Kroeber Anthropological Society Papers, Nos. 71-72, 1990

A Case Study of the "erectus" - "sapiens" Transition in Asia: Hominid Remains fromHexian and Chaoxian Counties, Anhui Province, China

Dennis A. Etler

Newly discovered fossil hominidsfrom the late middle Pleistocene of China demonstrate that archaicforms of Homo sapiens were either coexistent with or slightly more recent than advanced forms ofHomo erectus. This is most clearly seen at sites in Hexian and Chaoxian counties, Anhui province,which date to between 150-200,000 YBP by uranium series tests. The replacement of H. erectus byarchaic H. sapiens in China is characterized by the retention of a significant component of heritagefeatures in archaic H. sapiens, overlaid by a mosaic of changes in craniofacial anatomy that trendtowards modern humans. This new evidence gives powerful support to the regional continuity theory ofhuman evolution in east Asia. Furthermore, it suggests that the transitionfrom H. erectus to archaic H.sapiens in China was relatively quick and possibly modulated by a heightening of gene flow betweenwestern and eastern Asia at approximately 250,000 YBP.

INTRODUCTION

The accelerated pace of discovery of latemiddle Pleistocene - early late Pleistocene humanfossil remains in China over the last two decadesmakes it possible to approach the question of the"erectus" - "sapiens" transition in east Asia froman entirely new empirical basis. Human skeletalremains from this period of time includeadvanced forms ofHomo erectus from Zhoukou-dian (Skull V) (Qiu et al. 1973) and Hexian(Huang et al. 1981, 1982; Wu and Dong 1982),and cranial material attributed to archaic Homosapiens from Changyang (Jia 1957), Maba (Wuand Peng 1959), Xujiayao (Jia and Wei 1976; Jiaetal. 1979; Wu 1980), Dali (Wu 1981), Yingkou(Lu 1985, 1989; Wu, R. 1988) and Chaoxian(Xu et al. 1984, 1986a, 1986b) (Figure 1).1 Ofparticular note are the hominid remains fromLongtandong, Hexian county and Yanshan,Chaoxian county, Anhui province which aretaken to represent H. erectus and archaic H.sapiens, respectively. Not only are these twosites in close spatial proximity (Figure 2), theymay also be close in age. The Hexian fauna(Tables 1 and 2) has been correlated with theupper layers at Zhoukoudian (ZKD) and oxygenisotope Stage 8 which would give an age of 240-280,000 years before present (YBP) (Xu andYou 1984). The Chaoxian fauna (Table 2) isthought to be somewhat younger due to the ab-sence of archaic species such as Megantereon andTrogontherium which are encountered at Hexian.Otherwise, the faunas both reflect a similar mix-ing of northern and southern elements.

Recent uranium series dating of the hominidbearing strata at both sites, however, raises thepossibility that they are virtually synchronousand, moreover, somewhat younger than the up-

per layers at ZKD Locality 1 (Chen et al. 1987;Chen and Yuan 1988). The uranium seriesanalyses, based on the internal concordance of230Th/234U and 231pa/235U activity ratios ofbones and teeth associated with the humanremains, date the Hexian site to between 150-190,000 YBP and the Chaoxian site to between160-200,000 YBP. While uranium series datesare still subject to considerable controversy, thetechniques employed to date the organic remainsare identical at both sites, and the physical condi-tions encountered at Longtandong and Yanshanare similar. It is, therefore, reasonable to acceptthe dates as being broadly comparable. Nonethe-less, given the vagaries of uranium series datingthese ages should best be taken as minimum agesfor both sites. The close spatial and possibletemporal proximity of the Hexian and Chaoxianhominids coupled with their morphological dif-ferences (vide infra) raise obvious questionsconcerning their relatedness to one another. Inaddition, the suggested late age of occurrence ofH. erectus at Hexian raises questions about thetemporal span of H. erectus in east Asia, and themode and tempo of its replacement by archaic H.sapiens. In order to more fully appreciate theimplications of these new finds for an under-standing of the course of human evolution in eastAsia, this contribution will concentrate on descri-bing and interpreting the human fossil remainsfrom both Hexian and Chaoxian.

While I had the opportunity to view theHexian calvaria during a recent visit to China, Ihave not seen the material from Chaoxian, norhave I had the chance to personally study thespecimens in question. I have, however, availedmyself of the Chinese descriptions of the spe-cimens which are quite detailed in extent andwhich follow the format and termiinology of Wei-

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denreich's (1937, 1943) classic descriptions ofthe hominid remains from Zhoukoudian. Muchof the analyses presented in the Chinese accountshas not been previously available in English. Ihope that this presentation alleviates, to an extent,this circumstance, thereby facilitating discussionof these important hominid remains by a westernaudience.

HEXIAN COUNTY

The Longtandong cave site located on Wang-jiashan, a hill near Taodian village, Hexiancounty, Anhui province (E 1180 20', N 310 45')(Huang et al. 1981, 1982; Wu and Dong 1982;Wu 1983; Zheng 1982; Zheng 1987; Xu and You1984; Chen et al. 1987) has produced the mostprolific remains of H. erectus unearthed in Chinasince the height of excavations at ZKD in the1930's. An inventory of recovered specimensincludes: a nearly complete calvaria (PA 830), the

supraorbital portion of a frontal (PA 840), a rightparietal (PA 841), a fragment of a left mandibularcorpus retaining M2 and M3 (PA 831), a rightupper P4 (PA 832), a left upper M2 (PA 833), aleft lower Ml and M2 of one individual [PA 834(1-2)], a right upper I1 (PA 835), a left upper Ml(PA 836), a right upper M2 (PA 837), and twoleft lower M2's (PA 838 and PA 839).

The site has been known since 1973 when itwas blasted open during canal construction. Thecave was found to be richly fossiliferous and anumber of vertebrate specimens were sent to theInstitute of Vertebrate Paleontology and Paleo-anthropology (IVPP), Academia Sinica for study.In 1979, at the request of the Anhui ProvinceWater Conservancy Bureau, a team led by HuangWanpo was dispatched by the IVPP to furtherinvestigate the site. More fossil vertebrates werecollected along with bone that has been claimed tobe artifactual in nature (Huang et al. 1982).Excavations were resumed in January 1980. InJuly the first human fossil, an upper molar, was

Figure 1. Localities in China yielding late middle Pleistocene or early late Pleistocene fossil hominids.

1. Hexian, Anhui 2. Chaoxian, Anhui 3. Zhoukoudian, Beijing 4. Xujiayao, Shanxi 5. Dingcun, Shanxi6. Dali, Shaanxi 7. Yingkou (Jinniushan), Liaoning 8. Miaohoushan, Liaoning 9. Changyang, Hubei10. Jiande, Zhejiang 11. Maba, Guangdong 12. Tongzi, Guizhou

3

Figure 2. Location of the Hexian and Chaoxian sites.

recovered by Qin Wanju of the Anhui Water Con-servancy Bureau from previously excavated backdirt within the cave. Later that fall during thethird excavation the calvaria and other human fos-sils were found. Further excavations led to therecovery of additional teeth and cranial remains(Wu 1983).

