Totem: The University of Western Ontario Journal ofAnthropology

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The Homo habilis ParadoxJoanna EberlyThe University of Western Ontario

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Recommended CitationEberly, Joanna (2007) "The Homo habilis Paradox," Totem: The University of Western Ontario Journal of Anthropology: Vol. 15: Iss. 1,Article 3.Available at: http://ir.lib.uwo.ca/totem/vol15/iss1/3

The Homo habilis Paradox

KeywordsHomo habilis, classification, subspecies, paleoanthropology

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The Homo habilis Paradox: IsHomo habilis anAustralopithecine Moonlightingas the First Migrant Out ofAfrica?

IntroductionWhen Bed I of the Olduvai

Gorge was excavated in 1959, hominidremains were recovered that wouldrevolutionize the field ofpaleoanthropology. Oldowan toolsassociated with the fossils promptedresearchers to name this new speciesHomo habilis, the 'handyman', and toposition it at the base of the humangenus (Leakey et al. 1964). However,recent articles forwarded by Wood andCollard (1999a, 1999b) challenge thishominid's traditional taxonomic positionby re-assigning it to the genusAustralopithecus. The unrelated, butequally important, discoveries of theDmanisi and Longgupo hominids propelinterpretations of Homo habilis in adifferent direction. These findingsintroduce the possibility that a group ofhabiline migrants formed the firstexpansion out of Africa (Gabunia et al.2002, Wanpo et al. 1995). Certainly theliterature presents an interestingparadox: is Homo habilis primitiveenough to be classified as anaustralopithecine, or advanced enough tocolonize Eurasia? In an endeavour toclarify Homo habilis' evolutionary role,this paper will explore recentamendments to its conventionalphylogenie position. The plausibleclassification of the Dmanisi andLonggupo remains will be discussed,along with their implications forhominid migratory models. Finally, this

paper will attempt to reconcile theHomo habilis paradox by proposingtaxonomic revisions and identifyinga new subspecies: Homo habilistransmigro.

Literary Review: Old Assumptionsand New Discoveries

Homo habilis' membershipwithin genus Homo is based upon anumber of putative features such as a600 cubic centimetre cranium, acapacity for language, precision grip,and a habitual bipedal gait (Wood &Collard 1999a: 198). Traditionally, thishominid was considered to be the basalspecies of a single lineage thatgradually evolved into anatomicallymodern Homo sapiens (Conroy2005:302). Yet in the 1970s, fossilsfound at the Koobi Fora Formation inKenya presented a major difficulty forthe long-standing notion of anagenesis.Though initially identified as earlyHomo habilis, subsequent comparativestudies exposed a great deal ofvariation between these specimens andthe Olduvai sample. The divergencesbetween the two skeletal assemblages,found particularly in the postcranialand craniodental complexes, are somarked that a new species wasestablished to accommodate some ofthe Koobi Fora hominids: Homorudolfensis (Conroy 2005:315-318).

Since the 2001 description ofKenyanthropus platyops, manypaleoanthropologists support thereassignment of Homo rudolfensis tothis autonomous evolutionary lineage(Leakey et al 2001:439, Lieberman2001:419-420, Tattersall 2003).Nevertheless, this revision does notspell the end for a 'bushier-looking'cladogram of the genus H 0 mo.Numerous Plio-Pleistocene sites

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continue to produce evidence of multiplecontemporaneous species dispersed acrossAfrica and Eurasia. Most researchers nowagree that the morphological diversityrepresented by the post-australopithecinesample is too great to be contained by asingle species (Brown et al. 2004, Lahr &Foley 2004, Tattersall 2003). Indeed, therejection of anagenetic models also callsinto question Homo habilis' comfortableposition at the base of its genus.

In 1999, Wood and Collardpresented a new interpretation of Homohabilis' morphology that removed thisspecies from the human lineage entirely.They argued that the traditional criteria foradmission into the genus Homo arearbitrary, and confuse the boundarybetween highly-evolved hominids and theancestral austra10pithecines (Wood andCollard 1999a: 199-200, 1999b:65-66).When Homo habilis was first described inthe 1960s, the minimum cranial capacityfor the genus Homo was set at 600 cubiccentimetres. The biological significance ofthis brain size is questionable, especiallysince Leakey et al. (1964) adjusted theoriginal parameter in order to positiontheir 01duvai habi1ines at the base of thehuman line (Wood & Collard 1999a: 199and 1999b: 66). Furthermore, endocranialcasts do not produce reliablerepresentations of the brain's linguisticareas; so Homo habilis' possession oflanguage is also debatable (Wood &Collard 1999b:66). Even the species''handyman' status is challenged by thepossibility that a contemporaneous anddexterous australopithecine may haveproduced 01dowan tools (Wood & Collard1999b:66).

