Homo erectusin East Asia: Human Ancestor or Evolutionary ......In order to better understand the...

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evolution, part of an intractable debate that continues to swirl over the origins ofanatomically modern people.

Many readers are assuredly familiar with the contours of this debate,between those who advocate a recent, uniregional origin for modern Homo sapi-ens in Africa (henceforth termed the “recent out of Africa,” or ROA, hypothe-sis) versus those who support a more deeply rooted, multiregional origin for ourspecies (termed MRE for “multiregional evolution”). The implications of thesetwo differing interpretations of modern human origins for understanding thecourse of human evolution and the status of H. erectusduring the last 1.8 millionyears of earth history are manifold.

One major premise of the uniregional ROA hypothesis is that modernhumans originated in sub-Saharan Africa between 100-200,000 years ago byevolving a set of unique physical features and cultural innovations that allowedthem to disperse throughout the world, replacing older, more archaic human lin-eages that had established themselves in various regional settings. Most advo-cates of this hypothesis view the genus Homo as subject to the same evolution-ary processes of divergence and differentiation as any other biological entity.ROA supporters are thus prone to use cladistic analysis, which attempts to parti-tion fossil specimens into distinct evolutionary lineages by documenting unique-ly derived features (autapomorphies) that separate one from another (Harrison1993; see box 1).

In order to better understand the nature of Homo erectus, the sig-nificance of its evolutionary history in East Asia, and the role it playedin human evolution, it is first necessary to come to grips with what ismeant by the term itself, as H. erectus has come to mean differentthings to different people. For some, H. erectusrepresents the first trulypandemic human species and the direct progenitor of archaic andmodern H. sapiens. For others, H. erectus is an interesting footnote tothe saga of human evolution, a distinct species that emerged in theEast, only to go extinct without issue when modern humans expand-ed their range out of Africa to encompass the far reaches of the OldWorld. From this latter perspective H. erectus serves as the easternanalog of what is thought, by some, to be another failed experimentin humanity - the Neanderthals. The very manner in which H. erec-tusis conceived is thus held captive to two competing views of human

Homo erectus in East Asia:Human Ancestor or Evolutionary Dead-End?

Dennis A. EtlerDepartment of Anthropology

Cabrillo College, Aptos, California

Box 1: Pleistocene hominid phylogenies: multilineal and recent out of Africa theoriesThe phylogeny in figure 1 is consistent with Tattersall’s (1999) mul-

tilineal paradigm that envisions multiple species of Homo throughout thePleistocene. In this rendition Asian descendants of H. erectus are viewedas possessing derived features that distinguish them from their commonancestor and each other. Post-erectine Chinese hominids (H. daliensis) arederived in having larger cranial capacities in association with changes inthe structure of the braincase and cranial base. Australasian descendants ofH. erectus (H. soloensis) likewise show significant increases in cranialcapacity, but retain ancestral erectine-like features of the cranium (Swisheret al. 1997). The establishment of separate species to accommodate thisdiversity is in keeping with the multilineal interpretation of hominid phy-logeny.

Fig.2:Summation of Pleistocene hominid phylogeny as conceptualized by ROA. The authorpresents this as a logical interpretation of ROA, but disagrees with its basic premises.

Figure 2 attempts to summarize current views of hominid phylogeny held by themajority of ROA advocates. In this scheme H. ergaster serves as the Last CommonAncestor (LCA) of all later Pleistocene hominids. It is now thought to have dispersedout of Africa to western Eurasia (Dmanisi, Georgia) as early as 1.8 mya (Gabunia et al.2000, 2001). It is hypothesized that this eastern dispersal led to a speciation event withthe establishment of H. erectusin East Asia (China) (Huang et al. 1995) and Australasia(Java) (Swisher et al. 1994). Further west H. ergaster is thought to have evolved into H.heidelbergensis in Europe and Africa, the European branch leading to the latePleistocene Neanderthals (H. neanderthalensis) and the African branch to H. sapiens.Many proponents of ROA interpret post-erectine, pre-modern hominids of China aspart of an eastern range expansion of H. heidelbergensis that led to cohabitation with,or the replacement of, H. erectus. A fossil gap between pre-modern and modernhumans in East Asia is thought by some advocates of ROA to represent a true lacunaattributable to an extinction event that led to a depopulation of East Asia prior to thearrival of modern H. sapiens (Jin and Su 2000).H. erectus is thought to have survivedin Australasia as a distinct taxon well into the late Pleistocene (Swisher et al. 1997).

Fig.1: Proposed full extension of multilineal hominid phylogeny during thePleistocene. The author presents this phylogeny as a logical extension of the mul-tilineal paradigm, but does not subscribe to it.

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played in the origins of modern people. FromBräuer’s standpoint, proponents of ROA can besorted into two camps, those who support “hard”or “soft” replacement. The hard ROA positionemphasizes the near total replacement ofEurasian and Australasian archaics by dispersingAfrican moderns (Stringer 1984, 1988), whilesoft ROA (the Afro-European model of Bräuer,1984, 1992, 2001) allows for varying degrees ofgene flow and hybridization between variousregional groups and advancing moderns, as wellas the persistence of some regional traits acrossthe archaic/modern boundary. This “hybridiza-tion and replacement” model, while more mal-leable than the “total replacement” model,nonetheless still emphasizes replacement versusin situ evolution, and unidirectional extra-African versus multidirectional pandemic dis-persals, as the predominant mode by which themodern human phenotype spread throughoutthe world.

Although proponents of MRE continue todocument regional continuity in Europe (Frayer1993), Western Asia (Kramer et al. 2001),Eastern Asia (Wolpoff et al. 1984; Wolpoff1985; Etler and Li 1994; Etler 1996), andAustralasia (Hawks et al. 2000), its basic, under-lying premise is that modern humans do notshare an exclusively African origin. Instead,MRE contends that all human populations of thepast made differential contributions to all popu-lations of the present. In this context there is noneed to assert that Pleistocene humans from dif-ferent regions of the world were necessarily thelineal antecedents of their present day inhabi-tants. MRE does not deny the possibility that var-ious local evolutionary histories, some entailingthe replacement of one population by another,may have unfolded in different regions of theworld at different times in the past. In this senseMRE is not necessarily contingent on regionalcontinuity, i.e. the direct linkage of predecessorwith successor populations, in any given regionof the world, although this may well be the case.While the focus of MRE has generally been onregional continuity, a greater emphasis on thepandemic nature of the human species through-out the Pleistocene and the trans-regional timingof major transitions in the fossil record may bet-ter help elucidate factors that led to the emer-gence of modern humans.

Debating the evidence:Advocates of ROAand MRE have been very aggressive in chal-lenging each other on a number of fronts, includ-ing genetics, phylogenetics, archaeology, and the

Notwithstanding the pitfalls of identifyingtrue autapomorphies (Nowaczewska 2000), thecladistic mindset has led some anthropologists tosuggest that there was a proliferation of humanspecies throughout the Pleistocene (see Tattersall1999). In this cladogenetic, multilineal schemethere are ancestral species that occupy nodes ofdivergence, terminal species that represent theend products of lineages diverging from oneanother, and intermediate or transitional speciesthat serve as links between the nodes and termi-nations. If the multilineal paradigm is strictlyadhered to an elaborate geneology of Pleistocenehominids can be created (fig.1). Most multineal-ists, however, do not subscribe to as species richa phylogeny as so depicted. A favored interpre-tation of Pleistocene hominid phylogeny formany multilinealists is presented in fig.2. Nomatter how many Pleistocene species of Homoare recognized, the crux of human evolutionarystudies for multilinealists is to try to understandtheir defining features and in particular what fac-tors led to the emergence of modern humans, suigeneris, from their not quite fully formed fore-bears (see Foley and Lahr 1997).