Longtandong is located on the northernslope of Wangjia Hill, 30 km northwest of theYangtze River at an elevation of 23 meters abovesea level. The deposits within the cave can be di-vided into four layers. The hominid materialcomes from the second stratigraphic level whichconsists of 0.5-1.0 meters of brown to yellowclays (Huang et al. 1981, 1982) (Figure 3). TheHexian fauna produced from the cave is bio-geographically transitional, containing bothpaleoarctic and subtropical elements (see Tables 1and 2). The micro-mammalian componentshows considerable faunal mixing with northern,southern and "alpine" species seen frequently inthe western montane region of China all present(Zheng 1982; Xu and You 1984). Palynologicalsampling at Longtandong likewise indicates thepresence of both northern and southern taxa(Huang and Huang 1985). Since the 1950's ithas been anticipated that this area of the NorthChina Plain, situated as it is between the Chang-jiang (Yangtze) and Huanghe (Yellow) Rivers,would produce a transitional faunal complex con-taning both northern and southern taxa. Pei

(1957) referred to the area in question asthe Huai River Transitional Zone. Themammalian fauna from Sihong in JiangsuProvince, now known to be of Mioceneage, was initially thought to be repre-sentative of just such a Pleistocenetransitional zone. It was later shown,however, that castorid and proboscideanremains at Sihong had been misinter-preted and were not of Pleistocene age (Liand Zhou 1978). The faunal remainsfrom both Hexian and Chaoxian are thefirst verification of the Huai River Transi-tional Zone and have contributed greatlyto a better understanding of the paleobio-geographic zonation of east Asia duringthe middle and late Pleistocene (Liu et al.1982; Zheng 1987).

HUMAN REMAINS FROM HEXIANCOUNTY

U The Hexian calvaria (Figure 4) hasbeen studied in detail by Wu and Dong

_ (1982). The following account of thespecimen is based on their analysis.

Preservation

The heavily fossilized specimen wasoriginally fragmented into eight pieces. Recon-struction revealed a well-preserved calvaria thatretains a substantial part of the cranial base. Therestoration consists of: 1) a nearly complete fron-tal including the supraorbital region, but lackingthe inferior portion of the nasal process; 2) anearly complete left parietal; 3) the right parietalsomewhat damaged at pterion, near the parietaleminence and asterion; 4) a left and righttemporal, both lacking zygomatic and mastoidprocesses due to breakage -- the left temporal isrelatively complete, the right temporal squamahas been lost at the sphenoidal margin and alonga portion of the parietal margin; 5) an occiput thatpreserves the posterior margin of the foramenmagnum and much of the nuchal and occipitalplanes; and 6) the lateral portion of the greaterwing of the left sphenoid. The vault between thetemporal lines anterior to the vertex and posteriorto bregma is flattened. It has been suggested thatthis may be due to compressive forces duringfossilization (Wu and Dong 1982).

The relatively large and robust constructionof the calvaria (cranial capacity approximately1025 cc), the thick and well-developed supraorbi-tal and occipital tori, the well-expressed tempo-ral and nuchal lines and the sloping forehead

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Table 1. Taxonomic list of non-mammalian and micimammalian vertebrates from Longtandong, Hexicounty, Anhui province (after Huang et al. 1981, l9fZheng 1982; Xu and You 1984; Huang and Hua1985).

REPTILIA

Ocadia sp.Amyda sp.Alligator cf. sinensis

AVES

Crossoptilon sp.

MAMMALIA

InsectivoraBlarinella quadraticauda (Milne-EdwardChodsigoa youngi Huang et al.Anourosorex squamipes Milne-Edwards?Scaptochirus sp.

ChiropteraRhinolophus cf. ferrum-equinum Thomas?Myotis sp.Hipposideros sp.Miniopterus schreberii Kuhl

RodentiaTrogontherium cuvieri FischerCricetulus varians (Zdansky)Apodemus argrarius PallasRattus rattus (Linnaeus)Rattus norvegicus BerkenhoutRattus edwardsi ThomasMicrotus brantioides YoungEothenomys melanogaster Milne-EdwardsEothenomys inexEothenomys evaEothenomys proditorTamias wimani Young

combine to suggest a male specimen. Wu andDong (1982) judge it to be a young adult, as bothexternal and internal sutures (excepting the leftsphenoidal suture) are open.

Lateral Aspect

Wu and Dong (1982) describe the calvaia aslow vaulted with the supraorbital and occipitaltori being respectively the most anterior andposterior projecting points on the skull, so thatgreatest cranial length (g-op) is coincident withthe distance between glabella and inion (i.e.,inion coincides with opisthocranion). The tem-poral lines are well-expressed and separated by

ro- 77 mm at their closest approximation.ian The posterior course of the temporal line82; terminates in a distinct angular torus at theng mastoid angle of the parietal. The above

features are all characteristic of previouslyknown crania of H. erectus from ZKD.The pterionic region, preserved on the leftside, is of the sphenoparietal type andassumes an "H" shape. The sphenoparie-tal suture is 11 mm long (Wu and Dong1982).

The temporal squama is relativelyhigh and arched superiorly at the parietalmargin unlike the condition in Pekingman (with the exception of Skull V) inwhich the squamosal margin is low andstraight. The left squamous portion is 70mm long and 42 mm high, yielding a

s) length/height index of 60, compared to anaverage of 49.6 in Peking man (Weiden-reich 1943), 64.6 in the Dali cranium (Wu1981) and an average of 65.2 in modernman (Wu and Dong 1982). The Hexiantemporal, therefore, approximates that ofmodern humans in both shape and size.The parietal notch separating the temporalsquama from the mastoid region is deeplyincised. The root of the zygomaticprocess of the temporal slants posterosu-periorly, merging with the supramastoidcrest which in turn unites with the angulartorus. The sulcus processus zygomatci,which positions the zygomatic processlaterally away from the squama, is wideand shallow rather than narrow and deepas in modern man. The supramastoid andmastoid crests are well-developed as isthe supramastoid sulcus. The digastricfossa is broad, shallow and opens upposteriorly, differing from the deeper andmore narrow incisura mastoidea seen inmodern humans. What is preserved of the

mammillary process of the mastoid is relativelysmall and, as in other specimens of H. erectus,lies below the level of the supramastoid crest. Itextends further backwards than in modern man.The configuration of the external auditory meatusand the overhanging suprameatal tegmen is as in"Sinanthropus" (see Weidenreich 1943:53). Theaperture of the meatus is verdcally elliptic. Thepostglenoid process is poorly expressed as a lowprotuberance.

