6 3 Hominids from the Lower Pleistocene of South China Geoffrey G. Pope For a number of reasons, the study of hominid evolution in China has always been a simultaneously fascinating and frustrating pursuit. Since the end of World War II and the disappearance of the largest single sample of Homo erectus fossils in the world, a large quantity of new information bearing on the pre- sence of lower and middle Pleistocene hominids has been published. Similarly, new information about the faunas which shared the habitat of Pleistocene man has also been forthcoming. Unfortunately, the absence of K/Ar dating and the lack of paleomagnetic profiles in China has necessitated a reliance on the "character" of faunal assemblages and the "evolutionary stages" of various taxa as indicators of the chronological relation- ships of various hominid fossils. With the exception of the Lantian calotte and mandible, hominid fossils at- ributable to the lower Pleistocene (between 1.80 and .69 million years ago) are very rare. The oldest known fossils on the east Asian mainland come from two localities in south China and consist of isolated teeth. While these specimens tell us little about the mor- phological evolution of Homo, their very presence indi- cates that by the early Pleistocene an early grade of Homo or a late grade of Australopithecus had already attained a wide distribution in the Old World. Before 1965 and the discovery of the first fossils discussed in this paper (Yuanmou, Yunan), no hominids were definitely known from the lower Pleis- tocene of the southern faunal area (an area in China bordered on the north by the Qin Ling Mountains; see Figure 1). The lower Pleistocene (including the lower middle Pleistocene of Chinese palaeontologists) mammalian faunas are characterized by a number of presumably tropical and subtropical elements which shared a number of affinities with other faunas of southeast Asia. The Stegodon-Aliluropoda fauna, as it has been called, has been divided into earlier and later components according to (1) evolutionary grades of Fig. 1 Pleistocene Faunal Localities of Southern China the members of various phylogenies (primarily size) and (2) the first and last appearance of various taxa. These subdivisions which are based primarily on the work of Pei (1961, 1963 and 1965), Kahlke (1961) and Colbert and Hooijer (1953) may be summarized as follows: Earlier (possibly including the late Pliocene) (1) Pongo (2) Gigantopithecus (smaller teeth) (3) Ailuropoda (smaller form) (4) Tapirus (smaller form) (5) Stegodon praeorientalis (6) Equus yunnanensis (7) Gomphotheriidae (8) Cuon (larger form) Later (1) Macaca (first appearance) (2) Gigantopithecus (larger teeth?) (3) Pongo (4) Hylobates (? first appearance) (5) Ailuropoda (larger form) (6) Tapirus (= Megatapirus) (7) Elephas namadicus (8) Stegodon orientalis (9) Cuon (smaller form with well- developed metaconid) It must be noted that Chinese palaeontologists seem to use a 3 million year long Pleistocene (in only one publi- cation, Zhou and Zhang 1974, has this been explicitly stated). The beginning of the middle Pleistocene also remains undefined and the use of both epoch subdivi- sions varies with different authors. More recent au- thors seem to be using a "date" for the beginning of the Pleistocene similar to the one used in this paper. Addi- tionally, some of the above listed taxa may be present in both epoch subdivisions in spite of the fact that they are currently unknown from one or the other. The assumption of rectilinear phyletic sequences and con- tinuous change in size of these taxa has played a par- ticularly important role in assigning relative ages to the members of presumed lineages. However Tapirus and "Megatapirus" probably do not constitute a single lineage (Colbert and Hooijer 1953) and both are known from the Trinil faunal zone. The presence of a large form of Ailuropoda, the presence of Macaca and the presence of Elephas namadicus may prove to be the best guide fossils for a period of time equivalent to the late lower Pleistocene or early middle Pleistocene. However, since there are few well-studied stratig- raphic