Citation: Zhao L X, Wang C B, Jin C Z, et al. Fossil Orangutan-like hominoid teeth from Late Pleistocene human site of Mulanshan cave in Chongzuo of Guangxi and implications on taxonomy and evolution of orangutan. Chinese Sci Bull, 2009, 54: 3924―3930, doi: 10.1007/s11434-009-0653-x
Fossil Orangutan-like hominoid teeth from Late Pleis-tocene human site of Mulanshan cave in Chongzuo of Guangxi and implications on taxonomy and evolution of orangutan
ZHAO LingXia1†, WANG CuiBin1, JIN ChangZhu1, QIN DaGong2 & PAN WenShi2 1 Institute of Vertebrate Paleontology and Paleoanthropology, Chinese Academy of Sciences, Beijing 100044, China; 2 Chongzuo Biodiversity Research Institute, Peking University, Beijing 100871, China;
Fossil records indicate orangutan-like hominoids have been widely distributed in south China during Pleistocene, although currently only surviving in the tropical forests of Kalimantan and Sumatra in Indonesia. This paper describes the recently discovered hominoid fossil teeth from human site of Mu-lanshan cave in Chongzuo of Guangxi, whose geological age is the Late Pleistocene, about 11000 yeas age based on associated mammal fauna and U-series dating. Compared with those of modern and subfossil orangutans from Indonesia, other fossil great apes from China, the hominoid teeth from Mu-lanshan cave are orangutan-like, but show somehow different from Indonesia's orangutans, the aver-age sizes of cheek teeth larger and occlusal enamel wrinkles less and simpler. They are classified temporarily as the subspecies of Pongo pygmaeus weidenreichi. Concerning the variations of mor-phological features and dental sizes of orangutan-like teeth from southern China and neighboring northern Vietnam, different subspecies or species or genus possibly, but the key evidence is necessary to be identified.
Late Pleistocene, Mulanshan Cave in Chongzuo of Guangxi, Pongo pygmaeus weidenreichi, hominoid tooth
Orangutans are the only great apes in Asia, surviving in the tropical forests of Kalimantan and Sumatra of Indo- nesia. But fossil records show that orangutan-like ho- minoids have been widely distributed in Southeast Asia and southern China during Pleistocene[1−7]. The fossil orangutan-like hominoids in Asia play key roles in un- derstanding the taxonomy and phylogeny of extant and fossil hominoids, their evolution and extinction with climate and environmental changes since Neocene. Be- fore the 1980s, the Miocene orangutan-like hominoid fossils, from Siwalik area of India-Pakistan and Yunnan in southwest China, were considered the earliest mem- bers of hominidae. In recent decades, molecular anthro- pology and rich fossil evidence of early hominid from African strongly suggest that African modern great apes are much closer to, or even included into Hominidae,
Asian Miocene large-bodied hominoids are linked to
Pongidae than Hominidae by most of anthropological
scholars. However still fewer scholars take contrary opi-
nion, as Grehan and Schwartz[8] insist that there are
many features that suggest affinities between human and
orangutan and orangutan-like hominoids. Xu and Lu[9] place Lufengpithecus as an early member of hominidae
in recently published monograph. In addition, Ciochon
et al.[10,11] suggests a mystery ape existing in Pleistocene
Asia, which is different from orangutan. All views indi-
Received July 6, 2009; accepted September 8, 2009 doi: 10.1007/s11434-009-0653-x †Corresponding author (email: [email protected]) Supported by the National Basic Research Program of China (Grant Nos. 2006CB806400 and 2007FY110200) and Key Knowledge Innovation Project of Chinese Academy of Sciences (Grant No.KZCX2-YW-106)
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cate the complexity and uncertain problems of orangutan- like hominoid issues in Asia. In recent decades, there are more and more orangutan-like hominoid fossil teeth discovered in south China, especially which coexist with human artifacts in some sites such as Longgudong cave of Jianshi in Hubei, Mohuidong cave of Tiandong in Guangxi, Longgupo of Wushan in Chongqing. Orangu-tan-like fossils are the common members of Pleistocene fauna in south China, their distribution ranges from the north of Yanhui cave in Tongzi of Guizhou (28°15′N) to the south of Honglin quarry (19°15′N) of Changjiang in Hainan island, from the east of Shuangyan cave (113°24′E) in Yingde of Guangdong to the west of Heshang cave (102°30′E) in Fumin of Yunnan, up to now more than 30 sites discovered[1−6] (Figure 1). Fewer detailed study on the new-discovered Pleistocene orangutan-like teeth of south China has been done, which are usually listed as a member of Pongo sp. in
fauna lists, except that the study by Gu et al.[1] and Zhou[2] with a few fossils. Detailed study is necessary for such kinds of orangutan-like hominoids from south China, which is helpful to interpreting the early hominid origin and hominoid evolution in East Asia. This paper will describe the recently discovered hominoid teeth from Mulanshan cave in Chongzuo of Guangxi, where a human mandible fragment fossil is discovered, and also compare them with those of modern and early Holocene subfossil orangutans from Indonesia, and Pleistocene Gigantopithecus and late Miocene Lufengpithecus from
south China, and try to interpret implications for the taxonomy and evolution of orangutan.