As one may suspect, the opponents of theuniregional multilinealists are the multiregionalunilinealists. Rather than seeing modern humans

as the result of a relatively recent, unique specia-tion event, the unilinealists see our species as theend product of a continuously evolving lineagethat has been arbitrarily divided into time succes-sive chronospecies, generally speaking H. erec-tusand H. sapiens. These species are seen as partof an evolutionary continuum so, as logic dic-tates, many proponents of MRE recognize butone species of human during the Pleistocene, i.e.the ever-evolving H. sapiens, of which we arethe latest incarnation (Wolpoff et al. 1994a) (seebox 2). Human evolution during the last 1.8 mil-lion years has hence been both anagenetic andincremental. For proponents of MRE, the task ofhuman paleontology is to better understand thetransitions that have catapulted early H. sapiens(i.e. H. erectus) from a marginal player on theworld’s stage to our present role as producer anddirector of not only our own future but that of theworld-at-large (Hawks and Wolpoff 2001a,2001b).

Refinements of ROA and MRE: Althoughmany ROA supporters reject the idea that therewere meaningful interactions between archaicand modern humans subsequent to the latter’sextra-African dispersal, some of its advocates aremore amenable to compromise regarding therole that regionally distributed archaic humans

Homo erectus in East Asia

Box 2:Pleistocene hominid evolution according to MREThe diagram in figure 3, adapted from Templeton (2002), reflects the underlying premise of multi-

regional evolution (MRE), that human evolution during the Pleistocene is characterized by discrete dis-persal events intercalatedwith recurrent gene flowwith isolation by distance.The two latest “out ofAfrica” dispersals may beassociated with majortransitions in the fossilrecord, from erectines toarchaics and archaics tomodern humans. Thegenetic signals of otherdispersals may have beenlost through selectivesweeps associated withmajor transitions in thehuman fossil record. Inthis scheme human evolu-tion (i.e. the evolution ofthe genus Homo) from thebeginning of thePleistocene on is charac-terized by one evolvinglineage with no speciation.

Fig.3: Reticular interpretation of hominid evolution during the Pleistocenebased on recent genetic research.


fossil record. The genetic evidence in favor ofROA rests primarily on studies of mitochondrialDNA (mtDNA) and the Y-chromosome.Advocates of ROA claim that these studiesprove a late Middle Pleistocene to early LatePleistocene exclusive African origin for all livingpeople (Cann et al. 1987; Hammer 1995; Jin etal. 2000; Brookfield 2001; Harpending 2001;Jorde et al. 1998; 2000; Ke et al. 2001; Vigilantet al. 1991). Proponents of MRE, however, havecountered with studies of autosomal genes of thenuclear genome, and the X-chromosome, thatshow evidence of deep population structurewithin our species that greatly predates theappearance of modern H. sapiens (Harding et al.1997; Harris and Hey 1999).

Phylogenetic studies have been used byadvocates of ROA to negate the reality of region-al traits while attempting to validate the existenceof derived, species defining features (autapomor-phies) of Asian H. erectus (Andrews 1984;Bilsborough 2000). On the other hand advocatesof MRE use similar studies to demonstrateregional continuity in Europe, Asia, andAustralasia (Wolpoff et al. 1994b, 2001).

Certain aspects of the archaeological recordhave been cited by supporters of ROA to showa clear divide between the cognitive powers ofarchaics and moderns (Foley and Lahr 1997),while supporters of MRE have used otheraspects of the archaeological record to demon-strate continuity between archaic and moderntechnology (Kramer et al. 2001).

Finally, the fossil evidence of human evolu-tion during the Pleistocene has been used by par-tisans of ROA to show that the earliest record ofmodern humans is in Africa and the Levant tothe exclusion of other regions (Stringer andAndrews 1988), while supporters of MRE usethe human fossil record to demonstrate the tran-sitional nature of pre-modern forms of H. sapi-ens in various regions of the Old World. Belowan attempt is made to review some of theseissues.

Genetic evidence: Over the last twodecades the sequencing of various segments ofthe human genome has resulted in the accumu-lation of increasingly refined datasets relevant todeciphering the relationships of extant humanpopulations to one another and to extrapolatinghuman population histories (Harpending 2001;Harpending and Rogers 2000; Harpending andEller in press; Relethford 1998; Relethford andHarding 2001). The analysis of these datasets hasgenerally supported ROA’s contention that the

modern human genome emerged as the result ofa population bottleneck in sub-Saharan Africaduring the last 100,000 years. The primarydatasets that have been employed are themtDNA genome, non-coding and/or non-recombining segments of the Y-chromosomeand various genes or non-coding segments of thenuclear DNA genome.

These datasets, however, do not necessarilyreflect the same population histories, as there isno intrinsic relationship between the lineagesthey represent. Matrilineal mtDNA lineages andpatrilineal Y-chromosome lineages, for instance,track different demographic components of ourshared genetic heritage, while the nucleargenome, which is subject to recombination, fol-lows a different set of rules entirely. It is not sur-prising, therefore, that the nuclear genomereflects deep population structure within modernhuman populations that predate the emergenceof anatomically modern humans (Ayala 1996;Harding et al. 1997; Harris and Hey 1999;Hawks et al. 2000; Relethford and Harding2001). What therefore seem to be contradictoryindicators of coalescence in our genetic past mayin reality be different facets of our shared heritagereflected through the prism of modern genomics(see Hawks et al. 2000b; Hawks and Wolpoff2001b).

Another concern that needs to be addressedis the underlying premise employed in the analy-ses of various genomic datasets. Since the 1970sthe neutral hypothesis, which posits that much ofthe human genome is selectively neutral andhence evolves at a statistically calculable rate (i.e.the molecular clock), has been the prevailing par-adigm used to interpret haplotype trees demon-strating branching orders in the sequestration ofextant populations. The rooting of thesegenealogical trees in Africa has been used as evi-dence in support of ROA, while the neutralhypothesis supports the idea that a populationbottleneck was the only effective means bywhich genetic variation in modern human popu-lations has been constricted.

Recent studies, however, are showing thatvarious non-coding segments of the genomeplay a significant role in gene regulation (Chu1998; Dermitzakis et al. 2002). In addition,mtDNA has been implicated in various meta-bolic diseases of the nervous system, such asParkinson’s syndrome, etc. (Wallace 1999), andin apoptosis (programmed cell death) associatedwith the aging process (Green and Reed 1998).These studies highlight a new appreciation of the

role played by the mitochondrial genome in theregulation of neurological and other basic lifeprocesses that have certainly been altered in theevolution of modern humans. Changes in thefunction and expression of mitochondrial genesmay therefore have been of decisive importancein the evolution of modern neurochemistry and,concomitantly, various parameters of modernhuman physiology and behavior (Ruiz-Pesini etal, 2004) . If this is the case, the entire mitochon-drial genome may be subject to positive direc-tional selection, in effect neutralizing the neutralhypothesis (Wise et al. 1998).

The remaking of the human genome in thetransition from pre-modern to modern H. sapi-ens may thus have been the result of multipleselective sweeps rather than a single populationbottleneck. Reinterpreting the genetic data basedon a different set of premises than those held byadvocates of ROA results in an entirely differentset of conclusions that favor a more multifacetedinterpretation of recent human evolution thatemphasizes multiple dispersals and recurrentgene flow with isolation, all of which redound infavor of MRE (Templeton 2002; Hawks et al.2001b). Other factors such as “cultural hitchhik-ing” in which cultural evolution is hypothesizedto have “reduced the diversity of genes whichhave similar transmission characteristics to selec-tive cultural traits” (Whitehead et al. in press)may also help to explain the process by whichgenetic homogenization of modern human pop-ulations has taken place.

Regional traits vs. species defining autapo-morphies: It is interesting to note the similaritiesin how another dataset relating to MRE andROA has been evaluated and disposed of. Thisdataset consists of morphological features, men-tioned above, that on the one hand are used tosupport the existence of Asian regional traitschampioned by supporters of MRE, while on theother hand are used to support the existence ofuniquely derived, species defining features(autapomorphies) of H. erectus, championed bysupporters of ROA.