Vertical Aspect

The frontal has a low and sloping squamousportion. The frontal profile (m-g-i) (Martin 1928:

5

639, No. 32a) measures 580 and the inclination ofthe frontal squama (b-g-i) [Martin 1928:640, No.32(2)] measures 410. These measures are consi-derably less than in the early H. sapiens Dalispecimen in which the above angles measure 720and 500 respectively (Wu 1981), but within therange of variation seen at ZKD (Weidenreich1943:109) (Table 3).

The supraorbital torus is wide laterally andthick inferosuperiorly. Although the superiormargin of the glabellar region is slightly de-pressed, the left and right supraorbitals unite in asingle body. Thickness measures of the supraor-bital torus are as follows: left medial 19 mm, leftmedian 16 mm, left lateral 12 mm; right medial18 mm, right median 17 mm, right lateral 13 mm.

Greatest thickness is medial and least thickness islateral as in specimens of H. erectus from ZKD.The frontal squama above the supraorbitals has apronounced "bump-like" protuberance as in mate-rial from ZKD (Weidenreich 1943:225). Thisdiffers from Javanese specimens of H. erectus,which show a flat frontal squama. There is a dis-tinct post-toral sulcus in the Hexian calvaria but itis much more weakly expressed than in similarspecimens from ZKD. Javanese specimens lackthis structure entirely as the flat squamous portionof the frontal merges smoothly with the supra-orbital torus.

The frontal eminences are only slightlyexpressed and thus are similar in degree to malespecimens from ZKD. A metopic suture is evi-

Table 2. Taxonomic list of macro-mammalian vertebrates from Longtandong, Hexian county and theupper (hominid bearing) levels at Yanshan, Chaoxian county, Anhui province, China (after Huang et al.1981, 1982; Zheng 1982; Huang and Huang 1985; Xu et al. 1984, 1986a, 1986b).

FAUNA LOCALITIES

Hexian Chaoxian

PrimatesMacaca robustus Young

CarnivoraCanis sp.Cuon alpinus LinnaeusVulpes sp.Arctonyx collaris CuvierLutra sp.Hyaena brevirostris sinensis OwenMegantereon sp.Felis chinensis GrayPanthera pardus LinnaeusAiluropoda sp.Ursus arctos LinnaeusUrsus thibetanus kokeni (Zdansky)

ProboscideaStegodon orientalis Owen

PerissodactylaEquus sp.Tapirus sinensis OwenMegatapirus sp.Dicerorhinus sp.

ArtiodactylaSus lydekkeri ZdanskySus cf. xiaozhuCervus (Pseudaxis) grayi ZdanskyMegaloceros pachyosteus (Young)Hydropotes inermis SwinhoeElaphurus davidianus Milne-EdwardsBison sp.Caprinae

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dent from a point level with the frontal eminencesto bregma. An unfused metopic suture is alsoseen in Skull XI from ZKD (Wu and Dong1982).

Wu and Dong (1982) note that the breadth ofthe frontal is expanded over those from ZKD.Least frontal breadth (ft-ft) measures 93 mm,greatest frontal breadth (co-co) 118 mm. Bothmeasurements are nearly 10% greater than theaverage for ZKD calvariae (Weidenreich 1943:106). The degree of postorbital constriction canbe gauged by an index that measures maximumpostorbital constriction (minimum breadth of thefrontal behind the orbits) against the distancebetween the outer edges of the brow ridges (101mm and 111 mm respectively in the Hexianspecimen) (Wu 1981). In the Hexian specimenthis index stands at 91 versus a range of 80.7-82.9 in specimens from ZKD and 85.1 for theDali cranium (Wu 1981). This relatively minordegree of postorbital constriction is considered adistinguishing feature of the Hexian calvaria byWu and Dong (1982).

A sagittal keel is apparent along the mid-sagittal plane from a position level with the

frontal eminences to bregma at which point itgradually disappears. The sagittal keel in Pekingman is more prominent extending from a positionlevel with the frontal eminences to the obelionregion of the parietals. The robust constructionof the vault in Peking man is further reflected bythe presence of a cruciate eminence at the junctionof the coronal and sagittal sutures. The Hexianspecimen lacks this latter structure as well as theparasagittal depressions seen along the sagittalsuture in specimens of H. erectus from ZKD andJava (Wu and Dong 1982).

Greatest cranial width is low on the skull inline with the supramastoid crests. The oudine ofthe neurocranium at this level is ovoid. The pan-etal eminences are well-developed which may bean indication of the relative youth of the indivi-dual. Parietal foramina are not in evidence.

Occiput and Cranial Base

Wu and Dong (1982) state that the occipitaltorus is well-developed forming a continuous,smooth protuberance. The most projecting point

Figure 3. Stratigraphic profile of Longtandong Cave, Hexian county, Anhui.

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LongtanY I .2 3 3 .4 -"-21 5 i 6i

1. Talus deposit 2. Fossil boaing deposits 3. Brownish-red day4. Brownish-red day and sandy clay Intercalated with brecda5. HominId clvaria 6. Cambrian dolomite

A. Trial excavaton, Oct.1979 B. Excavation, Jan. 1980C. Excavation, Oct.-Nov. 1980

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Figure 4. The Hexian calvaria.

along the torus is at midpoint and the median por-tion is relatively straight while the lateral portionscurve inferiorly, diminishing towards asterion.There is a clear supratoral sulcus. The occipitalangle separating the occipital and nuchal planes isacute. The inferior margin of the torus forms aslightly arched line equivalent to the superior nu-chal line in modem man. It courses laterally,forming the lateral occipital crest. The externaloccipital protuberance is indistinct. A large inter-sutural bone is positioned on the right side atasterion.

In basal aspect a broad and shallow incisurasupraorbitalis is positioned medially on the lowermargin of the left and right supraorbitals. It isbound laterally by a processus supraorbitalis inthe fashion described by Weidenreich (1943:29).However, these structures are not as distinct as inthe ZKD material. Only the anterior surfaces ofthe orbital roofs are preserved, and they are dis-tinguished by their overall flatness. The frontalsinus is relatively small and is restricted to the up-per and lateral portions of the interorbital regionin the same manner as the majority of specimensfrom ZKD (cf. Weidenreich 1943:31) (left an-teroposterior length 7.5 mm, right anteroposteriorlength 11.5 mm, left width 14 mm, right width11 mm).