Figure 1 Distribution of Mulanshan cave (★) and other Pleistocene orangutan-like fossil sites (●) in south China. 1, Yanhui cave of Tongzi; 2, Shuangyan cave of Yingde; 3, Honglin quarry of Changjiang; 4, Heshang cave of Fumin.
1 Fossil site and geological age
Mulanshan cave (22°17′13.6″N, 107°30′45.1″E) is lo-cated at the west-southern slope of Mulanshan hill, about 2 km northwest from Ecological Park of Chong-zuo, Guangxi. It is a karst cave developed in Lower Tri-assic limestone landscape. Altitude of the cave roof is 179 m above sea level, 34 m high over the local river. The fossil deposition contains two different strata units A and B, which formed in different periods. Human fos-sil mandible fragment and mammal fauna fossils are discovered in unit B. The hominoid teeth described in this paper also are excavated from unit B. Based on mammal fauna analysis and U-series dating, the geo-logical age of hominoid teeth is early Late Pleistocene, about 110000 years ago[12].
2 Description and measurement
Total 14 isolated hominoid teeth are analyzed. Only one deciduous tooth (CZ0408a-4) is upper molar, and all other teeth are permanent teeth, including 5 upper mo-lars, 1 lower incisor, 2 premolars and 5 lower molars (Table 1). Most of tooth crowns are well preserved with little part of roots (Figure 2). Worn condition in most tooth crowns is slight, so that morphological features are
well preserved. All those teeth represent 5―7 individu-
als at least. The morphological description is following, and the mesiodistal crown diameter (MD) and perpen-dicular buccolingual diameter (BL) are measured and crown area are calculated (MD× BL) (Table 1).
I1: The crown (CZ0408a-5) is high and narrow, ant its shape is nearly symmetrical with upper medial corner a little sharper than the lateral one, the cutting edge is the widest part of crown. The labial surface is slightly con-vex from above downward and transversely. The lingual surface is flat transversely in above half and becomes convex downward where the cingulum forms a triangu-lar concave in the lower one-third. The lateral and me-dial surfaces of crown are triangular with an acute angle at the cutting edge.
P4: There are 2 P4 (CZ0408e-10, 8). The crown is four-sided with blunt angles. Maximum diameter is from anterior-buccal to posterior-lingual. On the buccal sur-face the enamel border is produced toward the roots, more visible in CZ0408e-10 than CZ0408e-8. The two cusps, buccal protoconid and lingual metaconid, are placed on a transverse line distinctly nearer to the ante-
Figure 2 Hominoid teeth from Mulanshan cave of Chongzuo in Guangxi. From left to right, upper: CZ0408e-10, CZ0408e-8, CZ0408a-4, CZ0408e-6, CZ0408e-9, lower: CZ0411-5, CZ0408-1, CZ0408a-3, CZ0408-13, CZ0408e-11.
Table 1 Measurement of hominoid teeth from Mulanshan cave
Specimen Tooth type Worn condition MD (mm) LB (mm) Crown area (mm2) Crown area of modern Pongo[12] (mm2)
CZ0408a-4 Right dM2 slighter 12.5 12.5 156.3
CZ0408a-7 Right M1 heavy 11.8 13.7 161.7 160.8±23.91
CZ0408e-6 Right M2 unworn 14.7 15.8 232.3
CZ0408e-14 Right M2 moderate 12.9 14.4 185.8
CZ0408e-9 Left M2 slight 14.0 15.0 210
170.0±30.97
CZ0411-15 Right M3 unerupted 12.0 13.4 160.8 151.8±32.14
CZ0408a-5 Right I1 moderate 8.8 9.3 81.8 88.1±15.22
CZ0408e-8 Right P4 slight 10.9 13.1 142.8
CZ0408e-10 Left P4 slighter 10.9 12.3 134.1
131.9±22.87
CZ0408-1 Right M1 slight 13.4 11.2 150.1 155.1±22.95
CZ0408-13 Left M2 moderate 15.1 13.7 206.9
CZ0408a-3 Right M2 slight 16.3 14.8 241.2
173.9±32.05
CZ0408a-2 Right M3 unworn 16.6 15.0 249
CZ0408e-11 Right M3 unerupted 15.8 13.2 208.6
168.0±34.27
rior than to the posterior border of the crown. The transverse ridged in the two cusps are connected and separate the anterior small trigonid fovea and the poste-rior large talonid basin. The talonid basin occupies more than two-third of occlusal area. Enamel wrinkles are visible in the talonid basin.
