THEHUMANCAREER

ByRichardKlein

Thenounphylogenyanditsadjectivephylogeneticareoftenusedassynonymsforevolution
andevolutionary.However,strictlyspeakingaphylogenyisatreediagramillustratingthe
evolutionaryhistoryandrelationshipsofaspeciesorspeciesgroup.Itisthustoaspecies
whatagenealogyorfamilytreeistoanindividual.
ThezoologicalorderPrimatesincludes230livingspeciesclassifiedinformallyaslemurs,
tarsiers,monkeys,apes,andhumans.Manyadditionalprimatespeciesareknownonlyin
fossil form, and many more probably await discovery. Knowledge of living and fossil
nonhumanprimatesiscrucialtoanunderstandingofhumanevolution.
Intheclassificationschemeemployed,thelivingAnthropoideaorhigherprimatesbelongto
threesuperfamiliestheHominoidea,comprisingpeopleandapes;theCercopithecoidea,
comprisingtheAfroEurasianorOldWorldmonkeys;andtheCeboidea,comprisingthe
SouthandCentralAmericanorNewWorldmonkeys.
AlandconnectiondevelopedbetweenAfricaandEurasiasometimebetween20and17Ma,
itbecamepermanentabout15Ma,anditallowedhumanoidstospreadfromAfricainto
broadleavedwoodlandsthatstretchedacrosssouthernEurasiafromSpainonthewestto
Chinaontheeast.TheyflourishedinsouthernEurasiauntil87Ma,whenadverseclimatic
changeeliminatedthemfromEuropeandreducedtheirAsiandistributiontotheextreme
southeast,includingsouthernChina.
THEAUSTRALOPHITSANDHOMOHABILIS:In1939,GregoryandHellman(1939)
proposed that ancient primitive human fossils from South Africa be allocated to the
subfamily Australophithecinae, in distinction from a second subfamily, the Homininae,
containing later, more advanced humans. The Australopithecinae and the Homininae

together comprised the family Hominidae separate from the family Pongidae, which
includedthegreatapes.
Theaustralopithsaretheoldestknownundoubtedhominis,withafossilrecordthat
extendsfrombefore4.4millionyearsago(Ma)toatleast1.2Ma.Inthevernacular,they
havesometimesbeencalledmanapesbecausetheycombinedbipedallocomotionthe
definingcharacteristicofthehominintribewithsmall,apesizedbrainsandapelikeskulls
andfaces.Itisbecomingincreasinglyclearthatmostifnotallaustralopithsresembledapes
inotherrespects,includinganapelikeupperbodyformwithlongpowerfularms,andthat
theyprobablymixedhumanstylebipedallocomotiononthegroundwithapestyleagilityin
thetrees.Inshort,morphologyandbehaviourtogethersuggestthattheaustralopithswere
bipedalapesmorethanprimitivehumans.Humansinthenarrowsenseassignedtothe
genusHomofortheirlargerbrains,theirsmallerfaces,andothersignificantdeparturesfrom
theapeconditionappearedonlyabout2.5Ma,anditisprobablynocoincidencethatthe
oldeststonetoolsdatefromaboutthesametime.
Based on the presumed rate at which living humans and chimpanzees diverged
energetically,theirlastsharedancestorlivedbetween8and5Ma.Theputativehominin
fossils dated to roughly 6 Ma in Chad, Ethiopia, and Kenya may confirm this, but if
unambiguousevidenceforbipedalism,theoldestunquestionablehomininspeciespostdate
4.5Ma.
Morethanahundredyearsago,ThomasHuxleyand,later,Darwinpostulatedthat
humans originatedinAfrica, sinceitwas thehome oftheir closestlivingrelatives, the
chimpanzeeandthegorilla.TheaccumulateddataamplysupportHuxleysandDarwins
prescience.TheknownAfricanearlyhomininsitesareconcentratedintwoareasinthe
northeasternportionofSouthAfricaandalongtheeastAfricanRiftValley,fromnorthern
TanzaniainthesouthtoeastcentralEthiopiainthenorth.
Thesuggestionthatlowlatitudevegetationalchangeselectedforbipedalismisoftenknown
asthesavannahhypothesis,sinceitassumesthatsavanna(grasslandwithscatteredtrees
or tree clumps) is the environment in which bipedalism (and other uniquely human
characteristics)wouldhaveconferredthegreatestadvantage.Savannagrasslandsofmodern
aspectdevelopedinAfricaonlyafter1.81.7Ma,whenaridityandseasonalityincreased
sharplyovermuchofthecontinent.However,fossilpollens,sediments,geochemistry,and

