Kroeber Anthropological Society Papers, Nos. 71-72, 1990 A Case Study of the "erectus" - "sapiens" Transition in Asia: Hominid Remains from Hexian and Chaoxian Counties, Anhui Province, China Dennis A. Etler Newly discovered fossil hominids from the late middle Pleistocene of China demonstrate that archaic forms of Homo sapiens were either coexistent with or slightly more recent than advanced forms of Homo 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 by archaic H. sapiens in China is characterized by the retention of a significant component of heritage features in archaic H. sapiens, overlaid by a mosaic of changes in craniofacial anatomy that trend towards modern humans. This new evidence gives powerful support to the regional continuity theory of human 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 between western and eastern Asia at approximately 250,000 YBP. INTRODUCTION The accelerated pace of discovery of late middle Pleistocene - early late Pleistocene human fossil remains in China over the last two decades makes it possible to approach the question of the "erectus" - "sapiens" transition in east Asia from an entirely new empirical basis. Human skeletal remains from this period of time include advanced forms of Homo 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 Homo sapiens from Changyang (Jia 1957), Maba (Wu and Peng 1959), Xujiayao (Jia and Wei 1976; Jia etal. 1979; Wu 1980), Dali (Wu 1981), Yingkou (Lu 1985, 1989; Wu, R. 1988) and Chaoxian (Xu et al. 1984, 1986a, 1986b) (Figure 1).1 Of particular note are the hominid remains from Longtandong, Hexian county and Yanshan, Chaoxian county, Anhui province which are taken to represent H. erectus and archaic H. sapiens, respectively. Not only are these two sites in close spatial proximity (Figure 2), they may also be close in age. The Hexian fauna (Tables 1 and 2) has been correlated with the upper layers at Zhoukoudian (ZKD) and oxygen isotope Stage 8 which would give an age of 240- 280,000 years before present (YBP) (Xu and You 1984). The Chaoxian fauna (Table 2) is thought to be somewhat younger due to the ab- sence of archaic species such as Megantereon and Trogontherium 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 hominid bearing strata at both sites, however, raises the possibility that they are virtually synchronous and, moreover, somewhat younger than the up- per layers at ZKD Locality 1 (Chen et al. 1987; Chen and Yuan 1988). The uranium series analyses, based on the internal concordance of 230Th/234U and 231pa/235U activity ratios of bones and teeth associated with the human remains, date the Hexian site to between 150- 190,000 YBP and the Chaoxian site to between 160-200,000 YBP. While uranium series dates are still subject to considerable controversy, the techniques employed to date the organic remains are identical at both sites, and the physical condi- tions encountered at Longtandong and Yanshan are similar. It is, therefore, reasonable to accept the dates as being broadly comparable. Nonethe- less, given the vagaries of uranium series dating these ages should best be taken as minimum ages for both sites. The close spatial and possible temporal proximity of the Hexian and Chaoxian hominids coupled with their morphological dif- ferences (vide infra) raise obvious questions concerning their relatedness to one another. In addition, the suggested late age of occurrence of H. erectus at Hexian raises questions about the temporal span of H. erectus in east Asia, and the mode and tempo of its replacement by archaic H. sapiens. In order to more fully appreciate the implications of these new finds for an under- standing of the course of human evolution in east Asia, this contribution will concentrate on descri- bing and interpreting the human fossil remains from both Hexian and Chaoxian. While I had the opportunity to view the Hexian calvaria during a recent visit to China, I have not seen the material from Chaoxian, nor have I had the chance to personally study the specimens in question. I have, however, availed myself of the Chinese descriptions of the spe- cimens which are quite detailed in extent and which follow the format and termiinology of Wei- 2 denreich's (1937, 1943) classic descriptions of the hominid remains from Zhoukoudian. Much of the analyses presented in the Chinese accounts has not been previously available