There is some difference in morphological traits of 2 P4. Trigonid fovea is larger and longer transversely in CZ0408e-10 than in CZ0408e-8 in which is a small pit. Relative to the whole crown area, the talonid basin oc-cupies a larger part in CZ0408e-10 than in CZ0408e-8. The posterior part is more lingually produced in CZ0408e-10 than CZ0408e-8.
Upper molars: There are four main cusps: two buccal
cusps, the paracone and the metacone are relatively high and sharp, while two lingual cusps, the protocone and hypocone are low and blunt. The protocone and the pa-racone are connected by the anterior transverse ridge, and the protocone and metacone are connected by the central olique ridge. The trigonid fossa is deep in central, anterior fovea is small after the anterior marginal ridge and the posterior fovea is prominent in front of the pos-terior marginal ridge. The occlusal enamel wrinkles are visible but not so dense. The buccal vertical groove be-tween the paracone and the metacone runs from up downward and disappears at one-half height of the crown and form a small pit with a short horizontal ledge in CZ0408a-7 and CZ0408e-6. The lingual vertical
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groove between the protocone and hypocone runs almost throughout the whole crown height. The buccal surface is steep and the lingual surface is oblique and round- going. There is a short oblique ledge at the anterior- lingual middle surface of protocone in CZ0408e-6, which is the residual trace of cingulum. The morphology of M3 CZ0411-15 is very different from other molars, the metacone reduced and hypocone almost absent. CZ0408a-4 is an upper second deciduous molar, tooth crown and cusps are low, there are residual traces of cingulum as small ledges at the anterior-buccal surface of protocone and middle lingual surface between para-cone and metacone.
Lower molar: There are five main cusps: the buccal protoconid and hypoconid and hypoconulid, and the lingual metaconid and entoconid. 5 cusps are arranged in “Dryopithecus type” and clearly separated by occlusal grooves. The three buccal cusps are blunt and the two lingual cusps are relatively sharp. The tips of cusps are located laterally and so the central talonid basin is wide and large and occupies most part of the total occlusal area. The morphology of mesial fovea and distal fovea is various or absent. The occlusal enamel wrinkles are coarse and sparse and different from the pattern of fine and dense enamel wrinkles in modern orangutan. The two buccal grooves are clear and somehow deep, and the lingual groove is almost invisible and shallow. Of the three lower molars, M2 and M3 are bigger than M1.
CZ0408-1 is M1, and there is clear mesial fovea after the mesial marginal edge and distal fovea before the distal marginal edge. The buccal groove between proto-conid and hypoconid runs up downward and disappears at the one-half of crown height where a small pit is formed. This feature is different from the other lower molars, in which the buccal groove runs almost throughout the whole crown height. The enamel wrin-kles are sparse and simple. The crown shape of the two M2 (CZ0408-13, CZ0408a-3) are more square than that of M1 and M3, and the talonid basin is large and wide opened. There is no clear mesial fovea and distal fovea in CZ0408a-3 M2, CZ0408a-2 M3 and CZ0408e-11 M3. The crown of CZ0408-13 M2 is moderately worn, its mesial fovea is visible. The crown shape of two M3 (CZ0408a-2, CZ0408e-11) becomes narrow distally be-cause of the reduced hypoconid and entoconid, its talo-nid basin is narrower and longer than those of other lower molars. There is a small cusp between metaconid and entoconid in CZ0408a-2, between hypoconuid and
entoconid in CZ0408e-11.