Regional traits have been used, ever sinceWeidenreich’s classic studies of “Peking Man”(Weidenreich 1936, 1937, 1943), as a lodestonepointing towards continuity in the evolution ofregional populations in Europe, East Asia, andAustralasia. Supposed autapomorphies of H.erectus in Asia, have been used by supporters ofROA to buttress their position that archaic popu-lations throughout Eurasia were distinct at thespecies level and uninvolved in the origins of



if it is shown that a particular regional trait occursin other regions of the world or that the trait is ple-siomorphic (i.e. an ancestral retention) for thetaxon under consideration, its value as a regionalmarker is not necessarily diminished if it can beshown to occur in greater frequency and for agreater length of time in the region under study.Species defining autapomorphies are, on theother hand, by definition both unique andderived. If it can be shown that the trait is neither,its value as a species defining character is totallycompromised.

In his monograph, A Theory of Human andPrimate Evolution, Groves’ (1989) critique ofregional traits falls victim to the first error. He dis-misses most of them by demonstrating that theyare not unique to the region they are supposed tocharacterize or that they are plesiomorphic forhumans as a whole. This sort of analysis would

obviate the usefulness of a character said to beautapomorphic, but does little in the way ofnegating a character said to be regional. Lahr’s(1994) critique, published in the Journal ofHuman Evolution, is more substantive as shedemonstrates that a number of supposedly EastAsian regional traits are redundant, not geneti-cally controlled or do not follow the expectedregional frequency pattern. On the other hand,her study re-substantiates the well-known,empirically obvious fact that the East Asian faceis characterized by transversal flatness and high,laterally flaring cheek bones, a suite of featuresseen in nearly all East Asian hominids, both liv-ing and fossil (Wang and Tobias 2000, 2001).Unfortunately, Lahr’s analysis includes only onedental feature, third molar agenesis. By notincluding upper incisor shoveling the impact ofher study is diminished. Her research does, how-


modern humans. What is interesting to note isthat many of these traits are virtually identical.Moreover, both interpretations have been chal-lenged by a variety of studies and shown to bewanting to varying degrees.

The basic problem with utilizing morpho-logical features as determinants of evolutionaryrelationships is the polytypic nature of the humanspecies, the polymorphic character of the traitsunder consideration, and the peripatetic amblingsof human beings during the Pleistocene. Thereis, moreover, an epistemological differencebetween the two approaches. Regional traits, nomatter how misconstrued by their critics, neednot be unique to any given region, nor must theynecessarily be derived relative to the ancestralcondition; i.e. a regional trait can be a primitiveretention that is maintained at a high frequency inone part of the world while lost in another. Thus,

Box 3: The earliest Chinese hominid: truth or consequences?Since the “recent out of Africa” (ROA)

hypothesis of modern human origins gained pop-ularity in the 1990s, vigorous attempts have beenmade to isolate Asian H. erectus from the main-stream of human evolution. One such attempt hasbeen the recognition of an early species of Homoat the Longgupo cave site in Wushan, Sichuanprovince, China. In 1995, fragmentary fossilremains from Longgupo Cave, consisting of amandibular fragment containing a well worn P4and M1 (fig.4) and an unassociated, isolated upperincisor, all dated to approximately 1.9 mya, wereattributed to an early form of the genus Homo, saidto be ancestral to an exclusively Asian H. erectusthat eventually went extinct without issue (Huanget al. 1995; Ciochon 1995). This identification ofthe Longgupo remains, however, as belonging toa possible ancestor of an Asian delimited H. erec-tus, is controversial. In point of fact, the Longgupomandibular specimen has been shown to favor-ably compare with Asian apes from late Miocenethrough early Pliocene sites in Yunnan, while lack-ing features commonly seen in early specimens of

Homo (Wu 2000; Etler et al. 2001). The Wushanmandibular P4 (fig.4) is characterized by a slit-likeanterior fovea; anteriorally placed, twinned proto-conid and metaconid; and a broad, recessed talonidbasin. The Wushan mandibular M1 is characterizedby 5 cusps arrayed around the periphery of a broad,deep talonid basin, a buccally placed hypoconulidand differential wear on the buccal (cheek-side)cusps. Comparing the Wushan mandible to the fos-sil hominoid specimen from Yuanmou, Yunnan,dated to 6-5 mya (fig.5), note the overall similarityof lower P4 morphology, in particular the constrict-ed anterior fovea; the anteriorally positionedtwinned protoconid and metaconid; the broad, deeptalonid basin; and the sub-oval shape of the crown.M1 morphology does not differ in any discernableway from that seen in the Wushan specimen.Compared to the Wushan specimen, the P4 of theDmanisi jaw (fig.6), dated to 1.8 mya, has centrallyrather than anteriorally placed bulbous protoconidand metaconid, and a delimited talonid basin con-stricted into a distally placed posterior fovea, analo-gous to the mesially positioned anterior fovea. The

Dmanisi M1 is distinguished by bulbous cuspsthat fill in the talonid basin, and has a distally ratherthan buccally placed hypoconulid. The DmanisiM1 occlusal wear is evenly distributed over thetooth crown.

In addition, the isolated upper incisor found atLonggupo is indistinguishable from that of a mod-ern human (Wang 1996; Wu 2000; Etler et al.2001) and was most probably “brought in byflowing water or other forces into the fissure of thecomparatively old Longgupo Cave deposits . . .and mixed with the Longgupo fauna” (Wang1996; Etler et al. 2001). Hence, there is a strongpossibility that the Longgupo specimens are apalimpsest and do not represent a valid hominidtaxon (see also Schwartz and Tattersall 1996). It is,therefore, extremely unlikely that the Longguporemains are an early hominid ancestor of an exclu-sively Asian delimited H. erectus.

Fig.4: Two views of the Wushan (Longgupo) denti-tion, with mandible and associated fourth premolar(P4) and first molar (M1)( photo: H Lanpo).

Fig.5: Mandibular dentition, with fourth premolar (P4)and three molars, of hominoid fossil from Yuanmou,Yunnan, dated 6-5 mya (photo: Li Kunsheng).

Fig.6: Mandibular dentition of the Dmanisihominid, dated to 1.8 mya (photo: L. Gabunia).

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supporters of ROA has not, however, left themnonplused in their advocacy of ROA. In likemanner, the abandonment of many traits linkedto regional continuity should not be miscon-strued as a vindication of ROA. If, for the sake ofargument, the Neanderthal and Ngandong (i.e.late Javan occurrence of “H. erectus”) questionsare set aside, there remains a swath of archaichumans, distributed throughout much of the OldWorld, from Africa through the Middle East toSouth and East Asia, that could have graduallyevolved into modern H. sapiens without anymajor replacement event. This position does notdeny that some important local replacements, insuch peripheral areas as Europe or insularSoutheast Asia, may have taken place. In thisregard it would be foolish to suggest that popula-tions have not frequently been supplantedthroughout recorded history. The history of thelast few millennia, or the past few centuries forthat matter, gives stark testimony to this ratherself-evident fact. Is it so surprising then that cer-tain marginal populations in western Europe (e.g.Neanderthals) or Java (e.g. the Solo/Ngandongpopulation) may have met a similar fate?

The intermittent movement of both genesand people, due to changing paleoclimatic orother paleogeographical and paleoecologicalfactors, throughout the central “zone of sapi-enization” identified above, could account for

apparent disparities in both the tempo and modeof human evolution in various regions of the OldWorld. It is unlikely that this process took placein the same, exact manner or at the same, exacttime in, for instance, Africa and East Asia. Therecould very well have been staggered transitionstowards a more modern morphology, variousregions at times leading the way and at othertimes lagging behind. This position is consistentwith Bräuer’s soft replacement model, that sug-gests the emergence of modern humans was“multi-causal, different in various regions andhardly rapid . . .” or for that matter uniform. Atthis point, however, MRE and “soft” ROA partways. “Soft” ROA still emphasizes extra-African dispersal and replacement, whileaccommodating limited amounts of hybridiza-tion and continuity. MRE, on the other hand,hypothesizes that there was a continuous humanpresence in central regions of the Old Worlduninterrupted by any major replacement event.To the contrary, the introduction of exogenousgenes into local gene pools may have played asignificant role in initiating major transitions inthe fossil record rather than near or total replace-ment of one population by another.