especiallyassociatedanimalremainsindicatethattheearliestunquestionablehominins(the
australopiths and HomoHabilis),wholivedbetweenroughly4.5and1.6Ma,variously
inhabitedwoodedgrasslandsorwoodland/grasslandmosaicsthatarebroadlyconsistentwith
thesavannahhypothesis.
Thesavannahhypothesispredictsonlythatnaturalselectionwouldhavefavoured
anatomicalandbehaviouralchangesthatenhancedsurvivalingrassier,lesswoodedsettings.
Itdoesnotexplainwhybipedalismwouldhavebeenadvantageousinmoreopenvegetation,
butthepotentialbenefitsinclude:(1)theabilitytocarrymeatorotherfooditemstotreesor
otherrefugestoothergroupmembers;(2)anenhancedabilitytopicksmalldiameterfruits
fromshorttrees;(3)areductionintheamountofskinsurfaceexposedtodirectsunlightat
midday,thusreducingthedangerofheatstress,particularlyforthebrain;(4)thefreeingof
thehandsfortooluseorforcarryingtheyounglowdistances;(5)adecreaseintheenergy
requiredtowalkatnormal(low)speeds;(6)theabilitytoseemoreclearlyorfurtheraway
duringpassagefromplace;or(7)thefacilitationofthreatdisplays thatreducedviolent
conflictsoverscarceresources.
Thevariousalternativesaredifficulttofalsify,andormorecouldhaveoperatedin
concert. The least likely is perhaps selection to reduce midday stress, since a range of
paleoenvironmentalindicatorsagreethattheearliesthominidsinhibitedrelativelywooded,
wellshadedenvironments.Itwasprobablyonlyabout1.81.7Ma,withtheadventofHomo
ergaster,thathomininsfirstinvadedthekindofarid,moreopensettingswheredaytime
foragingcouldhaveproduceddangerousheatloads.Heatadaptationplausiblyexplainswhy
H.ergaster wasthefirsthominintopossessanexternalnose,whyitwasalsothefirstto
achieveessentiallymodernbodysize,andwhyitwasdistinguishedbythesamelong,linear
bodyformthatcharacterisehistorichumansinequatorialgrasslandsandsavannas.If H.
ergasterwasspeciallyadaptedtoheat,itmayalsohavebeenthefirsthominintopossessa
largelyhairless,nakedskin.Thisisessentialtopromoteheatlossthroughsweating,andfor
creaturesthatmustremainactiveatmiddayunderdry,sunnyconditions,sweatingisthe
mosteffectivemechanismformaintainingtolerablebodyandbraintemperatures.Effective
sweating was probably crucial for the significant brain expansion that followed the
appearanceofH.ergaster.
Alinkbetweenbipedalismandtoolusemayseemunlikely,sincethefossilrecord
suggeststhatbipedalismantedatesstonetoolmanufacturebymorethan2my,but,likethe

toolsthatmodernchimpanzeesusetoprobefortermites,cracknuts,diguptubers,andso
forth,theearliesthumantoolsmayhavebeenmadeofwoodorotherhighlyperishable
materials.Inaddition,evenonesinstoneorbonemighthavebeeninsufficientlyshapedto
catchthearcheologistseye.Finallyitsimportanttorecallthattheearliestpeople(broadly
defined)probablylackedthetypicallyhumanhabitofoccupyingasinglelocaleforseveral
days(ornights),whichmeanstheymaynothaveleftstonetoolsinarcheologicallyvisible
clusters.
For decades, paleoanthropologists have often divided the genus Homo among three
successivespecies:Homohabilis,nowdatedbetweenabout2.5millionyears(Ma)and1.7
Ma;Homoerectus,nowplacedbetweenroughly1.7Maand500thousandyearsago(ka);
and Homo sapiens after 500 ka. In this view, each species was distinguished from its
predecessorprimarilybylargerbrainsizeandbydetailsofcraniofacialform,including,for
example,achangeinbraincaseshapefrommoreroundedinH.habilistomoreangularin
HomoerectustomoreroundedagaininH.sapiens.Anearlyorarchaicbeforeabout250
kawassetoffbyatendencytoretainmanyH.erectusfeatures,includingthickcranialwalls,
alow,flattenedfrontal(forehead),abroadcranialbasefromwhichthesidewallssloped
inward,andarelativelymassive,chinlessjawwithlargeteeth.Differentvariantsof H.
sapiens weresometimesdesignatedassubspecies,suchas H.sapiensneanderthalis (the
Neanderthals)betweenroughly130and4030ka.Authoritiesdebatedwhether H.erectus
evolvedinto H.sapiens inthegradualmodethatDarwinproposedorwhether H.erectus
remainedlargelyunchangedforperhapsamillionyearsandthenevolvedintoH.sapiensin
aspurtlikethosethatpunctuatetheevolutionofmanyotherspecies.
Theconventionalviewstillhasitsadherents,andoneprominentauthorityhaseven
sunkH.erectusintoH.sapiens,primarilybecausetheboundarybetweenthetwoisfuzzy.In
his view,H.sapiens originatedabout1.7Maandsubsequentlydiversifiedintoregional
formsthatmaintainedgeneticcontactandthatgraduallyevolvedintolivingpopulationsin
thesameregions.
Thischapteroutlinesthepatternofhumanbiologicalandbehaviouraldevelopment
between the appearance of H. ergaster 1.81.7 Ma and the emergence of the classic
Neanderthals(H.neanderthalis)andtheircontemporariesby130ka.Forheuristicpurposes,