As in specimens of H. erectus from ZKD,the mandibular fossa is deep and narrow and thearticular eminence is weakly expressed (Weiden-reich 1943:47). Of particular note is the presenceof a deep groove medially along the anterior mar-gin of the right mandibular fossa. The size of thetympanic plate, its position in relation to the mid-sagittal plane of the cranium and its spatialorientation are all as described for "Sinanthropus"(Weidenreich 1943:52-57), i.e., it is very thick,positioned perpendicular to the mid-sagittal planeof the cranium (the plesiomorphic condition forhominoids -- in modem humans the tympanicplate is flexed posteriorly with reference to themid-sagittal plane) and oriented intermediatelybetween the vertical condition seen in modem hu-mans and the horizontal condition seen in apes.The angle between the tympanic plate and thepetrous process, however, is more acute in theHexian specimen (300) than in specimens knownfrom ZKD (50° in Skull III). In modem humansthe tympanic plate and petrous process are nearlyparallel to one another due to the increasedobliquity of the former and the more obtuse posi-tioning of the latter in relation to the mid-sagittalplane. In the Hexian specimen the petrous pro-cess is positioned less obliquely in relation to themid-sagittal plane than in material from ZKD (600

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vs. 400), in this respect falling within the upperrange of variation seen in modem human samples(38°-63°) (Tobias 1967). Both a styloid andvaginal process are lacking. The foramen spino-sum of the sphenoid is preserved on the rightside. The occipitomastoid crest is not as welldeveloped as in Peking man. As described byWeidenreich (1943:63) the foramen processusstyloidei (which occupies the position of themissing styloid process) and the digastric fossa(incisura mastoidea) of "Sinanthropus" lie on astraight line while the stylomastoid foramen lieswithout it. In the Hexian specimen these threestructures lie on the same line, concordant withthe modem human condition in which the styloidprocess, stylomastoid foramen and digastricfossa lie in tandem. The foramen magnum is atleast 32 mm wide. Its margin is strongly lipped,merging with the external occipital crest postero-laterally.

Endocranial Construction

The endocranial surface of the Hexian cal-varia preserves a low, broad frontal crest. It isdivided into two branches separated by a sagittalsulcus along the lower 9 mm of its course. Thisis similar to the condition in ZKD Skull III asreported by Weidenreich (1943:32). The sagittalsulcus fades out towards the lower third of thefrontal squama. There is noforamen caecum,consistent with the absence of this feature in"Sinanthropus" noted by Weidenreich (1943:34).There are clear cerebral impressions.

Along the sagittal margin of the parietals aresmall granular depressions, i.e., thefoveolae gra-nulares. The cerebral impressions and juga arenot as distinct as those seen on the frontal. Alongthe sagittal margin of each parietal is a low crestseparated one from the other by a sagittal sulcus.The sphenoidal angle of the left parietal is well-preserved. Internally, a Sylvian crest arises at theborder of the sphenoid and slants posterosuperi-orly following the anteroposterior diameter of theparietal for half its course. The anterior ramus ofthe middle meningeal artery cuts across the ante-rior portion of the Sylvian crest. The presence ofa developed Sylvian crest was observed by Wei-denreich in his description of the ZKD crania(1943:36).

The internal surface of the occipital showsthe typical division of the upper (cerebral) andlower (cerebellar) occipital fossae by a clear cru-ciate eminence. The central prominence of thislatter structure constitutes the internal occipitalprotuberance. As in other specimens of H. erec-tus the cerebellar fossae are distinctly smaller than

the cerebral fossae, the reverse of the condition inmodern man. In Peking man the ratio of the areaof the cerebral to cerebellar fossae is more or less2:1, while in the Hexian specimen it is closer to4:3, indicating an expansion of the relative area ofthe cerebellar fossae in relation to the cerebralfossae and a closer approximation to the modernhuman condition.

There is a 22 mm separation of the internaland external occipital protuberances from eachother in the Hexian specimen. This is clearly lar-ger than in the Dali cranium (11 mm) but smallerthan in Peking man (27.5-38.0 mm, mean = 32.8mm). In modern man these structures are coin-cident.

The sagittal sulcus on the internal surface ofthe occipital is broad and shallow. It unites witha similarly broad and shallow right transversesulcus. The left transverse sulcus is broad anddeep and positioned slightly higher than the righttransverse sulcus. It is not in communicationwith the sagittal sulcus. This configuration of theoccipital sulci conforms to the pattern observedfor the ZKD material (Weidenreich 1943:42).The transverse sulci do not extend upward to themastoid angle of the parietal as in modern manbut rather to the mastoid region of the temporaland occipital. The sulcal impressions of the cere-brum and cerebellum are poorly expressed.

The pyramidal process of the temporal isrelatively stout. The height of the posterior sur-face from the base of the superior border of thesigmoid sulcus to the superior petrosal sulcusmeasures 22 mm in the Hexian specimen. Thisvalue does not exceed 18 mm in "Sinanthropus"but in modern man it may approach 23 mm (Wei-denreich 1943:67). The apertures of both theinternal acoustic meatus and the internal vestibu-lar canal are intact. The sigmoid sulcus is broadand deep.

The ramification of the middle meningealartery follows the pattern seen in "Sinanthropus"in which the posterior branch (ramus temporalis)is somewhat more robust than the anterior branch(ramus fronto-parietalis) and the sub-branchesfrom both are much less abundant than in modernman.

Cranial Measurements

Cranial measurements of the Hexian calvariain comparison to relevant samples of H. erectusare given in Table 3. A number of these mea-surements are close to those for H. erectus fromZKD and Java. These include cranial length [g-op(i)], cranial height (po-b), inclination of thefrontal squama (b-g-i angle), horizontal curvature

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of the cranium (biauricular breadth/auriculo-bregmatic arc), fronto-parietal breadth index[minimum frontal breadth (ft-ft)/maximum cranialbreadth (eu-eu)], and parietal and occipital cur-vature. In a number of features the Hexianspecimen differs from material from Java and ismore similar to material from ZKD, e.g., mea-sures of biauricular breadth (au-au), cranialcircumference, transverse arc of the cranium,sagittal arc of the cranium, inferior inclination ofthe frontal, occipital angle and cranial capacity.The only index of the Hexian specimen that isdifferent from specimens of H. erectus fromZKD and more similar to specimens from Java isthe parieto-occipital breadth index. This indexdemonstrates the relatively larger breadth of theocciput in the Hexian cranium when compared tocrania from ZKD and the overall spherical shapeof the cranium versus the more elliptical shapecharacteristic of material from ZKD. Relativelyprogressive metric features of the Hexian craniuminclude an increase in minimum and maximumfrontal breadth, a greater degree of sagittal andfrontal curvature, and a relatively low value forthe parieto-occipital arc index (indicating that theparietal and occipital arcs are nearly equal to oneanother versus the greater length of the occipitalarc in most H. erectus and the greater length ofthe parietal arc in modem humans). In addition,the Hexian cranium shows a much greater maxi-mum breadth than either the ZKD or Javanesematerial and its length/breadth index falls withinthe brachycephalic range rather than the dolicho-cephalic range of most of the ZKD crania and themesocephalic range of Javanese specimens fromTrinil and Djetis. These latter traits, however, arenot thought to possess much phyletic weight(Huang et al. 1981).

