Early Pleistocene large-mammal fauna associated with Gigantopithecus at Mohui Cave, Bubing Basin, South China
Introduction
Gigantopithecus blacki was named by von Koenigswaldv (1935), based on a huge lower third molar found in a pharmacy in Hong Kong. Subsequent in situ discoveries have been made at more than ten karst cave localities since 1956, including Daxin, Liucheng, Wuming, Bama, Jianshi, Longgupo, Mohui, Bijie, Chuifeng, and Sanhe in southern China, and Tham Khuyen in northern Vietnam.
Daxin is the first locality to yield in situ G. blacki in southwest Guangxi. This cave was found by Pei Wenzhong and his field team from the Institute of Vertebrate Paleontology and Paleoanthropology, Chinese Academy of Sciences (IVPP, CAS) in 1955, where three Gigantopithecus teeth were recovered from reddish clay deposits. Because of the 26 associated mammalian species, the fauna is placed into the general category of ‘‘Stegodon/Ailuropoda Fauna’’, which is biostratigraphically estimated to date to the middle Pleistocene (Pei and Woo, 1956, Han, 1982). At Daxin Cave, three species are held in common with the earlier Gigantopithecus occurrence at Liucheng (Pongo sp., Dicoryphochoerus ultimus, and Megalovis guangxiensis), but none of these taxa occur in the Wuming assemblage that is also dated to the middle Pleistocene. In addition, measurements of the Gigantopithecus dentition from Daxin and Wuming are larger than those from the Liucheng series. Zhang (1985) argues that there is a general increase in tooth size within Gigantopithecus from the early Pleistocene to middle Pleistocene (Zhang, 1985). Recently, the Daxin faunal assemblage was dated to 308–380 ka based on coupled ESR and 230Th/234U (Rink et al., 2008).
The Liucheng Cave, also named Gigantopithecus Cave (Juyuandong), was discovered by a local farmer in spring 1957, and subsequently excavated by Pei Wenzhong and his team from 1957 to 1963. This excavation revealed the largest sample of G. blacki (three mandibles and 1006 teeth) associated with a huge number of other mammalian specimens of the early Pleistocene in south China (Woo, 1962, Han, 1987, Pei, 1987). This site has long been recognized to contain an early Early Pleistocene faunal assemblage based on the presence of archaic taxa such as Gomphotherium serridentoides (Sinomastodon yangziensis; Wang et al., 2013), Nestoritherium praesinensis, Tapirus sanyuanensis, Ailuropoda microta, Pachycrocuta licenti, Dorcabune liuchengensis, Megalovis guangxiensis, and Cervavitus (Cervoceros) fenqii (Chow, 1957, Kahlke, 1961, Huang, 1979, Li, 1981, Han and Xu, 1985, Jin et al., 2008).
The Wuming location was found by a hydrogeological survey team from Guangxi in 1965, followed by a test excavation by IVPP the same year. Twelve G. blacki teeth and a small number of other mammal fossils were collected from a hard yellow sandy-clay. Wuming mammalian fauna consists of 14 species, including Ailuropoda melanoleuca fovealis, Stegodon sp., and Rhinoceros sinensis (Zhang et al., 1973). This location is dated to 481–745 ka based on ESR coupled 230Th/234U (Rink et al., 2008).
In the Bama Cave Site, only one G. blacki tooth (lower third molar) was discovered by the survey team that discovered Wuming. The mammalian fauna includes 15 species, containing Ailuropoda melanoleuca baconi and Stegodon sp. This faunal assemblage lacks the ancient taxa found in the Liucheng locality. Zhang et al. (1975) argued that age of the Bama fauna is contemporaneous with that of Wuming, middle Pleistocene age (Zhang et al., 1975).
The Jianshi Cave Site (Longgudong), located in west Hubei Province, central China, has long been known for its rich mammalian fossils. This site was initially discovered and excavated by a team from IVPP in 1970 (Xu et al., 1974), and more recently excavated by a joint IVPP-Hubei Museum team in 1999 and 2000 (Zheng, 2004). Although some teeth were assigned to Australopithecus (Gao, 1975) or Meganthropus (Zhang et al., 2004), the phylogenetic position of these early Asian hominids are actually ambiguous and disputed (Wang, 2009). The mammalian fauna consists of 35 species of large-mammals and 52 species of small-mammals (Zheng, 2004). The geological age of this fauna is estimated biostratigraphically to be middle Early Pleistocene (Xu et al., 1974, Jin et al., 2014).
The Longgupo Site, located in Wushan County of Chongqing City, was discovered and excavated by Huang and his team from 1984 to 1988 (Huang and Fang, 1991). The fauna also includes a mandibular fragment and an isolated incisor originally allocated to Homo. The mandible is now widely considered to be that of an ape, closely related to the Miocene hominoid Lufengpithecus. The mammalian fauna consists of 115 species, including 14 G. blacki teeth. The Longgupo faunal assemblage is similar to that of Liucheng (Han, 1987, Pei, 1987) and Chuifeng (Wang, 2009), biostratigraphically dated to be the early Early Pleistocene (Huang and Fang, 1991, Wei et al., 2014). Paleomagnetic and ESR dating indicate that this mammalian fauna is around 1.92 Ma (Huang et al., 1995). The ESR/U-series dating results show that fossil teeth from different archaeological layers of the lowest geological unit represent an age between ca. 1.4 and 1.8 Ma (Han et al., 2012).