3 Discussion and conclusion
Large-bodied hominoid fossils have been richly discov-ered in south China, mainly including three categories, Miocene Lufengpithecus from Kaiyuan, Lufeng, and Yuanmou of Yunnan, Pleistocene Gigantopithecus blacki from Guangxi, Guizhou and western Hubei-the Three Gorges area, and Pleistocene Pongo or orangutan-like hominoids which are widely distributed in Guangxi, Guangdong, Yunnan, Guizhou and Hainan
Hominoid teeth from Mulanshan cave are clearly dif-ferent from those of Gigantopithecus in the crown size, crown height and enamel thickness. The premolars and molars from Mulanshan cave are significantly smaller in tooth size, lower in crown height, thinner of enamel b than that of Gigantopithecus. Although there is no sig-nificant difference in tooth morphology between homi-noids from Mualnshan cave and Miocene Lufengpith-ecus from Yunnan, it is obviously different in the tooth size. 10 isolated lower buccal teeth of Lufengpithecus kaiyuanensis from Xiaolongtan of Kaiyuan are smaller than the smallest of corresponding hominoid teeth from Mulanshan of Chongzuo[13,14]. The similar situation oc-curs in Lufengpithecus hudienensis from Shihuiba of Lufeng[15]. There is a little overlap in range of tooth measurements between that of Lufengpithecus lufengen-sis[9,16] and hominoids from Mulanshan cave, the mean size of buccal teeth of Lufengpithecus lufengensis as a whole are smaller than that of Mulanshan hominoids. In addition, the lower central incisor from Mulanshan is obviously wider than that of Lufengpithcus.
The size and morphology of hominoid teeth from Mulanshan are comparable to that of modern and sub-fossil orangutans from Indonesia. That is why we call here orangutan-like hominoid. Extant orangutans live in Sumatra (Pongo pygmaeus abelii) and Borneo (Pongo pygmaeus pygmaeus) only, and orangutans or orangu-tan-like hominoids have been widely distributed in Southeast Asia and south China during Pleistocene. The enamel wrinkle pattern of buccal teeth is coarser and simpler in Mulanshan hominoids, and much denser and finer in Indonesia orangutans. But there are similar teeth with denser and finer wrinkles found in other sites in southern China, for example, a lower molar (V147 in Guizhou Museum) from Guanyin cave in Guizhou. The two different patterns of enamel wrinkles are also visible
in Lufengpithecus. Beside the different pattern of enamel wrinkle system, comparisons of tooth size also show
somewhat difference as in Tables 1―4. Table 1 shows
that crown sizes of M2 and M2 and M3 from Mulanshan are significantly bigger than that of mean values of modern and subfossil orangutan, while other tooth types are nearly around the mean values, few teeth less than the mean of the latter. However, all measurement data of Mulanshan hominoids are within the range of latter[5,17].
Compared to the large sample of each tooth type of modern and subfossil orangutans, the sample from Mu-lanshan cave is so much less, and it is better to analyze it
as a whole. Here we compare the frequency distribution of all tooth crown sizes of hominoids from Mulanshan cave with that of modern and subfossil orangutans after the analysis method by Harrison in study of prehistoric primate teeth from Indonesia[12]. Table 2 shows the whole sample of Mulanshan cave falls within the range of modern and subfossil orangutans, but most specimens (11/13) move upward above the mean of modern oran-gutan, and only 2 specimens are below the mean of modern orangutans within one standard deviation while nearly 30% in the subfossil orangutans. The data indi-cates that, hominoid teeth from Mulanshan, as a whole
Table 2 Comparison of distribution of the crown areas (MD × LB) of the permanent teeth of hominoid teeth from Mulanshan cave with those of modern and subfossil orangutans from Indonesia
Standard deviation intervals from the mean values f modern orangutans
CZ0408-013 15.1 13.7 206.9 Mulanshan cave Early Late Pleistocene
CZ0408a-003 16.3 14.8 241.2
Ganqianyan[2] Late Pleistocene 15.3 13.2 202
Mean of Chinese fossil Pongo (n = 10) 241.7
Mean of Modern Pongo (n = 61)[17] 173.9 9 (32.05)
Table 4 Comparison of the distribution of crown areas of lower M2 of fossil orangutans of south China with those of Padang Highlands subfossil and modern orangutans of Indonesia
Standard deviation intervals from the mean values of modern orangutans
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sample from Mulanshan, are generally bigger than those of modern and subfossil orangutans.
In order to analyze the variation of tooth size of orangutans from different sites with different geological ages, we compare data of crown sizes M2 in Tables 3 and 4. On one hand there are more comparable data on fossil orangutans from south China, more importantly on the other hand, the size of M2 is highly related to body size in primate and usually be used to estimate the body size and weight. All the specimens of fossil orangutans from south China are significantly bigger than the mean of modern orangutans, and move more upward above the mean of orangutans than subfossil orangutans. The result implies that Pleistocene orangutans from south China are somewhat bigger than modern and subfossil orangutans from Indonesia. Zhou suggests that the body size of orangutan becomes probably smaller during tran-sition of middle to late Pleistocene, but it is still ques-tionable, because bigger teeth exist in Mulanshan Late Pleistocene site and the geological age dating of oran-gutan from Guanyin cave maybe changes as Late Pleis-tocene[18]. Anyway, more materials and dating work are necessary for analyzing the evolutionary trend of Pleis-tocene fossil orangutans in south China.