in English. I hope that this presentation alleviates, to an extent, this circumstance, thereby facilitating discussion of these important hominid remains by a western audience. HEXIAN COUNTY The Longtandong cave site located on Wang- jiashan, a hill near Taodian village, Hexian county, 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 You 1984; Chen et al. 1987) has produced the most prolific remains of H. erectus unearthed in China since the height of excavations at ZKD in the 1930's. An inventory of recovered specimens includes: a nearly complete calvaria (PA 830), the supraorbital portion of a frontal (PA 840), a right parietal (PA 841), a fragment of a left mandibular corpus retaining M2 and M3 (PA 831), a right upper P4 (PA 832), a left upper M2 (PA 833), a left 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 two left lower M2's (PA 838 and PA 839). The site has been known since 1973 when it was blasted open during canal construction. The cave was found to be richly fossiliferous and a number of vertebrate specimens were sent to the Institute of Vertebrate Paleontology and Paleo- anthropology (IVPP), Academia Sinica for study. In 1979, at the request of the Anhui Province Water Conservancy Bureau, a team led by Huang Wanpo was dispatched by the IVPP to further investigate the site. More fossil vertebrates were collected along with bone that has been claimed to be artifactual in nature (Huang et al. 1982). Excavations were resumed in January 1980. In July 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, Shanxi 6. Dali, Shaanxi 7. Yingkou (Jinniushan), Liaoning 8. Miaohoushan, Liaoning 9. Changyang, Hubei 10. 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 back dirt within the cave. Later that fall during the third excavation the calvaria and other human fos- sils were found. Further excavations led to the recovery of additional teeth and cranial remains (Wu 1983). Longtandong is located on the northern slope of Wangjia Hill, 30 km northwest of the Yangtze River at an elevation of 23 meters above sea level. The deposits within the cave can be di- vided into four layers. The hominid material comes from the second stratigraphic level which consists of 0.5-1.0 meters of brown to yellow clays (Huang et al. 1981, 1982) (Figure 3). The Hexian fauna produced from the cave is bio- geographically transitional, containing both paleoarctic and subtropical elements (see Tables 1 and 2). The micro-mammalian component shows considerable faunal mixing with northern, southern and "alpine" species seen frequently in the western montane region of China all present (Zheng 1982; Xu and You 1984). Palynological sampling at Longtandong likewise indicates the presence of both northern and southern taxa (Huang and Huang 1985). Since the 1950's it has been anticipated that this area of the North China 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 as the Huai River Transitional Zone. The mammalian fauna from Sihong in Jiangsu Province, now known to be of Miocene age, was initially thought to be repre- sentative of just such a Pleistocene transitional zone. It was later shown, however, that castorid and proboscidean remains at Sihong had been misinter- preted and were not of Pleistocene age (Li and Zhou 1978). The faunal remains from both Hexian and Chaoxian are the first verification of the Huai River Transi- tional Zone and have contributed greatly to a better understanding of the paleobio- geographic zonation of east Asia during the middle and late Pleistocene (Liu et al. 1982; Zheng 1987). HUMAN REMAINS FROM HEXIAN COUNTY at The Hexian calvaria (Figure 4) has been studied in detail by Wu and Dong -j (1982). The following account of the specimen is based on their analysis. Preservation The heavily fossilized specimen was originally fragmented into eight pieces. Recon- struction revealed a well-preserved calvaria that retains a substantial part of the cranial base. The restoration consists of: 1) a nearly complete fron- tal including the supraorbital region, but lacking the inferior portion of the nasal process; 2) a nearly complete left parietal; 3) the right parietal somewhat damaged at pterion, near the parietal eminence and asterion; 4) a left and right temporal, both lacking zygomatic and mastoid processes due to breakage -- the left temporal is relatively complete, the right temporal squama has