columns in south China and no radiometric dates, the use of first appearances is an extremely tenuous approach which cannot be verified. Yuanmou (Mai-Kai Valley) is located on the Long-Chuan River on the northern part of the Yunan Highlands (see Figures 1 and 2). Here an extensive series (at least 370 meters thick) of fluviolacustrine deposits has been exposed. The strata consist of a number of sands, gravels, clays and river terraces (see Figure 3). The lower 140 m of the section of the Yunamou deposits which are overlain by riverine gravels contain pollen indicative of tropical or subtrop- ical conditions (Hu 1973). The Yuanmou deposits may possibly cover a a time period which extends from the late Pliocene to the middle Pleistocene. Hominid incisors were recovered from deposits of brown, sandy clay situated approximately 59 m from the top of the section. Unfortunately, no explicit locality for the hominid remains is given in any of the literature. You and Qi (1973), however, state that the locality is in the vicinity of the village of Shang-Na-Bang and on the same stratigraphic level as locality 67003. While the uncertainty of the location of the finds points to a common frustration in assessing paleontological specimens in China, on the basis of the fauna and strata of localities 67001 and 67005 which reportedly are stratigraphically lower than locality 67003, a lower Pleistocene dating of the hominid fossils seems highly plausible (see Table 1). Presumably lower Pleistocene forms recovered from the two stratigraphically lower localities include Equus yunnanensis, Stegodon zhaotungensis and Hyaena licenti. The fauna from local- ity 67003 itself is not identified below Cervus spp., Gazella sp. and Bovinae indet. Of the above mentioned taxa, none can be definitely shown to be confined exclusively to the lower Pleistocene, but this may prove to be the case with E. yunnanensis and S. zhaotungensis. Few of the "characteristic" lower Pleistocene taxa of south China (i.e. Ailuropoda, Tapirus, Gigantopithecus, etc.) are present at these localities. This is due most probably to the fact that the portion of the Yuanmou deposits which contains the hominid incisors samples a cooler highland habitat. This interpretation is also supported by palynological evidence (Hu 1973). TABLE I Fossils and Fossil Localities of the Yuanmou Basin (From You and Qi 1973) Taxa Canis yuanmoensis Vulpes cf chiknshanensis Felis tigris Felis pardus Cynailurus sp. Hyaena licenti Hyaena sp. Stegodon zhotungensis Stegodon sp. Equus yunnanensis Rhinoceros sinensis Sus scrofa Cervus sp. A Cervus sp. B Bovinae indet. Gazella sp. Fossil Localities 67001 67001 67004 67001 67004 67001 67004, 67006 67005 67005 67001 67001 67004, 67006 67001, 67003 67001, 67002, 67003 67001, 67003, 67004, 67006 67003 6 4 6 5 1 Long-Chuan River 6 Ma-Da-Hai a 67005 A Jin-Sha River 2 Highway 7 Da Na-Niao b 67004 B Sha-Jie 3 Yuan-Mou 8 Shang-No-Bang c 67001 C Yuan-Mou 4 Yang-Liu Village 9 Xin-FO-Cun d 67002 D Wu-Ding 5 Xiao-Na-Niao e 67003 E Kun-Ming f 67006 F Long - Chuan River Fig. 2 Distribution Yuan Mou Quaternary Fossil Localities Approx. Scale 1:50,000 (After You & Qi, 1973) Horiz. Scale: 1:9000 Vert. Scale: 1 1500 * NW Yang-Liu Village Long-Chuan River T\ Sandy Soil Clay 1111 Verticlly M" Soil Sandy Clay - Mudstone J.' Sond -M Fossil Localty Mud arbonaceous Sand Pocket f3 Gravel Paleosol Bedded S Fault Fig. 3 YUAN MOU BASIN - Cross Section from Ma-Da-Hai to Long-Chuan River (After You & Qi, 1973) ,Loc. 67001 Brief Description of the Yuanmou Incisors The Yuanmou incisors are comparable in a number respects to the Zhoukoudian (= Choukoutien of the Wade-Giles System) Homo erectus incisors (described by Weidenreich 1937). The incisors are shovel-shaped and evidence a basal lingual tubercle (gingival emi- nence) and "finger-like prolongations" which are situated on the mesial and distal margins of the in- cisors. Both incisors are thought to represent upper central incisors belonging to the same individual. Al- though the Yuanmou incisors conform to the observa- ble morphology of "Sinanthropus", they are also differ- net in a few respects. While the