Fossil evidence: If the Longgupo remainsare discounted (see box 3), the earliest humanremains in East Asia belong to the partial crani-um from Gongwangling in Lantian county,

ever, confirm the expected regional patterning ofM3 agenesis (see also Liu and Zeng 1996), andit can be assumed it would have likewise con-firmed the prevalence of upper incisor shovelingin East Asia as well. Others have done significantresearch in respect to upper incisor morphologyto document the distinctive pattern of East Asianshoveling and its higher incidence in East Asiathan anywhere else (Crummett 1994; Liu 1995,1999a, 1999b). Nevertheless, Lahr’s study actu-ally lends support to the recognition of two majorphenetic complexes that characterize East Asianpeople both now and in the past, one facial andone dental. It suffices at present to say thatattempts to negate the reality of regional traits inEast Asia are less than devastating and, in fact,more often than not can be interpreted to supportthe very thing they are intended to criticize.

Although many East Asian traits once con-sidered to provide evidence for continuity havebeen challenged, the reality of autapomorphiesthat define an exclusively Asian H. erectus hasbeen effectively shown to be entirely wanting(Bräuer and Mbua 1992; Asfaw et al. 2002). Inpoint of fact, multilinealists have been hardpressed to present a case for species definingautapomorphies for any Middle through LatePleistocene hominids other than theNeanderthals. The abandonment of autapomor-phies of an Asian delimited H. erectus by many


Fig.7: Map of China showing homind sites discussed in the text. Yuanmou, Yunnan hasa diverse late Miocene/early Pliocene mammalian fauna that includes a large number ofhominoid fossils akin to late Miocene Lufengpithecusfrom the nearby site of Lufeng. Thelate Pliocene Wushan mandibular fragment, thought by some to have hominid affinities(Huang et al. 1995), actually compares favorably to fossil ape specimens from Yuanmouand Lufeng. Relatively complete material of H. erectus in China is known fromZhoukoudian, Lantian, Yunxian, Hexian, and Nanjing. Relatively complete material of apre-modern form of H. sapiens is known Dali and Jinniushan. Recent re-dating ofChinese hominid sites raises the distinct possibility that there is no overlap between H.erectus and pre-modern H. sapiens in China (contra Chen and Zhang 1991; Chen et al.1994), with all H. erectus specimens being older than 400,000 years and all pre-modernspecimens being younger than 250,000 years (Shen 2001).

Fig.8: Distribution of erectine and archaic Chinese fossil hominids during thePleistocene. Recent reinterpretations of the dating of East Asian hominid fossilsstrongly suggests that all specimens of H. erectus in China are older than 400,000years, while pre-modern archaics are younger than 250,000 years. Hence there wasno overlap between the two groups as once thought.


Saanxi province, China dated in excess of 1 mya(Etler 1996) (fig.7). Recent re-dating of otherhominid fossils in China has helped to clarifytheir temporal sequence (fig.8; see box 4). As inAfrica, there is clear evidence of transitionalhuman fossils in East Asia. Advocates of ROAhave a difficult time evaluating this material. Ifthese specimens are seen as descendants of anAsian delimited H. erectus, they have evolved inparallel to western archaics. If they are placed inH. heidelbergensis, there must have been areplacement event in East Asia preceding theadvent of modern H. sapiens. This would thenentail two major replacement events havingtaken place in this part of the world. MRE sug-gests that there were transitional forms betweenH. erectusand modern H. sapiens in Asia just asthere were in Africa. The new dates show that H.erectus in China is older than previously thoughtand does not overlap with more advanced archa-ic Chinese.

The major issue in Chinese paleoanthropol-ogy thus becomes a question of the relationshipof time-successive hominids to one another? Arethe archaic Chinese fossils transitional betweenearlier erectines and later moderns? Do theChinese archaics represent a continuation of theerectine lineage in East Asia that eventually goesextinct? Or, are the archaics representative of anincursion from the West, distinct from their erec-tine predecessors? To address these issues a nullhypothesis, that Chinese archaics dating fromapproximately 250,000 - 100,000 ya, are transi-tional forms between Chinese erectines and earlymoderns, can be proposed. In order to demon-strate replacement as the mode by which mod-ern humans became established in East Asia theabove stated null hypothesis must first be falsi-fied.

As mentioned previously, one means usedto discredit the transitional nature of Chinesearchaics has been to deny the existence of region-al characters in East Asia. But is it necessary todocument regional characters in order to demon-strate continuity? It is interesting to note that sup-porters of ROA have never been very much con-cerned with the presence of regional traits in theone region of the world they are most interestedin, i.e. Africa. They, nevertheless, believe thatcontinuity in human evolution occurred there, ifnowhere else. In point of fact the underpinning ofROA is a soft interpretation of continuity theoryin Africa.

As summarized by Lahr (1994), citingStringer and Andrews (1988), the ROA model is


Box 4: The Nanjing fossils and dating of Chinese hominids

New dates obtained for fossil hominids from Nanjing, Hexian, Zhoukoudian, and Yunxianindicate that all known specimens of H. erectus in China are older than 400,000 years. These datesmitigate against the likelihood that H. erectus and pre-modern forms of H. sapiens coexisted inChina as had been previously suggested.

The Nanjing fossil hominids (fig.9) were discovered in 1993-1994 at Hulu Cave, TangshanHill (N32°, E119°), on the outskirts of Nanjing (old Nanking). Based on the detailed descriptionand metric data supplied in the monographic treatment of the hominid remains by the TangshanArchaeological Team (1997) there can be no doubt regarding their great similarity to Homo erec-tus as known from the “Peking Man” site at Zhoukoudian (see also Wang and Tobias 2000).Interestingly, the associated fauna at Hulu Cave is so similar to that found at Zhoukoudian thatdirect correspondences can be made to layers 6-7 of Zhoukoudian Locality 1. These layers haveproduced only a scanty amount of hominid material and have been previously dated between400-350,000 ya. Mass spectrometric U-series dating of speleothems from Tangshan Cave, com-bined with ecological and paleoclimatic evidence, however, indicates that the Nanjing hominidsshould be dated to at least 580,000 ya, or more likely 620,000 ya (Zhao et al. 2001). This age is270,000 years older than previous estimates. Other Chinese sites yielding H. erectus, such as theupper layers at Locality 1 at Zhoukoudian and the Hexian site in Anhui, have recently been redat-ed in excess of 400,000 ya (Shen 2001; Shen et al.1996; Grun et al. 1998), while the Yunxianhominids from Quyuanhekou in Hubei have recently been dated in excess of 580,000 ya (Yan1993; Chen et al. 1996). The main sequence at Zhoukoudian that has yielded the majority of H.erectus specimens is now thought by some to be approximately 800,000 years old or older (Shen2001). These dates indicate that all Chinese H. erectus specimens are older than previouslythought and do not overlap with later more advanced archaic specimens known from Dali, Maba,and Jinniushan dated to approximately 250,000 - 120,000 ya. (Shen 2001). The conjectured con-temporaneity of H. erectus and H. sapiens in China (Chen et al. 1991, 1994) is thus most likely achimera.

The resolution of dating incongruities such as those discussed above goes a long way toresolving various conundrums associated with the Chinese human fossil record. If the above tem-poral ordering is accepted, the trajectory of human evolution in China becomes more lineal andstraightforward. What becomes most relevant are 1) the relative heterogeneity of H. erectus inChina at the demic level and 2) the transformation of human populations in China from the erec-tine to the archaic bauplan between 400,000 and 250,000 ya. Given the worldwide dearth ofhuman fossils remains between 750,000 and 400,000 ya and the margin of error associated withemployed dating techniques, it can be argued that there is no discrepancy between the course ofhuman evolution in the East and West during the Middle Pleistocene. In its broad outlines thetransformation of the human lineage from H. erectus to archaic and later pre-modern forms of H.sapiens should be viewed as simultaneous throughout the then inhabited world.