been lost at the sphenoidal margin and along a portion of the parietal margin; 5) an occiput that preserves the posterior margin of the foramen magnum and much of the nuchal and occipital planes; and 6) the lateral portion of the greater wing of the left sphenoid. The vault between the temporal lines anterior to the vertex and posterior to bregma is flattened. It has been suggested that this may be due to compressive forces during fossilization (Wu and Dong 1982). The relatively large and robust construction of the calvaria (cranial capacity approximately 1025 cc), the thick and well-developed supraorbi- tal and occipital tori, the well-expressed tempo- ral and nuchal lines and the sloping forehead 4 Table 1. Taxonomic list of non-mammalian and mici mammalian vertebrates from Longtandong, Hexi county, Anhui province (after Huang et al. 1981, l9f Zheng 1982; Xu and You 1984; Huang and Hua 1985). REPTILIA Ocadia sp. Amyda sp. Alligator cf. sinensis AVES Crossoptilon sp. MAMMALIA Insectivora Blarinella quadraticauda (Milne-Edward Chodsigoa youngi Huang et al. Anourosorex squamipes Milne-Edwards ?Scaptochirus sp. Chiroptera Rhinolophus cf. ferrum-equinum Thomas ?Myotis sp. Hipposideros sp. Miniopterus schreberii Kuhl Rodentia Trogontherium cuvieri Fischer Cricetulus varians (Zdansky) Apodemus argrarius Pallas Rattus rattus (Linnaeus) Rattus norvegicus Berkenhout Rattus edwardsi Thomas Microtus brantioides Young Eothenomys melanogaster Milne-Edwards Eothenomys inex Eothenomys eva Eothenomys proditor Tamias wimani Young combine to suggest a male specimen. Wu and Dong (1982) judge it to be a young adult, as both external and internal sutures (excepting the left sphenoidal suture) are open. Lateral Aspect Wu and Dong (1982) describe the calvaia as low vaulted with the supraorbital and occipital tori being respectively the most anterior and posterior projecting points on the skull, so that greatest cranial length (g-op) is coincident with the 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 line 82; terminates in a distinct angular torus at the ng mastoid angle of the parietal. The above features are all characteristic of previously known crania of H. erectus from ZKD. The pterionic region, preserved on the left side, is of the sphenoparietal type and assumes an "H" shape. The sphenoparie- tal suture is 11 mm long (Wu and Dong 1982). The temporal squama is relatively high and arched superiorly at the parietal margin unlike the condition in Peking man (with the exception of Skull V) in which the squamosal margin is low and straight. The left squamous portion is 70 mm long and 42 mm high, yielding a s) length/height index of 60, compared to an average of 49.6 in Peking man (Weiden- reich 1943), 64.6 in the Dali cranium (Wu 1981) and an average of 65.2 in modern man (Wu and Dong 1982). The Hexian temporal, therefore, approximates that of modern humans in both shape and size. The parietal notch separating the temporal squama from the mastoid region is deeply incised. The root of the zygomatic process of the temporal slants posterosu- periorly, merging with the supramastoid crest which in turn unites with the angular torus. The sulcus processus zygomatci, which positions the zygomatic process laterally away from the squama, is wide and shallow rather than narrow and deep as in modern man. The supramastoid and mastoid crests are well-developed as is the supramastoid sulcus. The digastric fossa is broad, shallow and opens up posteriorly, differing from the deeper and more narrow incisura mastoidea seen in modern humans. What is preserved of the mammillary process of the mastoid is relatively small and, as in other specimens of H. erectus, lies below the level of the supramastoid crest. It extends further backwards than in modern man. The configuration of the external auditory meatus and the overhanging suprameatal tegmen is as in "Sinanthropus" (see Weidenreich 1943:53). The aperture of the meatus is verdcally elliptic. The postglenoid process is poorly expressed as a low protuberance. Vertical Aspect The frontal has a low and sloping squamous portion. The frontal profile (m-g-i) (Martin 1928: 5 639, No. 32a) measures 580 and the inclination of the 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 Dali specimen in which the above angles measure 720 and 500 respectively (Wu 1981), but within the range of variation seen at