Pei[19] observed 13 drugstore teeth of fossil orangu-tans from South China and suggested these teeth are larger than those of extant orangutans. Weidenreich studied 5 orangutan teeth of late Pleistocene from He-shang cave in Yunnan and discovered that the occlusal wrinkles tend to be more coarse and sparse than that of extant orangutans[20]. Hooijer[5] studied 3170 subfossil orangutan teeth, early Holocene, from Sibrambang cave, Lida Ajer cave, Djamboe cave, and Padang Highlands, and assigned them as a new subspecies, Pongo pyg-maeus palaeosumatrensis. At the same time he assigned those fossils from China as another new subspecies, Pongo pygmaeus weidenreichi, whose typical feature is the larger tooth size, and the holotype specimen is a lower canine from Heshang cave of Yunnan discovered by Bein and Jia in 1938[21]. With more fossil orangutan teeth discovered in south China, analysis on some of them by Gu et al.[1] and Zhou[2] confirms the features of Pongo pygmaeus weidenreichi. Recently statistics analysis on a large sample of fossil orangutan teeth by Wang et al.[4] indicates the buccal teeth of Pleistocene orangutans from south China are significantly bigger than those of the modern and subfossil orangutans. The present results also show the increasing trend in dental
sizes of hominoids from Mulanshan cave. In addition, the occlusal enamel wrinkles of buccal teeth are rela-tively sparser and simpler when they are compared with the denser and more complex wrinkles in modern and subfossil orangutans.
Schwartz et al.[7] named 4 new subspecies of Pleisto-cene Pongo pygmaeus in Vietnam, P. p. ciochoni (Lang Trang), P. p. devosi (Hang Hum), P. p. fromageti (Tham Om) and P. p. kahlkei (Tham Khuyen), and also a new species P. hooijeri and a new genus and new species Langsonia liquidens. The first author thinks that Viet-nam specimens are within the variations of Pleistocene fossil orangutan-like teeth from south China. Although these new taxonomies have not yet been generally ac-cepted, it reflects the great variations of dental mor-phology of fossil orangutans. Among Pleistocene fossil large-bodied hominoids in south China, except for the unique Gigantopithecus teeth, other orangutan-like teeth have more variability in shape and size, therefore schol-ars have named some species in different genus and spe-cies to distinguish from Pongo, such as mandible frag-ment from Longgupo in Wushan[10,11], Hemianthropus peii teeth from Hong Kong drug stores[22], Australopith-cine teeth from Jianshi of Hubei[23], while a large num-ber of drugstore teeth from Guangdong and Guangxi were generally classified to Pongo sp., which have not been studied in detail. Of these samples, enamel wrinkles of buccal teeth in some are complex and very similar to modern orangutans, and while in some others are more simple and coarse, such as specimens from Mulanshan. Besides the enamel wrinkles, the tooth morphology are changed, a lower molar CZ0408-1 from Mulanshan is comparable and similar to Hemianthropus peii specimens[22]. Is Hemianthropus peii only a kind of variation of Pongo? In addition, the buccal teeth of Lu-fengpithecus are somehow similar with that of Pongo teeth in morphological characters, but skulls and mandi-bles and post-cranial bones are significantly differ- ent[9]. The fossil hominoid teeth from Mulanshan of Chongzuo are significantly different from Lufengpith-ecus teeth in tooth size, and also in the shape of lower central incisor, but the morphology of premolars and molars is somehow similar. In the absence of the skull and postcranial bones, it is difficult to establish new ge-nus or species based on Pleistocene orangutan-like iso-lated teeth. Thus we attribute Mulanshan hominoid specimens to Pongo pygmaeus weidenreichi temporarily. It is possible that more than one species or genus existed
among Pleistocene orangutan-like hominoid teeth from south China, however hard fossil evidence and more analysis research will be the key to holding it in future.
We are grateful to Academician Prof. Wu Xinzhi for detailed discussions on the manuscript. We would like to thank all members of the field team in Chongzuo led by Prof. Jin Changzhu for excavation work, and also thank two anonymous reviewers for comments on the manuscript.
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