Fig.9: This partial cranium of H. erectus from Nanjing (shown at left in profile, and at right in frontal view) is in alldetails nearly identical to the reconstruction of “Peking Woman” affected by Weidenreich nearly six decades ago. TheNanjing H. erectus site contains a fauna that correlates with layers 6-7 at Zhoukoudian and documents a southerndeployment of the Zhoukoudian deme half a million or more years ago (photo:Lu Zune).


predicated on the following three assumptions:“first, the presence of archaic-modern transition-al fossils in the African late Middle Pleistocene,represented by the group of Ngaloba, Irhoud,Omo 2, Florisbad, Eliye Springs; second, theappearance of early upper Pleistocene fossils inAfrica and the Middle East, like Klasies RiverMouth, Border Cave, Singa, Mumba, Qafzehand Skhul; and third, the morphological discon-tinuity between Neanderthals and modernsobserved in Europe.” Nowhere to be found isany mention of “regional traits” that characterizeboth fossils and modern Africans, reflecting anancestor-descendant relationship. That is to say,supporters of ROA do not bother trying todemonstrate “regional continuity” in Africa. It isleft to multiregionalists to try to demonstrate sucha relationship in other regions of the world, andif they encounter difficulties in doing so that issupposed to redound in favor of the ROA posi-tion.

MRE, however, does not necessarily rely ondocumenting regional traits to demonstrate con-tinuity in human evolution in particular regionsof the world, nor does MRE dismiss them.Rather, advocates of MRE can apply the samecriteria that supporters of ROA apply to theAfrican fossil record; i.e. first, there should befossils of a transitional nature in East Asia duringthe late Middle Pleistocene; second, the appear-ance of modern-looking fossils should occurduring the early Upper Pleistocene; and third,there should be the lack of any major disconti-nuity in the fossil record of the region understudy. If the above conditions are demonstratedin East Asia or any other region of the world out-side of Africa, then the null hypothesis of conti-nuity in the region under study will remain unfal-sified, and the whole foundation of ROA itselfbegins to founder. An analysis of these condi-tions in East Asia follows.

Condition 1: There should be archaic-to-modern transitional fossils during the late MiddlePleistocene of East Asia. This is readily demon-strated in China where a group of transitional fos-sils from the same broad time range as that inAfrica is well documented (fig.10). Theseinclude partial or complete crania from Maba,Dali, and Jinniushan, more fragmentary cranialremains from Xujiayao and Chaoxian, and a sig-nificant dental sample from Tongzi (Wu andPoirier 1995; Etler 1996) (figs.7,8). Whateverdoubts one may have about the exact ages ofthese specimens, there is no question that theycan all be dated to the late Middle Pleistocene or

very early Upper Pleistocene, i.e. between 250and 100 kya, broadly contemporaneous withsimilar fossils from Africa. Some ardent sup-porters of “hard” ROA have expressed doubtsabout the relationship between these archaicChinese fossils and earlier local H. erectus popu-lations and suggest that post-erectus/pre-modernhumans in China show closer affinities toEuropean archaics (i.e. H. heidelbergensis)(Stringer 1988; Rightmere 1998) than they do topredecessor populations of H. erectus. Very few,if any, researchers who have worked with theChinese material have, however, made similarassessments. In addition, the same could be saidabout specimens considered precursorial tomodern humans in Africa, i.e. they are more sim-ilar to archaics from other regions than they areto the populations that preceded them in Africa.Verdict: An unbiased, objective evaluation of thedata from China confirms Condition 1, that thereare late Middle Pleistocene transitional fossils inChina.

Condition 2:There should be modern look-ing fossils during the early Late Pleistocene inEast Asia.. Until recently, this was somewhatproblematic and it is still controversial. There isadmittedly a significant gap in the human fossilrecord in China from approximately 30-100,000years ago (Etler 1996). This is when the

European Neanderthals flourished, so it is puz-zling as to why there is such a dearth of fossilhuman remains in China from this period oftime. Nevertheless, recent redating of the fullymodern Liujiang material from the southernChinese region of Guangxi, consisting of a well-preserved cranium and partial skeleton discov-ered in the 1950s, indicates an antiquity in excessof 100,000 years and perhaps as old as 150,000years before present (Shen et al. 2002). Althoughthe original provenience of the Liujiang remainswithin the cave deposits from whence they wereretrieved is still debated, corroborating evidencefrom the immediate vicinity is supportive of theearly age assignment. This would place fullymodern Homo sapiens in southern China in atime frame identical to that of the earliest repre-sentatives of modern humans known from any-where in Africa. There are, moreover, otherremains that bear on this issue. The Xujiayaomaterial, dated to approximately 100 kya, is acase in point. Bräuer (1992) states that “theremains from Xujiayao . . . are too fragmentaryand morphologically quite heterogeneous to beof much help in clarifying what happened (inChina)” during the transition to modern humans.The Xujiayao remains, however, consist of twofairly complete parietals, a number of parietalfragments, a nearly complete temporal, two rela-


Fig.10: As in Africa, there is clear evidence of transitional human fossils in East Asia. Advocates of ROA havea difficult time evaluating this material. If these specimens are seen as descendants of an Asian delimited H. erec-tus, they have evolved in parallel to western archaics. If they are placed in H. heidelbergensis, then there musthave been a replacement event in East Asia preceding the advent of modern H. sapiens in East Asia. This wouldthen entail two major replacement events having taken place in this part of the world. MRE suggests that therewere transitional forms between H. erectus and modern H. sapiens in Asia just as there were in Africa.



Box 5: Chinese fossils throw light on phylogenetic affinities of Atapuerca-Gran Dolina hominidsThe hominid species, Homo antecessor, founded on a number of cranio-

facial specimens from Lower Pleistocene deposits at Atapuerca, Spain, wasdescribed as the common ancestor of Neanderthals (“H. neanderthalensis”)and modern humans (H. sapiens) (Bermúdez de Castro et al. 1997). In thiscontext, fossil hominids from Asia attributed to H. erectus were said to repre-sent a side branch of the human family tree, unrelated in any lineal sense tomodern Asian or non-Asian people (Gibbons 1997). The new taxon of H.antecessor, however, does not differ in any substantial way from previouslyknown Asian fossils attributed to H. erectus. In fact, the material fromAtapuerca-Gran Dolina is further evidence that H. erectus was a widespread,pandemic, polytypic human species (Asfaw et al. 2002), in the same fashionthat its inheritor, H. sapiens, has been and still is.

The diagnosis of H. antecessor as a distinct hominid species rests pri-marily on a number of craniofacial features, identified as “fully modern” incharacter, seen in the juvenile upperpartial face (ATD6-69) and associat-ed cranial remains (Bermúdez deCastro et al. 1997). These traitsinclude midfacial characters such asa coronally oriented infraorbital sur-face associated with a well-devel-oped canine fossa, and a horizontal,high rooted, inferior zygomaxillaryborder, as well as cranial vault fea-tures such as a double arched supra-orbital torus and a convex superiorborder of the temporal squama(Bermúdez de Castro et al 1997). Anumber of non-diagnostic dentaland mandibular traits are alsoincluded in the definition of the newspecies, some of which will be dis-cussed below.

The craniofacial features of so-called H. antecessor, which are saidto represent a unique combination of traits derived in the direction of laterEurafrican hominids, are, however, seen in the very same combination in atleast one Chinese fossil hominid attributed to H. erectus (i.e. Yunxian 2; fig.11)(Li and Etler 1992), while the specifically midfacial features said to distinguishH. antecessor from previously known Lower and Middle Pleistocenehominids are seen in all Chinese H. erectus fossils in which the relevant areais preserved (Etler 1996; Wang and Tobias 2000). Cranial vault features saidto distinguish H. antecessor are differentially seen in a number of H. erectusfossils from China (Etler 1996; Wu and Poirier 1995).