In light of these classificatoryshortcomings, Wood and Collard offerednew minimum requirements foradmittance into the Homo sapiens lineage.They argued that a hominid group belongs

within the genus H0m 0 when itexhibits a human-like body size, bodyshape, brain size, ontogeny, andmasticatory apparatus (Wood &Collard 1999b:67-70). These physicalcharacteristics are relevant totaxonomy because they indicate thatthe species also shares Homo sapiens'adaptive strategies (Wood & Collard1999b:70). Conversely, a fossil taxonwill diverge from this paradigm whenit occupies a different eco-niche andbelongs to a separate evolutionary line.According to Wood and Collard's

. (1999b:67-70) study, Homo habilisfalls short of the new criteria on everycount, save brain size. They observedthat Homo habilis more closelyapproximates the australopithecinepattern and would be betteraccommodated by the genusAustralopithecus (Wood & Collard1999b:70). Though their researchmakes a good case for the re-assignment of habi1ines, primitivefossils found outside of Africa suggestthat this revision is premature.

Located at the southern borderof the Republic of Georgia, theDmanisi site has yielded hominidremains that raise questions about thevalidity of Wood and Collard'scriteria. Archaeologists excavating amedieval village uncovered ancientcraniofacial fragments within the floorof a historic building's cellar. Stonetools affiliated with the fossils moreclosely resemble simple 01dowanimplements than the complexAcheulian hand axes of African andEuropean Homo erectus sensu lato(Gabunia et al. 2001:164, Vekua et al.2002:86, Swisher et al. 1994:1121).Relative dating established a reliableage of 1.75 million years, whichsignifies that these hominids lived

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contemporaneously with several Javanesepopulations of Homo erectus (Swisher etal. 1994:1118, Gibbons 1994:1087, Vekuaet al. 2002:85). Although these twoancient populations share a continent and avery early timeframe, they differnoticeably in terms of their grossmorphology. The Dmanisi specimens havesubstantially smaller cranial capacitiesthan Asian Homo erectus and exhibit anumber of craniofacial features thatcorrespond to the Homo habilis pattern(Rightmire et al. 2006: 115, Vekua et al.2002:85 and 88). Despite these primitiveretentions, the fossils share a number ofderived traits with the Javanese groups.

Fragmentary faciocranial remainsrecovered from the Longgupo Cave incentral China also exhibit a uniquecombination of primitive and derivedcharacteristics. One hundred and sixteenassociated faunal specimens secured arelative date range of 1.96 to 1.78 millionyears. Wood and Turner (1995:240)observed that the scanty sample offractured hominid bones offers 'meagerpickings' for taxonomic assignment;however, a lower premolar (Pm4)embedded within a piece of mandible isdiagnostic. This tooth, found alongside anassemblage of Oldowan-like tools,provides additional evidence for thecolonization of Asia by a habiline. In fact,the age and location of the fossils suggestthat these hominids formed the initialmigratory wave through the Levantinecorridor approximately two million yearsago (Wanpo et al. 1995:278).

The early dates and primitivemorphology of the Dmanisi and Longgupofossils came as a major surprise to thefield of paleoanthropology. Since trans-continental travel requires a hominid topossess human-like adaptive strategies, thefirst pioneers out of Africa were assumedto have had a human-like morphology. In

fact, this presumption is the very cruxof Wood and Collard's argument forthe taxonomic revision of genusHomo. The primitive-looking bones,teeth, and tools from the two Asiansites conflict with their revisions to thecriteria for a species' admittance intothe human genus. These assemblagesprove that small-brained hominidswith underdeveloped masticatorycomplexes and simple toolkits werecapable of occupying a Homo sapiens-like eco-niche. In a similar fashion,Homo habilis' primitive retentions donot necessarily exclude it from thebehavioural ecology of archaic Homo.In addition to contradicting Wood andCollard's major conclusions, theDmanisi and Longgupo fossils alsopresent problems for Homo erectus'long-established title as the 'firstmigrant' out of Africa.