ZKD (Weidenreich 1943:109) (Table 3). The supraorbital torus is wide laterally and thick inferosuperiorly. Although the superior margin of the glabellar region is slightly de- pressed, the left and right supraorbitals unite in a single body. Thickness measures of the supraor- bital torus are as follows: left medial 19 mm, left median 16 mm, left lateral 12 mm; right medial 18 mm, right median 17 mm, right lateral 13 mm. Greatest thickness is medial and least thickness is lateral as in specimens of H. erectus from ZKD. The frontal squama above the supraorbitals has a pronounced "bump-like" protuberance as in mate- rial from ZKD (Weidenreich 1943:225). This differs 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 it is much more weakly expressed than in similar specimens from ZKD. Javanese specimens lack this structure entirely as the flat squamous portion of the frontal merges smoothly with the supra- orbital torus. The frontal eminences are only slightly expressed and thus are similar in degree to male specimens from ZKD. A metopic suture is evi- Table 2. Taxonomic list of macro-mammalian vertebrates from Longtandong, Hexian county and the upper (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 Primates Macaca robustus Young Carnivora Canis sp. Cuon alpinus Linnaeus Vulpes sp. Arctonyx collaris Cuvier Lutra sp. Hyaena brevirostris sinensis Owen Megantereon sp. Felis chinensis Gray Panthera pardus Linnaeus Ailuropoda sp. Ursus arctos Linnaeus Ursus thibetanus kokeni (Zdansky) Proboscidea Stegodon orientalis Owen Perissodactyla Equus sp. Tapirus sinensis Owen Megatapirus sp. Dicerorhinus sp. Artiodactyla Sus lydekkeri Zdansky Sus cf. xiaozhu Cervus (Pseudaxis) grayi Zdansky Megaloceros pachyosteus (Young) Hydropotes inermis Swinhoe Elaphurus davidianus Milne-Edwards Bison sp. Caprinae x x x x x x x x x x x x x x x x x x x x x x x x x sp. sp. x sp. sp. sp. sp. x sp. x sp. x sp. x x x 6 dent from a point level with the frontal eminences to bregma. An unfused metopic suture is also seen in Skull XI from ZKD (Wu and Dong 1982). Wu and Dong (1982) note that the breadth of the frontal is expanded over those from ZKD. Least frontal breadth (ft-ft) measures 93 mm, greatest frontal breadth (co-co) 118 mm. Both measurements are nearly 10% greater than the average for ZKD calvariae (Weidenreich 1943: 106). The degree of postorbital constriction can be gauged by an index that measures maximum postorbital constriction (minimum breadth of the frontal behind the orbits) against the distance between the outer edges of the brow ridges (101 mm and 111 mm respectively in the Hexian specimen) (Wu 1981). In the Hexian specimen this index stands at 91 versus a range of 80.7- 82.9 in specimens from ZKD and 85.1 for the Dali cranium (Wu 1981). This relatively minor degree of postorbital constriction is considered a distinguishing feature of the Hexian calvaria by Wu 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 it gradually disappears. The sagittal keel in Peking man is more prominent extending from a position level with the frontal eminences to the obelion region of the parietals. The robust construction of the vault in Peking man is further reflected by the presence of a cruciate eminence at the junction of the coronal and sagittal sutures. The Hexian specimen lacks this latter structure as well as the parasagittal depressions seen along the sagittal suture in specimens of H. erectus from ZKD and Java (Wu and Dong 1982). Greatest cranial width is low on the skull in line with the supramastoid crests. The oudine of the neurocranium at this level is ovoid. The pan- etal eminences are well-developed which may be an 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 occipital torus is well-developed forming a continuous, smooth protuberance. The most projecting point Figure 3. Stratigraphic profile of Longtandong Cave, Hexian county, Anhui. Wangliashan L tf c I Longtandong well Longtan Y I .2 3 3 .4 -"-21 5 i 6i 1. Talus deposit 2. Fossil boaing deposits 3. Brownish-red day 4. Brownish-red day and sandy clay Intercalated with brecda 5. HominId clvaria 6. Cambrian dolomite A. Trial excavaton, Oct.1979 B. Excavation, Jan. 1980 C. Excavation, Oct.-Nov. 1980 liniinishing towards asterion. articular eminence is weakly expressed (Wei'd pratoral sulcus. The occipital reich 1943:47). C)f particular note is the prese ^ occipital