The thickness of the cranial vault bones atvarious points along the cranium is as follows: atthe median point of the frontal squama 7 mm; atthe left parietal eminence 13.5 mm; at the mastoidangle of left parietal 18 mm; at the midpoint of theoccipital torus 18 mm; at the cerebellar fossa ofoccipital 6 mm; and at the median point of thetemporal squama 10 mm. These values demon-strate that cranial vault thickness is overall lessthan in Peking man and much less than in Lantianman.

Additional Remains

The supraorbital fragment (PA 840) des-cribed by Wu (1983) preserves much of the righttorus and a small portion of the frontal squamaimmediately superior to it. Medially it is brokenat glabella. The nasal process of the frontal is

preserved as is a small portion of the roof of theethmoidal sinus. The external surface of the boneis exfoliated in places. Wu (1983) observes thatwhile the supraorbital torus is strongly devel-oped, it is not quite as robust as in the calvarialspecimen PA 830. It differs from PA 830 andhomologous ZKD mateial in that the thickest partof the torus is situated along the supraorbital mid-line rather than further medially. Although mostof the frontal squama is lost, Wu (1983) notesthat, as in PA 830, postorbital constriction is notas great as in crania from ZKD. In like mannerthe post-toral sulcus is reduced in both PA 830and PA 840, differing in this respect from theZKD crania. In all other respects the PA 840specimen is said to be similar to both PA 830 andmaterial from ZKD.

The right parietal fragment (PA 841) des-cribed by Wu (1983) is 40 mm long and 60 mmwide. It preserves the area around the parietaleminence and a small portion posterior to theparieto-temporal suture. The bone surface issmooth without evidence of exfoliation. It isdarker in color than either PA 830 or PA 840,coming as it does from darker sediments towardsthe eastern part of the cave. The other cranialspecimens come from lighter colored sedimentstowards the western part of the cave. All thematerial, however, was excavated from the samestratigraphic level. The wall of PA 841 is rela-tively thick -- 11 mm in the vicinity of the parietaleminence, well within the range of Peking man(5-16 mm). The temporal margin is robust butthe bone thins out posteriorly.

The left mandibular corpus fragment (PA831) (Wu and Dong 1982) retains M2 and M3 insitu although the mesial margin of the M2 crownis damaged. The teeth are relatively well wornindicating that the specimen belonged to a matureindividual. The corpus is said to be extremelyrobust and particularly thick and is most likelythat of a male. There are three mental foraminapositioned between P4 and Ml. Corpus heightbetween Ml and M2 is 32 mm, corpus thicknessat the same position is 20.7 mm, yielding a ro-busticity index of 64.7 (Wu and Dong 1982).This is greater than in all ZKD and Javanesespecimens (both male and female), as well as theChenjiawo mandible from Lantian. The teeth re-tained by the specimen will be discussed belowalong with other dental specimens from the site.

Dental Remains

The dental remains from Hexian (Table 4)are remarkable for their overall robustness andlarge size. In nearly all instances (except M3) the

11

Table 4. Measurements of the Hexian dentition.

CrownLength Breadth

Right I' (PA 835) 11.7 9.4Right Pi (PA 832) 9.0 13.4Left M1 (PA 836) 12.3 13.7Right M2 (PA 837) 12.5 15.5Left M2 (PA 833) 12.0 14.0Left Ml (PA 834-1) 12.5 13.1Left M2 (PA 834-2) 13.3 13.6Left M2 (PA 838) 13.6 13.9Left M2 (PA 839) 14.3 13.4Left M3 (PA 831) 11.3 10.7

In millimeters (After Wu and Dong 1982; Nu 1963.)

teeth from Longtandong approach or exceed theupper limit of the range of metric variation forpreviously known dental specimens ofH. erectusfrom Asia. This is all the more significant giventhe relatively progressive nature of the Hexianremains and their purportedly young geologicage. Coupled with the large size of dental re-mains and the extreme thickness of cranial vaultbones from the upper Pleistocene site of Xujiayaoin Shanxi Province, it is apparent that mere mea-sures of overall size and robusticity do not serveas good phylogenetic indicators among east AsianPleistocene hominids.

Teeth recovered from Longtandong includethe socketed left lower M2 and M3 from the PA831 mandible, a right upper P4 (PA 832), a leftupper M2 (PA 833) and left lower M1-M2 [PA834 (1-2)] from the 1980 excavation, as well as aright upper IH (PA 835), a left upper Ml (PA836), a right upper M2 (PA 837) and two left lo-wer M2's (PA 838 and PA 839) recovered duringthe 1981 excavation.

The central incisor (Wu 1983) is particularlylarge, exceeding even the Yuanmou incisors inmesiodistal (md) length and labiolingual (11)breadth. Its overall size is most comparable tothe Krapina neandertals. The crown is well-pre-served except for a small piece of enamel lost atthe distal corner of the incisive margin which isotherwise only slightly worn. The specimen issaid to be clearly shovel-shaped with a broadincisive edge and thickened lateral margins whichtum inward towards the lingual surface of thetooth. The labial face of the crown projects bothlongitudinally and horizontally and has thre pro-minent longitudinal swellings. Lingually there isa pronounced basal tubercle which is confluentwith the lateral margins. There is a central lingual

fossa but it is not as well-expressed as in theYuanmou specimens. Four finger-like projec-tions of various lengths emanate from the basaltubercle and terminate in the central fossa. Theroot is extremely robust and conical. There aredistinct broad and shallow mesial and distalvertical grooves (Wu 1983).

The upper P4 (Wu and Dong 1982) is againvery large and robust, well exceeding the meansfor mesiodistal length and buccolingual breadthseen for this tooth in previous samples of H.erectus from Asia. The upper and lower molars,besides their large size, are morphologically con-sistent with other samples of H. erectus fromChina (Wu and Dong 1982; Wu 1983).

SUMMARY OF HUMAN REMAINS FROMHEXIAN COUNTY

It is clear from the above review that theHexian cranial and dental remains follow themorphological pattern of H. erectus in China asmeticulously outlined by Weidenreich (1937,1943). There can be no doubt as to their attribu-tion to the taxon H. erectus if that taxon is to haveany validity. The calvaria nevertheless doesshow certain progressive features that can best bedescribed as advanced over the condition seen atZKD, hence the appellation "advanced H. erec-tus" used in this article to describe the material.

CHAOXIAN COUNTY

The hominid bearing site in Chaoxian countyis located on Yanshan Hill outside of YanshanVillage, Daishan Township, Chaoxian county,

12

Anhui province (E 1170 52', N 310 33') (Xu et al.1984, 1986a, 1986b; Chen et al. 1987).

In April 1982, following reports of the dis-covery of Quaternary mammalian remains atYanshan Hill, a joint work-team of the IVPP andthe Anhui Archeological Institute was formed toexcavate the site. After one month's work a largequantity of mammalian fossils were recovered,including an incomplete human occipital bone.Further excavations during the winter of 1983 ledto the recovery of additional faunal remains aswell as a fragmentary human maxilla.