Some researchers havereconciled the existence of theseanomalous fossils by representingthem as an early grade of H 0 m 0

erectus sensu lato (Gabunia et al.2001, Lordkipanidze et al. 2006,Rightmire et al. 2006). However, suchinterpretations are often biased bypersonal beliefs about the splitting orlumping of African and Asian Homoerectus. For the sake of clarity, thispaper will employ the taxonomicsplitter's nomenclature when referringto these species; whereby, the termHomo ergaster refers to Africanspecimens and Homo erectusdesignates Asian specimens (Schwartz2004). The Dmanisi craniofacialremains share a significant number ofHomo erectus and Homo ergaster-liketraits: an elevated nasal saddle, a bar-like supraorbital torus on the parietals,a low temporal squama with a straightupper border passing downward

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toward asterion, flexion of the occiput, anda constricted foramen lacerum(Lordkipanidze et al. 2006: 1156). Gabuniaet al. (2001:165, Fig. 1) argued that therecently-discovered D2280 and D2282crania affiliate the Dmanisi populationwith East African Homo ergaster. TheD2280 and D2282 specimens possesscharacteristics that distinguish them fromAsian Homo erectus; such as their tall,thin-walled, and long cranial vaults, theirmoderate-sized supraorbital tori, andaspects of their lower dentition (Gabuniaet al. 2001:165-166). Yet, those who areeager to lump the Dmanisi assemblagewith Homo ergaster must first explain thesmall number of synapomorphies it shareswith the Javanese Homo erectus sample(Conroy 2005:417, Lordkipanidze et al.2006:1156). To add to the classificatoryconfusion, Vekua et al. (2002: 85) notedthat "the Dmanisi specimens are the mostprimitive and small-brained fossils to begrouped with [Homo ergaster]". In sum,these crania most closely resemble Homoergaster, but also incorporate severalHomo erectus specializations and someprimitive habiline retentions. The traitsthat approximate the Homo habilis patternare often represented as plesiomorphies,which situate the Dmanisi and Longgupohominids at the stem of the Homo ergasterspecies (Lordkipanidze et al. 2006: 1156,Rightmire et al. 2006: 139).

Figure 1: Lateral views of D2280 (right)and D2282 (left) crania, from DmanisiSite (Rightmire et al. 2006: 118).

Researchers who readilydismiss the Dmanisi hominids'habiline features run the risk ofmisinterpreting the significance ofthese primitive retentions. If the fossilcrania truly represent an early form ofHomo ergaster, there should be somedegree of brain size continuitybetween the two groups. However, theupper range of the Dmanisi cranialcapacities generally falls short of thelower size limit for Homo ergaster(Rightmire et al. 2006:139).Paleoanthropologists are hard-pressedto explain why the ancestral, smaller-brained hominids appear in thearchaeological record after theirlarger-brained African descendents.Moreover, Homo erectus has arelatively specialized morphology; yet,somehow this species co-existed withits unsophisticated 'ancestor'. Basedon affinities in brain size and certaincraniofacial features, Gabunia et al.(2002:88, Fig. 2) remarked that "it canbe argued that [the Dmanisi]population is closely related to Homohabilis". Hopefully, the Dmanisi sitewill produce well-preserved longbones that will clarify the phylogeneticrelationship between these hominidsand other species of Homo (Balter &Gibbons 2002).

Figure 2: Lateral views of D2700 (right)and KNM-ER 1813 (left). The Dmanisisubadult is slightly larger than theKoobi Fora [H. habilis], but the crania aresimilar in midfacial profile, supraorbitaldevelopment and rounding of the occiput.(Rightmire et al. 2006:127)

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In contrast to 'lumping' theDmanisi crania within Homo ergaster orHomo habilis hypodigms, some scientistsentertain the possibility that thisassemblage actually represents twospecies. In the year 2000, anarchaeological team unearthed mandibleD2600, which is considerably larger thanall other local specimens (Gabunia et al.2002). Those studying the fossil initiallyhypothesized that its unique dimensionscorresponded to a new species, H0 m 0

georgicus, which may have livedalongside a migrant group of H 0 m 0

ergaster (Gabunia et al. 2002:243).However, a comparison between thegracile D211 mandible and the robustD2600 specimen could not rule out thepossibility that this hominid populationexpressed extreme sexual dimorphism(Gabunia et al. 2002:244). Thus, theresearchers concluded that the Dmanisisite was occupied by only Homo georgicus(de Lumley & Lordkipanidze 2006,Gabunia et al. 2002:244-245). Thisproposition has not gained support in theliterature (Balter & Gibbons 2002). Oneoutlying individual does not substantiatethe creation of a new species, especiallysince the rest of the specimens at the siteare morphologically affiliated withAfrican members of genus Homo.

