are caringforthetoddlers.By thisage
young girlsare frequently carryingone- and two-year-olds(Denham 1974).
The five-to seven-year-olds playwiththetoddlers,keepthemoutofthefireand
do a littlegrub,seed or smallanimalcollectingas well. Mothersdo not like to
taketoddlerscollectingwiththembecauseofthenecessityofcarrying waterfor
them (Denham 1974). With this arrangementmotherscan leave unweaned
childrenforseveralhourswhilegatheringfood and thenreturnto nurse.Thus
one or two adultscanoverseeeightor tenchildrenaltogether withoutbecoming
exhausted,or actuallyfeeding,caringforor playingwith the children.The
motherswho areout collectingreturnto this'dinnercamp' andpreparefoodfor
themselvesand thechildren.Only a smallportionofwhatis collectedduringthe
day is actuallycarriedback to themaincamp at nightfall to be sharedwiththe
others.

Preparationoffood.A thirdhumanadvantageis theculturalcapacityto respondto

lactationalfailureby cooking,poundingand pre-chewingfood forinfantsto
digestmoreeasily.Humans have a culturalpatternwhichencouragesmothers
to providefoodfortheiryoung,bothin timesof milkshortage,and duringthe
weaning period. Among the Saniyo-Hiyowe who sometimesexperiencea
shortageof suitableinfantfood,non-motherswho findeasilydigestedgrubs,
etc., may bring them as a giftto the infant(Townsend I980). The human
patternsof processingfood may aid the youngin achievinga more complete
diet. Cooking, pounding,choppingand shellingnutsare all behaviourswhich
childrencouldundertaketo liberatecalories(Cohen 1980) andthuscontributeto
theirown and theirsiblings,dietarysuccess.Chimpanzees(and otherapes) do
notundertakethesetypesofbehaviouron theiroffspring's behalf.The bestthat
theydo is to allow themto sharein foodthattheyhavepreparedforthemselves.

Resourcelocation.A fourthdifferentiating factoris that most human food

resourcesarelocatedat or neargroundlevel. Motherswho are carryinginfants
do notusuallyhaveto climbtreesto gatherresourcessuchas nutsand fruit.The
divisionof labour is such thatchildren,men or non-mothering women will
usually obtain and share these resources.This means that neithernursing
mothersnoryoungoffspring have to negotiatetreesand runtheriskof falling,
whichis everpresentin thearborealprimates.
The valueofthesecontributions occursatbotha directand an indirectlevelof
influenceon thesubsistencebase. Ifchildrencan helpto wean themselvesearly
by eating solid food the frequencyand intensityof sucklingwill diminish,
allowingthemother'shormonalcyclingto beginagain. Slightlyolderchildren
can help to feedthemselvesand also theiryoungersiblings.They can gather
foodfordirectconsumptionandiftheymakeworthwhilecontributions to their
mother'sdiet,hervolumeofmilkmaybe increasedfora nursinginfant,and her
body storesof nutrientsnot so severelydepleted.A bettermilk supply will
promotethesuccessof thenursinginfant,while maintainingherreserveswill
allow herto become pregnantagain morequickly.Indirectcontributions such
as child care will allow mothersto gathermore efficiently,and spend fewer
552 A. C. ZELLER

caloriescarryinginfantson theirgatheringexpeditions.Leaving infantsin a

protectedcamp would also reducetheirriskof exposureto predation.In the
eventofa predatorencounterwhilegathering, motherswould onlyhaveto save
themselvesratherthanbe burdenedby a heavyhelplesschild.Assistancewith
food preparationwould allow a greatervolumeof food to be processedwitha
greaterprobabilitythattherewould be enough forall the young. The major
would be an increasein themother'senergy
influenceof all thesecontributions
levels,so thatshe could conceivemoreinfants,and moresuccessfully raisethe
ones she bore.