Yanshan Hill is part of a belt of low lyinghills, averaging 500 meters above sea level,which continue to its west and south. To the eastand north is the broad flood plain of the YuxiRiver which flows southeast towards theChangjiang (Yangtze) and drains Chaohu Lake.Terraces in the vicinity of Yanshan are poorlydefined but are clearly in evidence on either bankof the Yuxi. Yanshan Hill is an isolated karsticoutcrop that is riddled with a series of N/S trend-ing fissures, some of which have been widenedinto caverns. The fossil bearing locality is theremnant of one such cavern which has collapsedand been subsequently exposed on the westernslope of the hill. Five strata have been identified.The upper strata (1 and 2) are not in direct com-munication with the lower strata (3-5) (Figure 5).Their contained faunas also differ from the lowerstrata, yielding an assemblage with a definiteearly Pleistocene aspect and bearing a less diag-nostic but demonstrably younger fauna than thelower strata (Table 2). The human occipitalcomes from stratum 2 which has been correlatedfaunally with the upper layers at ZKD, although

Figure 5. Stratigraphic profile of Yanshan Hill,Chaoxian county, Anhui.

1

0 1itm

j,dcAi 4;3

2

1. Grayish deposits2. Reddish breociated deposits3. Yellowish deposits4. Reddish deposits5. Lght yellow deposits

i Umestone

3 Location of fossil hominid

there is no reason to reject a potentially youngerage (see discussion in Introduction above).

HUMAN REMAINS FROM CHAOXIANCOUNTY

Occiput

The occipital fragment recovered at Yanshandescribed by Xu et al. (1984) preserves much ofthe squama except for a small portion nearlambda which is damaged. The nuchal surfacebelow the inferior nuchal line is lost. The dam-aged area near lambda reveals that the inner plate,outer plate and diploe are all of equal thickness.The denticulated lambdoidal margin is well-preserved as is approximately 8 mm of the rightoccipitomastoid margin, allowing for the deter-mination of asterion on the right side. Theoccipital torus does not extend directly towardsasterion but is arched superiorly on either sidewith the lateral portions curving downwards.The torus does not project strongly and is restric-ted to the median portion of the occiput where itis of uniform thickness (ca. 15 mm). It fades outlaterally towards the asterionic region. The exter-nal occipital protuberance is indistinct. Althougha supratoral sulcus is undeveloped, a supratoralfossa can be seen above the median portion of thetorus. The superior nuchal line is indistinct as thesurface of the occipital torus merges smoothlywith the surface of the nuchal plane. The inferiornuchal line is somewhat more distinct. What ispreserved of the nuchal plane is relativelysmooth.

Internally, the cruciate eminence, sagittal andtransverse sulci, and internal occipital protu-berance can clearly be made out. The distanceseparating the external and internal occipital pro-tuberances is 22 mm, the same as in the Hexiancalvaria. Due to the incompleteness of the speci-men the relative proportions of the cerebral andcerebellar fossae cannot be made out.

In terms of overall dimensions the Chaoxianoccipital falls within the range of variation ofChinese H. erectus. Biasterionic breadth is re-constructed to be approximately 122.4 mm. Thisvalue is close to that of ZKD Skull V (124 mm),somewhat more than in other specimens fromZKD (111-117 mm), and considerably less thanin the Hexian specimen (141.8 mm). The lengthof the occipital chord from lambda to inion is50.2 mm, in keeping with specimens from ZKDwhich range between 47 mm and 52.5 mm.

Although the incompleteness of the specimendoes not allow for the direct calculation of theoccipital angle, its profile in mid-sagittal section

13

suggests that it was less sharply flexed than intypical specimens of H. erectus, approximatingmore closely the condition seen in archaic H. sa-piens. Thickness at the midpoint of the occipitaltorus is 7 mm. This is considerably less than inspecimens of H. erectus from China which rangefrom 12 mm to 20.4 mm.

In sum, the Chaoxian occipital specimen ismetrically similar to specimens of H. erectus pre-viously known from China in that the occiput isrelatively broad and the squamous portion is rela-tively short. However, a number of featuresdiffer from H. erectus and tend towards the con-dition seen in archaic forms of H. sapiens. Theseinclude: 1) the morphology of the occipital torus;2) the perceived greater degree of occipital cur-vature; and 3) the relative thinness of the bone.

Based on the preservation of the denticulatestructure of the lambdoidal margin, Xu et al.(1984) conclude that the lambdoidal suture hadnot closed. However, the relatively large breadthof the occiput suggests that the individual wasfully grown. Therefore, they infer that the spe-cimen belonged to a young adult. Given therelative gracility of the specimen and the undevel-oped nature of the occipital torus and otherectocranial buttresses, they further conclude thatit belonged to a female.

In assessing the phylogenetic affinities of theChaoxian occiput Xu et al. (1984) caution that itsrelatively progressive features may be a functionof both its age and sex. They conclude, how-ever, that the degree of occipital curvature,thinness of the cortical bone and abbreviation ofthe occipital torus exceed the range of variabilityobserved for male and female specimens, bothadolescent and fully mature, previously attributedto H. erectus in China. They therefore considerthe specimen to represent an anatomically primi-tive form of H. sapiens.

Maxilla

The maxillary fragment discovered in 1983,also from stratum 2, consists of the right maxillacontaining P3-Mi and the alveoli for Il-C, aswell as a portion of the left maxilla which retainsthe alveoli for I1-2 and the mesial wall of thecanine alveolus. The external surface of the rightalveolar process is lost posterior to P4, exposingthe whole root of Ml. Both lateral incisors arebroken off at the cervix. An isolated left P4, Mland M2 are also associated with the maxilla,yielding a total of 8 whole or partial teeth.

The maxillary specimen is well-preservedsubnasally and a moderate degree of alveolarprognathism can be observed in lateral aspect.

Xu et al. (1986a, 1986b) observe that the profilecontour of the nasoalveolar clivus is not straightnor slightly concave as in modern humans exhi-biting subnasal prognathism, but more convex asin Maxilla 5 of "Sinanthropus" (Weidenreich1943:76) or the Lantian maxilla. These latterspecimens, however, are said to be considerablymore prognathic than the Chaoxian specimen.Although the antenor nasal spine is broken away,Xu et al. (1986a, 1986b) infer it to have beenstrongly developed as reflected by the caliber ofits preserved base. Alveolar height (nasospinale-prosthion) is 27.8 mm, longer than in eitherPeking man (H. erectus) or Changyang man(archaic H. sapiens). The inferior margin of theright half of the piriform aperture is intact, as istwo-thirds of the left inferior margin. There is arather steep but short incline separating the naso-alveolar clivus from the inner floor of the nasalaperture, differing from the condition seen in "Si-nanthropus" in which the clivus is separated fromthe nasal floor only by a simple "margo limitans"(Weidenreich 1943:76). In this respect the Chao-xian specimen approximates the condition seen inmodern man. From the state of preservation ofthe right inferior nasal margin it can be estimatedthat the nasal aperture was relatively broad, muchas in "Sinanthropus".