allpopulationsthatexistedbetween1.8Maand130kawillbereferredtoasprimitive
Homo.
Thedifficultyofplacingmanykeyfossilsintime,combinedwiththerelativelysmall
sizeofthefossilsamples,explainswhyreasonableauthoritiescandisagreeonthecourseof
humanevolutionbetween1.8Maand130ka.Thediscussionhereaddressesgeologicageby
species,whichmeansineffectbyregion,startingwithHomoerectusintheFarEast.
The separation of Homo ergaster from Homo erectus is controversial, and even
adocatescommonlyrestrictH.ergastertoahandfulofrelativelycompleteAfricanfossils
datedtoabout1.81.4Ma.Tothese,thesamespecialistswouldprobablyaddhumanfossils
fromDmanisi,Georgia,thatmayberoughlyasold.
Intheviewthatistenatativelyacceptedhere,HomosapiensisastrictlyAfrican
offshootofH.ergaster.TheearliestspecimensaredistinguishedfromH.ergaster(andH.
erectua)byseveraladvancedfeatures,includinglargerendocranialvolume,amorerounded
(lessangulated)occipital,expandedparietals,andabroaderfrontalwithmorearched(vs.
moreshelflike)browridges.Thesefeaturesarenotexpressedequallyineveryspecimen,
perhapsinpartbecausethespecimensvaryinage.
As conceived here, Homo neanderthalis includes the classics Neanderthals, who
occupied Europe and southwestern Asia between roughly 130 and 5040 ka, and their
antecedents, who appear to have been strictly Europeans. Some of the pre (ante)
Neanderthal fossils have been dated by ESR on associated animal teeth, TL on heated
artifacts,andUseriesdisequilibriumonstratigraphicallyassociatedflowstones.Manyofthe
specimenscanbeorderedintimerelativetoothersbyassociatedmammalianfossils.In
addition,themammals,sometimessupplementedbyremainsofothervertebrates,plants,
freshwater mollusks, and even beetles, often reveal whether the climate was glacial or
interglacial.Incombinationwithnumericdatesandmammalianfossils,paleoclimatecan
then be used to correlate sites with the oxygenisotope stages of the global marine
stratigraphy.
Insum,datingandmorphologyshowthattheNeanderthalsdevelopedinEurope,
beginningasmuchas600ka,buttheavailablefossilscouldalsobeusedtoarguethatmore
primitivepopulationswithfewornoNeanderthalfeaturespersistedlocallyuntil250ka.
Alternatively,andperhapsmorelikely,thedatingandmorphologicalvariabilityimplythat
Neanderthalfeaturesaccumulatedpiecemeal,mainlyasaconsequenceofgeneticdriftin

smallisolatedpopulations,andthattheirfrequencyvariedstronglyfromplacetoplaceuntil
after250ka.Adenser,farbetterdatedfossilrecordwillbenecessarytochoosebetween
alternatives.
Amorecompleteadaptationtoterrestriallifeprobablyalsoselectedforincreased
bodysize,anditcouldevenexplainthedecreaseinsexualdimorphism.Inextanthigher
primates,strongsexualdimorphismrendtoreflectpolygynousmatingsystemsinwhich
malescompetevigorouslyforfemales,whilereducedsexualdimorphismisassociatedwith
monogamousmatingsystemsinwhichmalesandfemalespairforlongperiods.Decreased
dimorphisminH.ergastermaythussignalthebeginningsofthedistinctivelyhumanpattern
ofsharingandcooperationbetweenthesexesthatpromotedhistorichuntergatherersurvival
insavannahenvironments.
ReducedcheektoothsizeinH.ergaster/erectusprobablyreflectsagreaterreliance
ontechnology(artifacts)toprocessfood.Theremayhavebeenasimultaneousincreasein
the use ofthe incisorteeth for biting, gripping, and tearing, culminating in the heavily
shovelledupperincisorsofH.erectus.