The interestin Australopithecine

Australopithecinefertility. patternsis
fertility
based on theirstatusas ancestralhominidforms.Presentday Homo is un-
doubtedlyvery different fromearlyhominids,but the evolutionarylinkage
should allow some extrapolationto the past. Chimpanzeesare also used as
models sincetheirbody size and foragingpatternsmay morecloselyresemble
theAustralopithecine condition,and theirancestorswerecloselyrelated.
Thus fertilitypatternsof both chimpanzee(Pan) and modernHomo can be
examinedforany indicationstheymay give concerningtheAustralopithecine
pattern.AlthoughAustralopithecine levels of fatstorageand lactationalener-
getics are unknown, the similarityin pregnancylengthsbetween Pan and
modernHomo supportan argumentthattheAustralopithecine conditionwas
not radicallydifferent. They probablyresembledwild, ratherthan captive
chimpanzeesin not gaining much more weight than the level requiredto
become fertile.Accordingto theirskeletalremainsthislevel would have been
about 30 kg. whichis just a littleless thanchimpanzees.The lengthof birth
intervalsis based partlyon theamountof storedenergywhichcan be used for
lactation.Wildchimpanzeesrequirea meanof42 months(TutinI980) to regain
aftera previousbirth.Modernwesternwomenrequireonlyabout 12 to
fertility
14 monthsto resumecyclingeven when breastfeeding infantstotally(Kippley
1974). Post-birth amenorrheadoes notusuallyexceed I 8 monthsin anyhuman
population, althoughfertility may be delayed for several months (Leridon
1977). Cycle resumptionin Australopiths probablyfellbetweenthe Pan and
Homo levels,althoughtherangeis likelyto be closerto thechimpanzeeend due
to bodysizeand activitypattern.However,dietarysupplementation ofmothers
by kin, mate and possiblychildrencould operateto reducethe lengthof the
infertile
periodthroughthemechanismsmentionedin thisarticle.This change,
coupled with a greaterchance of survivalforpost-weanlingchildren,could
increasethenumberofoffspring successfully rearedby a mother.
Several authorshave investigatedthe possibilitythatfemalescould act to
increasetheirreproductive successby chosingmateswho would helpthemrear
theiroffspring (Hrdy& WilliamsI983; Irons1979; I983; TriversI972). Others
have argued that Australopithecusmay have had a more matrifocalliving
situationand utilisedaid frommaternalkinwho would be morecloselyrelated
to themthanweretheirmates(FischerI982; TannerI98I). This articlesuggests
thatoffspring who are equallyrelatedto themother,no matterwho she mates
with,may have contributedtheirefforts, both at maintainingthemselvesand
theirsiblingsand increasingtheirmother'spotentialforadditionaloffspring.
 A. C. ZELLER 553

Irons comments(I983) thata largersibshipmay have been advantageousto

earlyhominidsfortheformation ofalliancesas well as thespreadofpopulation,
as long as thespacingwas not so close as tojeopardiseindividuals.Successful
femaleAustralopithecines could benefitthemselvesby choosinga mate who
would help rearoffspring, as well as by havingchildrenwho would help both
themselvesand her. If the offspring were socialisedto assisteach other,they
would be in an advantageouspositionwithrespectto othersibshipswho didnot
operate in that fashion.If this behaviouraltraditionbecame common to a
majorityof thepopulationit could increasethenumbersof Australopithecines
and permitthemto expandfromtheiroriginalhabitat.

Conclusions
The datapresentedin thisarticlesupportan argumentthatsubsistenceactivities
ofhominidchildrencan directlyorindirectly save sufficientenergycostsforthe
motherto influenceherreproductive rateby shortening theinterbirthinterval.
Studies of greatape reproductionindicatethattheirapproximatelysix-year
birthintervalis barelysufficientto maintaintheirpopulationlevel. Fertilitydata
suggestthatnutritional factorsare a majorconstraint on humanreproduction,
especiallyin termsof energyexpendedforcaloriesgained.This is particularly
relevantafterthebabyhas beenbornand themotheris maintaining bothherself
and thechild.The presenceofweaningfoodsofsuitablequalityand consistency
are a major factoraffecting both survivalof childrenand the frequencywith
which they are produced. Length of nursing,frequencyof infanticideand
voluntarychildspacingareall interrelated determinants of overallfertility.
The actual contributionsmade by childrento the subsistencebase are
discussedin fourteenhunter/gatherer and partiallyhorticultural subsistence
systems.Direct contributions includeprovidingsome of theirown food from
an earlyage whichwill reducetheload on theirmother.This contribution will
be even more effective if older childrencontributeto theiryoungersiblings'
food supply. Childrenmay sharesmall but dietarilysignificant typesof food
whichtheyhave timeto collect,suchas berries,insectsand invertebrates, with
theirmothersand siblings.Moving to a moreindirectlevel childrenmay help
theirmothersgatherwood andwater,workin thegardensand perhapsdo some
housework.They can also relievetheirmothersof muchof theburdenof daily
childcarefortoddlersand youngchildren.All theseactivitiesallow themother
to conductherown subsistenceworkmoreefficiently in termsoffoodcollected
and caloriesexpended.
In his surveyof the economic value of childrenin JavanesecultureWhite
(1975) pointsout thatindirect contributionsarejustas valuableas directones in
freeingtheparentsto performvitaladultactivities.The mostvitalbehaviourin
an evolutionarysenseis theproductionand rearingof offspring. As thisarticle
indicatesthereare a greatmanyfactorsinvolvedin successfullyachievingthis
undertaking, even afterthemale and femalehave choseneach otherand begun
thematingprocess.In additionto themale-femalebond are levelsof maternal
diet, maternalactivitypatterns,lengthof lactationalperiod, infantdiet, and
successin defendingtheyoung,whichareall majorvariablesin determining the
numbersof offspring who can be raisedto maturity.These factorsaffectnot
554 A. C. ZELLER

onlypresent-day populationsbutalso earlierhominidswho had different

types
of resourcesbut thesame typesof problemsto face.The data presentedin this
articlesupport the suggestionthat one resource was their offspringwho
labouredwiththeirparentsto maximisethesize of theirsibshipand establish
hominidsas themajorprimatetaxon.

NOTE

This is a revisedand expandedversionof a paperpresentedto theCanadian EthnologicalSociety

Meetingsin Toronto,May i985. The paperis dedicatedto thememoryofmyfriendand colleague,
Dr MelissaJ.Knauer,who diedin a tragicaccidenton herway to conductfieldworkwiththe!Kung
inJulyi985.

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