The Yunxian hominids consist of two virtually complete crania initiallydated to the Middle Pleistocene based on their stratigraphic position and fau-nal associations (Li and Etler 1992). Later research, including the recovery oftypical Lower Pleistocene mammalian taxa from the hominid bearing stratum(Etler and Li 1994; Li 1997), paleomagnetic studies (Yan 1993), and ESR dat-ing of associated mammalian dental remains (Chen et al. 1996), however, indi-cate a late Lower or early Middle Pleistocene provenance for the Yunxianhominids, making them more or less contemporaneous with the newAtapuerca finds. Although the two Yunxian crania have been subject to vari-ous degrees of plastic deformation, substantial portions of the midface, supra-orbital region, and basicranium are preserved intact and can serve as a basis ofcomparison with ATD6-69 and other Atapuerca-Gran Dolina specimens.

The midfacial morphology of the Yunxian crania has been described asfollows: “The Yunxian crania show features of the mid-face common to non-neandertal late archaic and early modern H. sapiens (for example, the face isflattened and orthognathic with moderate alveolar prognathism; the maxillahas a distinct canine fossa; the lateral part of the maxilla is oriented coronallyand highly angled to the zygomatic; there is a high origin of the zygomaticroot; a horizontal inferior zygomaxillary border and a pronounced malarincisure, and so on)” (Li and Etler 1992). This description, in all its particulars,is identical to that of ATD6-69. Thus, the midfacial morphology identified byBermúdez de Castro et al. (1997) as “fully modern” and which they suggest“antedates other evidence of this feature by 650,000 years” was described ina penecontemporary human specimen from China five years prior to theirdescription of the Atapuerca juvenile! Other features mentioned by Bermúdezde Castro et al. (1997) as being in unique combination in the Atapuerca

remains, such as a double archedsupraorbital torus and a convexsuperior margin of the temporalsquama are also seen in theYunxian 2 cranium, as well asother specimens from Chinaattributed to H. erectus, includingLantian (double arched supraor-bital torus); and Hexian, Nanjing,and Skull V from ZhoukoudianLocality 1 (convex superior mar-gin of the temporal squama) (Wuand Poirier 1995; Etler 1996).There is, therefore, nothingunique about the morphologicalpattern of the Atapuerca remainsthat warrants creation of a newhuman species.

When the Yunxianremains were initially describedgreat emphasis was placed on the

modern character of their facial topography, as facial remains of H. erectuswere, and still are, relatively rare. It was stressed that modern human facialanatomy seemed to occur earliest in Asia, while the more obliquely set ances-tral hominid pattern persisted in the West, eventually giving rise to the derivedNeanderthal pattern seen in the Late Pleistocene. This conclusion must nowbe re-evaluated in light of the new Atapuerca-Gran Dolina specimens.Modern human facial topography should be recognized as a polymorphiccondition of H. erectus throughout its range.

Equal weight in assessing the phylogenetic affinities of the Yunxian cra-nia was also given to their basicranial morphology, a portion of the human cra-nial anatomy not sampled at Atapuerca-Gran Dolina. In this regard it wasobserved that both Yunxian crania preserve a morphology identical to thatseen in other specimens of Asian H. erectus. Li and Etler (1992), therefore,while noting the “modern” features of the Yunxian face and certain similari-ties with archaic western hominids in cranial vault morphology, attributed theYunxian specimens to a local variant of H. erectus and stressed the polytypicnature of that taxon.

The Atapuerca-Gran Dolina remains show that populations similar toYunxian and Lantian in mid-facial and cranial vault morphology were wide-spread throughout the northern hemisphere during the Lower and early MiddlePleistocene. In addition, a comparison of the Atapuerca-Gran Dolina mandibu-

Fig.11:Yunxian vs. Atapuerca-Gran Dolina mid-facial anatomy. Features of the ATD6-69 juve-nile mid-face from Atapuerca (right) are said to have a fully modern facial topography including“a prognathic (nonflat) mid-face, a well-developed canine fossa, a horizontal zygomaxillary bor-der and a sharp lower nasal margin” (Bermúdez de Castro et al. 1997). These very same featurescan be seen in the Yunxian 2 face (left) and in material of H. erectus from Zhoukoudian andNanjing.(photo: Li Tianyuan, J.M. Bemudez de Castro).


tively complete occipitals, the ramus and poste-rior body of a lower jaw, and a juvenile maxillawith intact dentition, all associated with a prolif-ic archaeological assemblage that includes proto-prismatic cores (Wu and Poirier; Etler and Li1994). If this material was found in Africa itwould surely not be so easily dismissed. In fact,Bräuer’s characterization of the Xujiayaoremains would appear to be equally applicable tomaterial such as that from Klasies River Mouth(which consists of a lower jaw, a fragmentaryfrontal bone, pieces of parietal, a zygomaticbone, and some isolated teeth), if not for the factthat the latter is so central to the argument formodern human origins in Africa.

Rather than dismissing the Xujiayao mater-ial, which has generally been attributed to latepre-modern H. sapiens in China, it needs to bere-evaluated in the light of new Chinese discov-eries, particularly the as yet undescribed skullfrom Laishui dated between 60-30 kya. TheLaishui cranium, a cast of which is on public dis-play at Peking University’s Sackler Museum ofArchaeology, has extremely robust supraorbitaltori that preserve an archaic morphology, a rela-tively sloping forehead, along with a robust butessentially modern cranial vault, cranial base,and face (Etler 1996; Bräuer 2001). This speci-men, as well as others from the late UpperPleistocene in China, such as a calva fromHuanglong in Shanxi and an occipital fromShiyu in Shanxi, shows features highly reminis-cent of those seen in homologous material fromXujiayao (Etler 1996). How modern specimensfrom Salawusu in Inner Mongolia, dated tobetween 65-35 kya, fit into the emerging pictureof Upper Pleistocene human evolution in Chinaalso requires further study. It can be argued, how-ever, that the earliest modern Chinese fossilsshow a higher retention of archaic features thananalogous material from further west.

While data are still limited, there is everyreason to believe that more fossils representingthis crucial period of human evolution in Chinawill eventually be found (contra Jin and Su2000). It is therefore very premature to dismissthe possibility of early modern humans in EastAsia having an antiquity equivalent to that seenin Africa and the Middle East. Verdict: The finaldisposition of Condition 2 is still pending, but allindications are it will be verified sooner ratherthan later.

Condition 3: There should be a lack of anymajor discontinuity in the human fossil record ofChina. Here we are on firmer ground. Advocates

of ROA originally held that Asian H. erectusandits late Middle Pleistocene descendants sharedautapomorphies that set them apart from westernarchaics and early moderns as an evolutionarydead-end, implicitly recognizing continuitybetween earlier more “primitive” and later more“advanced” pre-modern human populations inChina. (This assessment of the evolutionary rela-tionship between Asian H. erectus and laterarchaic humans in China such as Dali andJinniushan also flies in the face of the proposi-tion, mentioned above, that the latter specimensshow greater affinities to western archaics thanH. erectus.) Supporters of MRE, of course, havelong documented many features shared by thesetwo groups of fossils (Wu and Poirier 1995; Etler1996; Etler and Li 1994). It seems fair to state thatcontinuity between H. erectus and “archaic” H.sapiens in China has been generally accepted.Those features that differ between the twogroups are equivalent to the differences thatROA supporters would claim separate H.ergaster from H. heidelbergensis in Africa; i.e.they are transitional in nature.

The question then arises as to whether thereis any discontinuity between late pre-modernChinese and early moderns. The answer to thisquestion revolves around whether one accepts orrejects the validity of the trait complexes relatingto East Asian facial and dental morphology men-tioned earlier as local evolutionary markers inEast Asia. I would argue that their presence inerectine, pre-modern, and early modern speci-mens in China speaks to a definite degree ofmorphological continuity, and certainly gives noindication of discontinuity, between antecedentpre-modern and subsequent early modern popu-lations in East Asia. Verdict: It can be concludedthat there is no evidence for derived traits inChinese archaics that would preclude them frombeing ancestral to later moderns.