The anterior portion of the base of the nasalcavity is preserved. The elliptically shaped nasalorifice of the incisive canal is positioned near thefront of the nasal cavity. The floor of the cavityis slightly concave posteriad versus a flat config-uration in the ZKD and Changyang remains.

The condition of the maxillary sinus can beobserved on the right side where it is exposed. Itextends anteriorly to a position in line with P3and medially to the palatine process. The maxil-lary sinus in "Sinanthropus" is not as extensive.

The anterior part of the hard palate as well asa small portion near Ml is preserved. As in spe-cimens of "Sinanthropus" and modern man, it isrugose. The incisive foramen is complete andpositioned near the alveolar margin (i.e., behindorale) and the incisive canal can be seen to as-cend nearly vertically. Both these traits are fullymodern and differ from the condition in H. erec-tus in which the incisive foramen is positionedmore posteriorly and the canal is inclined at anangle of approximately 400 to the alveolar plane.

Dental Remains

A total of eight whole or partial teeth havebeen recovered at Yanshan. They are all pre-served in situ in the maxilla or clearly associatedwith it. The alveoli of the two central incisors

14

and the right canine are preserved intact, whilethe lateral incisors are broken off at their roots.The superior extremity of the canine jugum isfairly well removed from the nasal floor, unlikethe condition in "Sinanthropus" in which thecanine jugum reaches the level of the inferiormargin of the nasal aperture. Xu et al. (1986a,1986b) note that this discrepancy may be due toeither an ovell reduction in the length of the ca-nine root or to the increased height of the alveolarface in the Chaoxian specimen.

In addition to the three teeth preserved intactin the right maxilla (P3-Mi), three isolated leftupper cheek teeth were also recovered. Theseconsist of a P4, Ml and M2. Only P4 preservesa complete root. The crowns of all six teethshow varying degrees of wear but are none-theless in good condition. Metrically all six teethfall within the mid to upper range of variation ofChinese H. erectus (Xu et al. 1986a, 1986b).They are more robust than the two teeth (P3 andMl) associated with the Changyang maxilla butless robust than material from Xujiayao (Table5).

Morphologically, the teeth are virtually indis-tinguishable from those of H. erectus in China(Zhang 1986). Although both upper central inci-sors are lost, their alveoli are completely intactand a labiolingual diameter of no less than 8.3mm has been estimated (Zhang 1986). This isconsiderably greater than the mean for the labio-lingual diameter of upper Il's from ZKD. Theupper lateral incisors are broken off at the cervix.Small portions of the crowns of both teeth are,however, preserved mesially and a labiolingualdiameter of between 8 mm and 8.5 mm has beenestimated (Zhang 1986). This is near the meanfor Chinese H. erectus.

Although the Chaoxian upper right P3 isworn occlusally, its structure is still evident. Asin H. erectus from ZKD the buccal and lingualcusps are separated by a deep longitudinal fur-row, the inner slopes of the two cusps have

developed horizontal ridges, and there is a well-defined "tuberculum molare" on the buccalsurface (see Weidenreich 1937:37). The arrange-ment of ridges and wrinkles on the upper P4's isalso said to be highly reminiscent of Chinese H.erectus (Zhang 1986).

The two upper Ml's, being antimeres, aremorphologically identical. The crown is rhom-boidal in occlusal outline and broader anteriorlythan posteriorly. The cusps are differentiallyworn but still identifiable. The protocone andparacone are nearly equal in size, the metaconeslightly smaller. The boundary between the pro-tocone and paracone is along the cental axis ofthe occlusal surface of the tooth. The basalswelling of the buccal surface and a Carabelli'strait on the mesiolingual surface of the protoconeobserved in the Chaoxian specimens are commoncharacteristics of H. erectus upper Ml's fromChina (Zhang 1986). The upperM2 is also notedto be morphologically indistinguishable from H.erectus. The "erectus"-like character of the teethof archaic H. sapiens in China is a unique featureof the continuum ofhuman evolution in east Asia(Zhang 1986) and has been noted for specimensfrom Xujiayao (Jia et al. 1979) and Tongzi (Wu1984).

SUMMARY OF HUMAN REMAINS FROMCHAOXIAN COUNTY

The relatively modern morphology of theChaoxcian mailla is consistent with the attributionof the Chaoxian occiput to anatomically archaicH. sapiens. Features reminiscent of "Sinanthro-pus", such as the relatively pronounced alveolarprognathism, broad nasal aperture and robust"erectus"'-like dentition, are balanced by suchprogressive features as a strong anterior nasalspine, the anterior position of the incisive fora-men and vertical orientation of the incisive canal,the greater extent of the maxillary sinus develop-

Table 5. Robustness of teeth crowns (length x breadth) recovered from Yanshan, Chaoxian compared toother Chinese specimens.

P' P4 m2

Chaoxian specimens 107.0 104.6-106.7 154.1-159.6 166.4ZKD specimens (ave.) 102.0 105.9 141.1 138.4Hexian specimens - 120.6 168.5 168.0-193.8Changyang specimens 78.4 - 138.2 -Xujiayao specimens - - 187.6 157.3

(After Xu et al. 1986a, b; Wu and Wu 1985.)

15

ment and the diminution of the canine jugum.The differences between the Chaoxian andChangyang specimens, in terms of greater alve-olar height and more robust dentition in theformer and a flatter nasal floor in the latter, maybe due to sexual dimorphism, with the Chaoxianspecimen representing a male individual and theChangyang specimen representing a female (Xuetal. 1986a, 1986b).

CONCLUSION

The Hexian and Chaoxian remains arepowerful evidence for the regional continuitytheory of human evolution in China. There canbe no doubt that a significant transformation

affecting the human lineage in east Asia tookplace during the terminal period of the middlePleistocene. The nature of this transformationmust still be elucidated.

It appears that there is a considerable degreeof continuity in the transition from "advanced H.erectus" to "archaic H. sapiens" in China. Indi-viduals with more or less of the former or lattermorphologies seem to have lived in close spatialand temporal proximity. Recent discoveries ofmaterial of an archaic "sapiens"-like morphologyin China and the dating of these and previouslyknown specimens tend to reinforce this interpre-tation (Table 6, Figure 6). Archaic material fromDali, Yingkou (Jinniushan) and Changyang hasall been dated within a period of time duringwhich advanced forms of H. erectus still appar-

Table 6. Late middle Pleistocene - early late Pleistocene hominid sites in China serialized by uraniumseries age determinations (after Chen and Yuan 1988).