In addition to presentingpaleoanthropologists with a taxonomicconundrum, the Dmanisi and Longguporemains also defy traditional assumptionsabout early hominid migration. Clearly,Homo erectus' momentous departure fromAfrica, approximately one million yearsago, is no longer a tenable scenario. If theprimitive fossils from the two new Asiansites represent a taxon other than Homoergaster, be it Homo habilis or Homogeorgicus, then they signify that trans-continental movements were undertakenby multiple species (de Lumley &

Lordkipanidze 2006, Gabunia et al.2002, Schwartz et al. 2000).Furthermore, the habiline-like cranialcapacities of the Dmanisi specimensviolate the popular notion that earlyHomo required a relatively large brainto migrate northwards (Vekua et al.2002). Their diminutive crania alsoindicate that the Dmanisi hominidswere correspondingly small-bodied.Since the fossils retain a number ofhabiline craniofacial features, it islikely that they also possess H0 m 0

habilis-like limb proportions.Unfortunately, the Dmanisi site has notproduced enough postcranial materialto confirm this speculation. Futurelong bone discoveries will ultimatelysupport or reject the supposition thatshort legs physically prevented earlyhominids from traveling long distances(Balter & Gibbons 2002).

Material culture from theDmanisi and Longgupo site abolishesmany preconceptions about the firstpioneers into Eurasia. Acheulianimplements are no longer consideredto be the breakthrough technology thatfacilitated hominid expansion into neweco-niches. The 1.8 million-year-oldJavanese assemblages show that theearliest Homo erectus populationswere capable of migrating without thehelp of hand axes (Gibbons 1994). Infact, the lithic artifacts from theDmanisi and Longgupo sitesdemonstrate that the first trans-continental travelers were fabricatorsof simplistic Oldowan tools (Wanpo etal. 1995). Gowlett (2006:299)wondered how such small-brainedmembers of early Homo, equippedwith the most basic technology, werecapable of inhabiting temperateregions. He argued these primitivehominids were not hardy enough to

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cope with Pleistocene climatic changeswithout assistance (Gowlett 2006:299).

Thus, Gowlett (2006:304)proposed that adverse environmentalconditions created selective pressure forthe innovation of new tools, such as thecontrolled use of fire. This conjecture istoo far-reaching, since it suggests that thefirst colonizers of Eurasia possessed firetechnology long before the Africanhominids did. Indeed, the best-knownevidence of fire use prior to one millionyears ago is derived from the Swartkrans,Koobi Fora, and Chesowanja sites. Thehominid who originally wielded thistechnology has yet to be identified(Conroy 2005 :302). The Asianarchaeological record demonstrates thatHomo erectus began to employ fire as lateas the middle Pleistocene. Conversely, theDmanisi and Longgupo sites do not offer asingle trace of empirical support forGowlett's (2006) claim. He blamed thislack of evidence on the fact that thenegligible population densities of this timeequate to a very low probability thatancient campfires will ever be sampled(Gowlett 2006:305). This argument is alsoproblematic because the Dmanisi site isknown for its excellent preservation ofdelicate skeletal material. Archaeologistshave uncovered dozens of hominidspecimens and innumerable faunalremains; therefore, it seems unlikely thatburnt bone fragments and charcoaldeposits would completely escapedetection. Nevertheless, if Gowlett's(2006) speculations are correct,paleoanthropologists must revise theirassumptions about the origins of fire use.Early migrants with access to thistechnology would have adapted to theirnew territory in ways that approximate thehuman pattern, like withstanding coldertemperatures and cooking their food. Thediscovery of additional ancient sites will

hopefully clarify the ambiguous issueof Eurasian fire use.

DiscussionClearly, the fundamental tenets

of the traditional Homo erectusmigration model fail to accommodatethe Dmanisi and Longgupo specimens;thus, researchers are required toformulate alternate hypotheses. Thosewho assign these fossils to the base ofthe Homo ergaster cladesuggest that this species expanded intoEurasia shortly after its appearance inAfrica (Gabunia et al. 2001, Potts et al.2004). Homo erectus evolved from thegroups that remained in their ancestralcontinent, and formed a second waveof northward migration. Thegeographic and genetic isolation of theDmanisi pioneers explain theapomorphic features of the crania.Although these hominids are notunique enough to be split into theirown species, Homo georgicus,Rightmire et al. (2006: 140) believethat they merit the designation of anew Homo erectus (sensu lato)subspecies.