In conclusion, of the three conditions usedby advocates of ROA to demonstrate continuity,two are as demonstrable in East Asia as inAfrica: there are transitional fossils during the lateMiddle Pleistocene in China, and there is no evi-dence of discontinuity between Chinese archaicsand early Chinese moderns. The third prediction,the presence of early moderns during the earlyUpper Pleistocene, while not yet well docu-mented, has not been disproved. Moreover, thecontinuity of at least two significant trait com-plexes between late archaics and early modernsin China gives weight to the third prediction’seventual verification. It can therefore be argued


lar morphology as described by Bermúdez deCastro et al. (1997), with the penecontemporary H.erectusmandible from Lantian, Chenjiawo,shows complete identity in all salient features.Personal observation of the Chenjiawo mandibleat the IVPP in Beijing, China confirms that it hasa mylohyoid groove that “extends anteriorlynearly horizontal and courses into the mandibu-lar body as far as the level of M2/M3” and otherfeatures identified by Bermudez de Castro et al.(1997) as mandibular traits defining H. anteces-sor as distinct from their conception of H. erec-tus. The fact that the Atapuerca-Gran Dolinaremains also show evidence of shovel-shapedupper incisors and reduced mandibular M3s,common features of Asian H. erectus and alllater fossil and living Asians (Liu and Zeng1996; Etler 1996), is further evidence that thereis, in fact, no such thing as so-called Asian H.erectus in contradistinction to other purportedlate Lower - early Middle Pleistocene species ofHomo. On the contrary, the Atapuerca speci-mens more than adequately demonstrate that H.erectus was a wide-ranging, polytypic humanspecies ancestral to all later forms of humanity.H. erectuswas therefore not a side branch of thehuman family tree but part and parcel of its maintrunk.

In conclusion, as the Yunxian and Lantianspecimens are, in broad terms, contemporaneouswith the Atapuerca remains, and as they cannotbe adequately differentiated from one anothermorphologically, they can best be evaluated phy-logenetically as representative of a broadlydefined H. erectus, which served as the base forlocal evolutionary developments in Europe,Africa, continental East Asia, and insularSoutheast Asia. Furthermore, neither Atapuerca-Gran Dolina nor Yunxian have any direct bear-ing on the question of modern human origins,other than to demonstrate that late LowerPleistocene - Middle Pleistocene Homo had notyet differentiated to any considerable extent intoregional or species specific variants. Moreover,the similarities between the Atapuerca speci-mens, Yunxian, and Lantian (both atGongwangling and Chenjiawo) suggest thatspecimens of H. erectus as known fromZhoukoudian should not be misconstrued as dis-playing the “typical” or most widespread com-bination of H. erectus traits. In fact, these formerspecimens probably represent the nominal, gen-eralized morphology of H. erectusbetter than themore specialized specimens from Zhoukoudian,the description of which has served as the defacto basis for the definition of H. erectus eversince Weidenreich’s classic monographs on“Peking Man” in the 1930s and 40s(Weidenreich 1936, 1937, 1943).


that based on the fossil record continuity betweenarchaic and modern humans in East Asia is avery viable option. There is, in addition, othernon-fossil evidence that supports MRE, bothgenetic (see above) and archaeological (seebelow).

Archaeological evidence:Lahr (1994), as aspokesperson for the hard ROA position, arguesthat archaeological data can be used to buttressthe case for replacement of archaics by modernsin Europe, although it is conceded that the evi-dence is more equivocal in Africa and theMiddle East. As regards East Asia she states,“the lack of material and objective informationabout East Asian and Javanese technologiesdoes not allow the inclusion of these importantregions in the discussion, although the appear-ance of Upper Palaeolithic in Siberia at approxi-mately 35 kya suggests population movementinto this area.”

This statement by Lahr is characteristic ofthe selective vision of many supporters of ROA.Rather than confront an issue that is not in accordwith their a priori assumptions about modernhuman origins in Africa, they dismiss it out ofhand. To say there is a lack of material and objec-tive information about Paleolithic industries andtechnologies in East Asia is to negate over 70years of archaeological research in China and, inparticular, the last 50 years of intensive work byChinese archaeologists not only at the IVPP inBeijing, a world-reknowned paleontological andpaleoanthropological institution, but also thework of countless archaeologists at provincialand local levels throughout China, who haveaccumulated a wealth of data pertaining to theChinese Paleolithic. Many summaries of thesefinds and researches are available in English,written by both Chinese and foreign scholars

(Aigner 1981; Wu and Olsen 1985; Clark andSchick 1988; Ling 1996; Zhang 1999; Hou et al.2000; Keates 2000; Leng 2001).

What the Chinese Paleolithic shows is thatit developed, to a large extent, independentlyfrom outside influences. There is no evidence ofPaleolithic industries associated with the West tobe found in China, and attempts to identify cer-tain assemblages as “Acheulian,” “Mousterian,”or “Solutrean,” etc. were abandoned decades ago(Aigner 1981). This is not to say that varioustechnological modes identified in the West werenon-existent in China (see Hou et al. 2000). It isclear, however, that the Chinese Paleolithic, up toand including the Upper Paleolithic, must beseen in its own context (Clark and Schick 1988),a fact which does not meet the expectations of a“replacement” model for the origins of modernEast Asians, but is fully in accord with MRE.

There is one final point to be made regard-ing the assertion by many advocates of ROA that“a number of early modern specimens fromChina and Australasia exhibit basic similaritieswith early modern humans from Europe and

even Africa” (Bräuer 1992; see also Kamminga1992; Brown 2002), the implication being thatthis points to an external source of origin for earlymoderns in the East. This observation is, howev-er, totally in accord with the core concept ofMRE presented in this paper. If there was a “cen-tral zone of sapienization” extending from Africathrough the Middle East to East and SoutheastAsia during the late Middle Pleistocene/earlyLate Pleistocene transition, with heightened con-tact due to increased multilateral dispersals andconcomitant genetic communication, it shouldbe expected that all early moderns, be they fromAfrica, Europe, or Asia, would share in certain“grade” features that would tend to unite themphenetically to the exclusion of present-day pop-ulations in their respective regions of habitation.

Conclusion: In conclusion, evidence ismounting that the emergence of modern humansduring the Late Pleistocene was characterized bya “multi-causal and complex demographicprocess” and “a complex migration and mixingprocess, variable degrees of continuity in variousregions of the world, and strong ties betweeneastern and western parts of the Old World”(Brauer 1992). A similar conclusion wasreached from a study of the Yunxian crania, fromHubei province, China (Li and Etler 1992), inwhich it was noted that, “the differential distribu-tion of character states associated with H. sapiensin regionally disparate Middle Pleistocenehuman populations suggests that the events lead-ing to the emergence of modern humans werenot restricted to one region of the world alone. Inaddition, the mix of characters in the Yunxiancrania demonstrates that the taxon H. erectus isfounded on a set of ancestral hominid traits andregional polymorphisms. It hence has no mean-ing in a cladistic framework. In light of the aboveconsiderations we feel it is best to view all


Fig.12: a. (Asia): Dali, China; b: (Europe): Saccopastore, Italy; c. (Africa): Jebel Irhoud 1, Morocco.The threeexamples show regional variation in human cranial form and structure 120,000 years ago. Note the overall sim-ilarity between the Dali (a) and Irhoud (c) crania, in particular the flat and broad mid-face, squared eye sockets,and keeled braincase. This differs significantly from the Neanderthal (c) cranium from Europe, which has a nar-row projecting mid-face, rounded eye sockets, and globular braincase. In other features, the Dali cranium is sim-ilar to Saccopastore, in particular the broad shape of the nose and guttering of the lower border of the nasal open-ing. The Dali cranium also retains many H. erectus-like features seen in earlier human fossils from China.

Fig.13: Comparison of Yunxian 2 with archaic and modern human crania. Two hominid crania from Yunxian,Hubei province, China that have a mix of erectine and archaic features have recently been dated to the earlyMiddle Pleistocene between 800,000 - 560,000 ya. (Chen et al. 1996, 1997; Yan 1993). This scan of the secondYunxian cranium shows its overall similarity to the archaic Petralona specimen from Greece (center). Yunxian 2nevertheless retains ancestral features of the cranial vault and base not seen in the later more derived Petralonaspecimen. Cranial capacity of Yunxian 2 is estimated at no more than 1100 cc.

a cb


Middle Pleistocene hominids in a broad per-spective as an essential part of one evolving lin-eage in direct ancestry to modern humans”(figs.11-13; see box 5).