Site name, province FossilsAge range

(KYA)

Yingkou, Liaoning

*Zhoukoudian Locality 1 (Layers 1-3)

Dali, Shaanxi

Changyang, Hubei

Cranium and

Cranium andpartial skeleton

Calvaria (Skull 5)

Cranium

Maxillary fragmentand teeth

Dingcun, Shanxi

Chaoxian, Anhui

Teeth and juvenileparietal

Occipital, maxillaryfragment and teeth

*Hexian, Anhui Calvaria, cranialfragments, mandiblefragment and teeth

Zhoukoudian New Cave Tooth

Miaohoushan, Liaoning

Maba, Guangdong

Tongzi, Guizhou

Xujiayao, Shanxi

Teeth, juvenilefemoral diaphysis

Calvaria

Teeth

Cranial bones, cranialfragments, mandiblefragment and teeth

Jiande, Zhejiang Teeth

* advanced H. erectus, all others attributed to archaic H. sapiens.

230-300

220-2 90

180-230

170-220

160-210

160-200

150-190

135-175

130-290

119-140

102-191

100-125

90-117

16

Figure 6. Approximate ages of late middle Pleistocene - early late Pleistocene hominid bearing sites inChina based on uranium series age determinations.

South China East China North China

Xujiayao

INaw Cave

I DingcunChangyang Niaohoushan II I Dali

250

ZKD

Yinqkou

300

ently persisted. A picture is, therefore, emergingof a swift yet mosaic replacement of "erectus"-like morphologies by "archaic"-like morphologiescommencing approximately 250,000 years ago ineast Asia (cf. Zhang 1988). Whether "advanced-erectus" and "archaic-sapiens" individualsrepresent successive populations or merely poly-morphs within a single evolving lineage is aquestion which will be resolved only through therecovery and analysis of more fossil material.

In another context it is clear that there wasconsiderable evolutionary change within the H.erectus lineage in Asia even prior to the appear-ance of archaic H. sapiens. These changesinclude an increase in average cranial capacity,associated expansion and change in the propor-tions of the various cranial vault elements and agracilization of the ectocranial buttressing system.With the advent of archaic H. sapiens in China anew set of derived "sapiens"-like craniofacialmorphologies are laid over the earlier "erectus"pattern (Tables 7 and 8).

What were the factors mediating this change?One explanation may be the emplacement of exo-genous characters into the indigenous populationby gene flow from without (an admixture model)(cf. Wolpoff 1985, 1989; Wolpoff et al. 1984).Comparison of archaic H. sapiens specimensfrom China and elsewhere in the Old World (cf.Wu and Wu 1982; Wu, X. 1988) may helpelucidate the extent of this postulated gene flowbetween Asia, Europe and Africa during the latemiddle Pleistocene. An increase in populationmovements during this period may well explainthe impetus for the transformation, along more

modem lines, of regional populations in east Asiaat approximately 250,000 YBP. This transfor-mation is a major yet unheralded event in theannals of human evolution, predating the transi-tion to fully modem H. sapiens in east Asia byperhaps 200,000 years. How the advent of"archaic Homo sapiens" in east Asia relates to thelater occurrence of anatomically modem humansin this region of the world is another question thatmust now be addressed as we begin to unravelthe evolutionary biology of east Asian hominids.It is more and more clear, however, that theargument for regional continuity throughout aconsiderable portion of the human career in eastAsia has been reinforced by the discovery of thenew hominid remains discussed above.

In conclusion, it is incontrovertible that theincreasingly complete record of human evolutionin east Asia during the late middle Pleistocene -early late Pleistocene transition will play asteadily more important role in attempts to under-stand the dynamics of human evolution in aregional context.

ACKNOWLEDGMENTS

I would like to express my respect and admi-ration for the community of paleoanthropologistsand Paleolithic archeologists in China today who,under at times difficult circumstances, haveperservered in their important work of documen-ting the physical and cultural legacy of humandevelopment in China. They deserve great creditfor their important contributions to the study of

kya

100

150

200

La iI To zi

HaianChaoxian

17

Table 7. Distribution of cranial features of the Hexian calvaria between "erectus"-like and "sapiens"-likemorphologies.

Features "erectus"-like "sapiens"-like

1. Supraorbital construction x2. Inion coincident with opisthocranion x3. Well-expressed angular torus x4. Temporal squamous margin arched x5. Sulcus of zygomatic process shallow and wide x6. Broad digastric fossa (incisura mastoidea) x7. Mammillary process of mastoid small and

positioned below supramastoid crest x8. Construction of external auditory meatus x9. Construction of mandibular fossa and

accessory structures x10. Frontal profile and inclination x11. Well-expressed frontal tubercle x12. Supratoral sulcus reduced x13. Increased frontal breadth x14. Reduced post-orbital constriction x15. Reduced sagittal keeling (also lack of

cruciate eminence at bregma and para-sagittal depressions) x

16. Greatest cranial breadth low x17. Acute angle between nuchal and occipital

planes of occiput x18. Construction of occipital torus x19. Orientation of tympanic plate to

mid-sagittal plane x20. Orientation of tympanic plate to

petrous portion x21. Configuration of styloid region x22. Absence of foramen caecum x23. Ramification of middle meningeal artery x24. Expansion of cerebellar fossae in relation

to cerebral fossae x25. Distance between external occipital protuberance

and internal occipital protuberance reduced x26. Stout pyramidal process x27. Horizontal curvature x28. Parietal curvature x29. Occipital curvature x30. Sagittal curvature x31. Frontal curvature x32. Cranial capacity x

Table 8. Distribution of cranial features of Chaoxian specimens between "erectus"-like and "sapiens"-like morphologies.

Features werectusu-like usapiens"-like

1. Construction of occipital torus x2. Presence of suprAtoral fossa on occiput x3. Distance between internal occipital

protuberance and external occipitalprotuberance reduced x

4. Overall dimensions of occipital x5. Occipital thickness x6. Nasoalveolar clivus convex x7. Nasoalveolar clivus separated from floor

of nasal aperture by short incline ratherthan "margo limitans" x

8. Anterior nasal spine present x9. Incisive canal ascends vertically x

18

human origins and evolution. I would also liketo thank Prof. T.D. White, Mr. Monte McCros-sin, Mr. Gary Richards and the editor for usefulcomments which helped im-prove the character ofthis manuscript.

NOTES

1 The term "advanced Homo erectus" is used todesignate hominid specimens which evince to aconsiderable extent the total morphological pat-tern of H. erectus as defined by Weidenreich(1943) for the ZKD remains while at the sametime possessing a limited number of characterstrending in the direction of later H. sapiens. Theterm "archaic Homo sapiens" refers to specimensclearly derived in the direction of modern humansbut whose total morphological pattern cannot beaccommodated by inclusion within "anatomicallymodern Homo sapiens".

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