However, a major problemwith this hypothesis relates to thesynapomorphies shared between theDmanisi fossils and the derived groupsof Asian Homo erectus. Classifyingthese hominids as stem-grade Homoergaster precludes the possibility thatthis population could acquire suchspecializations. Parallel evolutioncannot explain the presence of thesederived features, since the migrantswere exploiting ecological zonesunlike those of Africa. Therefore,Dmanisi and Longgupo fossils areunlikely candidates for a taxonomicposition at the base of the H 0 m 0

ergaster clade.

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Several recently redated Javanesesites have generated a new perspective onhominid migration that drasticallydiverges from traditional models.Rightmire et al. (2006: 140) observed that,"dating does not presently rule out thepossibility that H. erectus [sensu lato]originated in Eurasia and that some groupsthen returned to Africa, where theyevolved toward H. erectus ergaster". Inother words, habiline migrants passedthough the Levantine corridor to establishIndonesian populations of Homo erectus,which later spread back into their ancestralcontinent. For this scenario to work, thederived form of Asian Homo erectus musthave evolved spontaneously from thecontemporaneous Dmanisi and Longgupogroups. Furthermore, this 'lumper'interpretation requires severalevolutionary reversals to occur, since thegeneralized African communities of Homoerectus were supposedly founded by morespecialized members of their species.

On the other hand, taxonomic'splitters' maintain the view that Eurasiawas first colonized by a stem-grade ofHomo ergaster (Wood & Turner 1995).These isolated populations later evolvedinto the separate Homo erectus clade,while their African ancestors maintainedtheir phylogenetic integrity. Once again,the contemporaneity of the Dmanisi andJava hominids presents theoreticaldifficulties. The numerous morphologicaldisparities between the crania from thetwo sites reveal the implausibility thatHomo erectus arose directly from such aprimitive-looking ancestor.

Surely, there must be a moreparsimonious alternative to suchinadequate migratory models. The smallbrain sizes and primitive craniofacialfeatures of the Dmanisi specimenspreclude an affiliation with the morehighly-evolved Homo erectus and Homoergaster species. Hence, these hominids

are derived from an earlier member ofthe genus Homo. Out of considerationfor Wood and Collard's (l999a,1999b) recent taxonomic revisions,this paper will refer to this ancestraltaxon as 'proto-habiline'. Proto-habilines were likely the earliestmembers of the human genus;however, there remains a possibilitythat this species was actually a lategrade of Australopithecus. Just beforeHomo ergaster split from the proto-habiline lineage to form its ownautonomous species, a group ofhominids left Africa to establish theDmanisi and Longgupo sites. For thisreason, the pioneers of Asia retainhabiline traits but also exhibit Homoergaster synapomorphies. The newly-established colonies becamereproductively isolated and no longercontributed to the humap line.Geographic seclusion and cranialapomorphies suggest that the Dmanisihominids are better described as asubspecies of Homo habiUs. Therefore,this paper proposes the tentative nameHomo habilis transmigro: the'migrating handyman' (Fig. 3). Moreskeletal material from the Longguposite is required in order to affiliate itsprehistoric occupants within this newcategory. Meanwhile, the populationsthat remained in Africa proceeded todiverge into two separate taxa: Homoergaster and Homo habilis. AfricanHomo ergaster formed a secondhominid pulse into Eurasia, whichultimately evolved into the erectinesthat first appeared in Java. TheIndonesian and Dmanisi taxaexperienced similar environmentalpressures, so any cranialsynapomorphies they share wouldhave been the result of parallelevolution.

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~H habihs

IllortiO habilistransn1igro

Figure 3: Adapted from Tattersall's (2003)phylogeny

ConclusionsUltimately, the Homo habilis

paradox may be resolved by majorrevisions to hominid migratory modelsand taxonomy. Contrary to Wood andCollard's (1999a, 1999b) arguments,Homo habilis is not primitive enough to beclassified as an australopithecine, asevidenced by their ability to travel acrosscontinents and colonize strange lands.These achievements approximate Homosapiens' adaptive strategies and firmlyestablish this taxon at the base of thehuman line. The Dmanisi and Longgupofossils present paleoanthropologists with aclassificatory nightmare. However, theirunique combination of primitive andderived traits is best accommodated by anew subspecies category: Homo habilistransmigro. The recognition of Asianhabilines challenges many traditionalpresumptions about the first hominidmigration out of Africa. Additional longbone and cranial material from theDmanisi and Longgupo sites will confirmor deny the existence of multiple,

contemporaneous taxa co-existingthroughout ancient Eurasia.

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