From a genetic perspective, the significantrole of African dispersals cannot be denied, butas Templeton (2002) has recently stated, “Acoherent picture of recent human evolutionemerges with two major themes. First is thedominant role that Africa has played in shapingthe modern human gene pool through at leasttwo - not one - major expansions after the origi-nal range extension of Homo erectus out ofAfrica. Second is the ubiquity of genetic inter-change between human populations, both interms of recurrent gene flow constrained by geo-graphical distance and of major populationexpansion events resulting in interbreeding, notreplacement.” It must be acknowledged, more-over, that evidence for ancient multilateral dis-persal events could very well have beenobscured by the recent population history of ourspecies. The resources of the human genome are,however, just now being fully accessed. Muchnew information will surely come to light thatwill better document the course of human genet-ic evolution. The fossil record of human evolu-tion will also continue to accumulate, likewiseshedding new light on our origins and the role ofH. erectus in human evolution.

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Author’s biographical note:

Dennis Etler has spent the past two decadesconducting paleoanthropological research inChina, working in collaboration with Chinesepaleoanthropologists and Paleolithic archaeolo-gists. He has been involved in a number of pro-jects that helped open the door to foreign partici-pation in paleoanthropological and archaeologi-cal research in China, including the descriptionand analysis of early Middle Pleistocene humancrania from Yunxian, Hubei; the joint Chinese-American study and excavation of earlyPaleolithic sites in the Nihewan basin of Hubei;

the joint Chinese-American study of fos-sil hominoid remainsfrom the Yuanmoubasin, Yunnan and there-interpretation of fos-sil hominoid speci-mens from the

Longgupo site in Sichuan. He has publishednumerous scientific articles that have helpedbring the rich fossil record of Chinese hominidsto the attention of the world-at-large.

Presently Dennis teaches PhysicalAnthropology at Cabrillo and Gavilan collegeson the central coast of California, where hemakes his home in Santa Cruz.He continues hisinterest in Asian paleoanthropology and hosts awebsite on the “Fossil Evidence for HumanEvolution in China,” that can be accessed atwww.chineseprehistory.org.


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Glossary:agenesis: failure of a biological structure todevelop.

alveolar: tooth-bearing portion of the upper jaw.

alveolar prognathism: forward projection of themaxilla, or upper jaw.

anagenesis: one form of speciation in which aspecies evolves through time into a new species.Anagenesis is contrasted with cladogenesis (sin-gle, continuous line vs. distinctive branching,respectively).

anterior fovea:depression in forward part of tooth.

autapomorphy: new morphological trait uniqueto a group in an evolutionary lineage; uniquelyderived feature.

autosome: any chromosome except a sex chro-mosome.

basicranium:part of the occipital bone at the bot-tom of the skull.

bauplan: The generalized, idealized archetypicalbodyplan of a major taxon.

buccal: relating to the cheek.

calva: uppermost portion of the braincase(equivalent to calotte).

canine fossa: a depression above the caninetooth on the cheek of modern humans resultingfrom the reduction in the size of the face.

cladistics: the school of evolutionary biology thatseeks to make evolutionary hypotheses throughinterpreting patterns of primitive and derivedcharacteristics; a method for determining the evo-lutionary relationships among animal groups bycomparing shared morphological features. Theserelationships are usually depicted in a cladogram.

cladogenesis: one form of speciation in whichone species evolves through time into two ormore descendant species. Cladogenesis is con-trasted with anagenesis.

coronally oriented infraorbital surface: The sur-face of the face below the eye orits is on a flatplane horizontal to the midline of the skull.cranial vault: combination of bones that encasethe skull.

cultural hitchhiking: co-evolution of culture andgenes; reduction in genetic diversity when neu-tral or nearly-neutral genes (such as mtDNA)and selectively advantageous cultural traits aretransmitted in parallel.

cusps: pointed or rounded bumps on the occlusalor chewing surface of a tooth.

deme: a population, or cluster of individuals,

with a high probability of mating with each othercompared with their probability of mating with amember of some other population, and thereforehaving a substantial amount of genetic exchange.

electron spin resonance (ESR) dating: datingmethod that measures the concentration of elec-tron traps in a material, which accumulatethrough natural radiation; method yields a mini-mum, rather than actual, age.

exogenous: originating from outside the organ-ism or system.

gene flow: the movement of genes into orthrough a population by interbreeding, or migra-tion and interbreeding, with another population.

grade: a level of organization based on the pres-ence of common biological features and used inassessing diffferent evolutionary lines of animals.

haplotype: combination of genetic markers orpolymorphisms present in a gene or genome, suchas the mtDNA, that are inherited together as a unit.

hypoconulid:one of several cusps on a lower tooth.

malar incisure: notch on the cheekbone (malar isequivalent to zygomatic).

mandible: lower jaw.

maxilla: upper jaw.

metaconid: one of several cusps on a lower tooth.

mitochondrial DNA (mtDNA): DNA in mito-chondria of cells, rather than nuclei. Becausemitochondria occur outside the cell of the nucle-us, mtDNA is typically only passed on throughthe female, and hence reveals matrilineal rela-tionships. Rate change for mtDNA is much fasterthan DNA of the nucleus, making it suitable foranalyses of recent evolutionary developments.

mylohyoid groove: longitudinally running grooveon the internal surface of the mandible (lowerjaw), where the mylohyoid muscle (flat, triangularmuscle forming the floor of the mouth) attaches.

non-coding: DNA sequence not involved incoding for a protein end-product.

non-recombining: those portions of the humangenome that do not combine, including mtDNAand the Y-chromosome. Because they are non-recombining, the mutations that occur in thesesegments of DNA are not reshuffled or mixedinto new combinations, and hence, accumulatein a linear or chronological fashion.

nuclear genome: full set of chromosomes, andcorresponding inheritable traits, from the nucle-us of a cell.

occlusal wear: wear on the chewing surface of atooth.

orthognathic: without forward projection of theupper or lower jaw; strait-jawed.

paleomagnetism: dating method based on theperiodic reversals of the earth’s magnetic polarity,often employed in dating of early hominine sites.

phylogeny: the study of evolutionary developmentof a species, often expressed with a family tree.

plesiomorphic: said of a trait that is an ancestralretention.

polymorphism: genetic variability for a trait.

polytypic:comprising several subspecies or geo-graphical variants.

population bottleneck: reduction in populationsize that may cause changes in gene frequencywithin a population.

protoconid: one of several cusps on a lower tooth.

proto-prismatic cores: blocks of lithic materialprepared in such a fashion as to produce an elon-gated flake when struck ancestral to UpperPaleolithic cores from which blade-like flakeswere struck.shoveling: characterized by upper incisor teethreinforced with extra enamel at their edges of thetongue side.

speleothem: a mineral deposit formed in a caveby the action of water.

supraorbital torus: ridge above the orbits on askull, very pronounced in H. erectus,Neanderthals, and some australopithecines (fromsupra: above; orbital: eye; torus: ridge).

talonid basin: feature on a lower tooth.

temporal squama: a portion of the temporalbone, which is located at the side of the skull nearthe ear, the squama is flat or fanlike in modernhumans and projects upward and forward.

trait complexes: group of attributes or characterswithin a species for which heritable differencescan be defined.

uranium series (U-series) dating: dating methodbased on the decay of a number of isotopes ofuranium.

Y-chromosome: a chromosome unique to males,on which sex-determination genes are located,consisting largely of non-recombining sequence.Genetic variation on the Y-chromosome canshow inheritance of specific genes or mutationsthrough the paternal line, and also track malemigrations through time and space.

zygomaxillary border: meeting place (or suture)where the zygomatic (cheekbone) meets themaxilla (upper jawbone).


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