incisor roots are round in cross section and robust in both the Yuanmou and "Sinanthropus" in- cisors, the Yuanmou roots are noticeably constricted at the neck. This trait adds to the overall triangular ap- pearance of the crown of the Yuanmou specimen (Hu 1973). In absolute measurements (see Table 2), the left Yuanmou specimen is longer than "Sinanthropus" male and female speciments (Weidenreich 1937), the Sterkfontein Sts. 52a specimen (Robinson 1956), the B-27 Omo specimen (Howell 1969) and the AL 200- la Hadar specimen Uohanson and Taieb 1976). In width, both the Yuanmou and "Sinanthropus" specimens are slightly smaller than the Sterkfontein and Hadar specimens. In length, the Yuanmou specimen is furth- est from the B-27 Omo specimen from the Brown Sands area which lies within the range of au- stralopithecines (Howell 1969). The Yuanmou speci- men is closest in length to the right incisor of the Hadar palate which is comparable with other material; assigned to Homo erectus; especially "Pithecanthropus IV" (Uohanson and Taieb 1976). In morphology as well as size the Yuanmou specimens are probably best assigned to an early form of Homo erectus. Farther to the north, four lower molars have been recovered from the Badong and Jian Shi districts of Hubei. Three of these teeth (PA-504, PA-502, PA-503) were found in cave deposits in association with isolated teeth which have been assigned to Gigantopithecus. The fourth molar, PA-507, from the Badong district, lacks provenience. All have been referred to Au- stralopithecus. Although they have been described as Table 2 Comparison of Measurements of Central Upper Incisors (mm) (Modified from Hu 1973, with Additions) 3 Length U Width Yuanmou male (left) 11.5 8.1 "Sinan- thropus" male (left) No. 4 "Sinan- thropus" Sterk- female fontein (right) (left) No. 2 Sts. 52a Omo (right) B-27 I a ~~i 94 9-4 10.7 9.8 9.5 9.2 Hadar (left) AL 200-la 10.8 0 0 m i 4 l -4 8.1 7.9 8.2 8.3 comparing favorably with A. africanus, Gao (1975) has sugested that they represent a new Asian species of Australopithecus. The cave deposits, which produced the PA-520, PA-503 and PA-504 molars, are characteristic of the Karst Cave deposits which are widespread in southern China (see Figure 4). The "dragon bone cave" itslef is located approximately 85 meters above Dragon Cave stream, a position characteristic of the older Karst Caves (see White 1974 for a review of the rationale regarding the temporal ordering of Karst Caves). The fossil content of the deposits is rather uniform regard- less of level. The fossils were either washed into the caves by running water as suggested by Hsu, et. al. 1974 or deposited as a result of the bone collecting habits of Hystrix. The hominid and other mammalian fossils consist largely of isolated teeth. The hominid and G.igantopithecus teeth were recovered from the pruple-red clay layer marked by the second lowest fossil symbol in Figure 4. The brownish-yellow sandy deposits and the stratigraphically lower cemented yel- low sands have produced exactly the same genera. The various strata probably do not represent discrete chronostratigraphic units. The fauna of Jian Shi (see Table 3) has been termed "late lower Pleistocene" (?ca. 2 m.y. BP) and has been considered to be intermediate in age between the ear- lier Xin-She-Chong (Liu Cheng Gigantopithecus cave) and the presumably middle Pleistocene Yan-Jing-Gou (see Figure 1) fissure fauna near Wan Xian, Sichuan (Hsu et al. 1975). Among the primates, the presence of Pongo and the absence of Macaca is taken as an indi- cator of the lower Pleistocene age of the fauna. In size and morphology, the Gigantopithecus teeth are nearly identical to the Xin-She-Chong specimens. Among the carnivores, Cuon is smaller in size than the genus rep- Table 3 Jian Shi Faunal List Gigantopithecus blacki Cuon javanwus antiquus Ursus sp. Ailuropoda cf. melanoleucafovealis Arctonyx sp. Hyaena licenti Felis sp. Machairodontinae Gomphotheriidae Trilophodon serridenstoides Stegodon sp. Equus yunnanensis Tapirus sinensis Rhinoceros sp. Sus sp. A Sus sp. B Ceruus sp. Muntiacus sp. Bonvinae Ovinae Hystrix sp. 6 6 I i 0 5 lo 15 M. N