The Bijie Location is located in Guizhou Province, southwest China. One isolated premolar of Gigantopithecus was found by IVPP paleontologists in a drugstore in 1964, but its provenience is unclear. In November 2006, Wei Wang recognized five G. blacki teeth. Zhao et al. (2006) described another tooth of G. blacki from the same site. The large-mammal faunal assemblage includes 5 species of Carnivora (Liu et al., 2011) and 8 species of Artiodactyla (Dong et al., 2010). After comparison to known mammalian faunas in south China, the Bijie fauna is somewhat later in age than that of Longgupo, earlier than that of Jianshi, and close to the age of Liucheng, early Early Pleistocene (Liu et al., 2011, Jin et al., 2014).
The Chuifeng Cave Site is positioned close to the Mohui Cave, with a distance of only 500 m. There are 92 G. blacki teeth associated with 24 species of other mammals at Chuifeng Cave. This faunal assemblage consists of some archaic taxa, such as Hystrix magna, Sinomastodon yangziensis, Stegodon huananensis, Ailuropoda microta, Pachycrocuta licenti, Tapirus sanyuanensis, and Sus peii, which are the typical early Pleistocene elements in south China, similar to the Liucheng and Longgupo faunas (Wang, 2009). The recent ESR results suggest that the lower layers at this cave can be dated to 1.92 ± 0.14 Ma, and the upper layers can be dated to older than 1.38 ± 0.17 Ma (Shao et al., 2014).
The Tham Khuyen Site, located in northern Vietnam, is well-known for its co-occurrence of Homo erectus and G. blacki. The canine of Gigantopithecus is morphologically and metrically similar to those from Liucheng (Ciochon et al., 1996). The mammalian fauna includes H. erectus ssp., G. blacki, and other 28 species, which permits it to be allocated to the “Ailuropoda/Stegodon Fauna” (Cuong, 1971, Schwartz et al., 1994). This site is dated to 475 ka based on ESR and U-series analyses (Ciochon et al., 1996).
More recent discoveries of Gigantopithecus faunas in the Chongzuo area in southwestern Guangxi Zhuang Autonomous Region, come from several sites found by an IVPP team during the past ten years. There have been several cave faunas reported, such as the early Pleistocene Baikong, Boyue, Queque faunas and the middle Pleistocene Hejiang faunas (Jin et al., 2014, Zhang et al., 2014). Systematic and detailed research has been conducted on only the Sanhe Cave fauna (Jin et al., 2008, Jin et al., 2009, Jin et al., 2010, Jin et al., 2014, Zhao et al., 2008, Wang et al., 2009, Wang et al., 2014, Dong et al., 2011, Dong et al., 2013, Mead et al., 2014, Qu et al., 2014). Fifty-six isolated G. blacki teeth associated with 32 species of large mammals and 52 species small mammals have been recovered. The geological age of the deposit is estimated to be middle Early Pleistocene based on faunal and stratigraphic correlation. Paleomagnetic dating of the fossil-bearing strata in the Sanhe Cave provides an age of approximately 1.2 Ma (Jin et al., 2009, Jin et al., 2014, Sun et al., 2014).
During the past half century, these discoveries of G. blacki have provided an unique opportunity for understanding the provenience, and temporal/spatial distribution of this extinct giant ape. However, the associated mammalian fauna is the key to comprehend the biostratigraphical position of Gigantopithecus and in what mammalian community it lived. Thus, finding and researching more Gigantopitcus faunas will permit a better understanding of the role that Gigantopithecus filled in the changing Pleistocene biotic communities of South China.
Here, we report an early Pleistocene mammal fauna obtained from the Mohui Cave of the Bubing Basin in western Guangxi Zhuang Autonomous Region, South China (Fig. 1). This basin is oriented northwest to southeast and is approximately 16 km long by 2 km wide. The Mohui Cave is situated in late Paleozoic limestone hills on the margins of the basin. It was excavated during three field seasons (October to November 2002, November 2003, and July 2008). Mammalian fossil teeth and bones, totaling 412, were recovered in situ in the Mohui Cave. However, 36 teeth were collected from sediments disturbed by local farmers who dug for fertilizer over a half century ago (Wang et al., 2005, Wang et al., 2007a, Wang et al., 2007b, Wang, 2013).
Section snippets
Geological setting
The chamber of Mohui Cave (107°00.13′E, 23°34.891′N) is 50 m in length, 2–6 m in breadth, and 5–6 m in height. The eastward facing cave entrance is 215 m above sea level and 65 m above the nearby 1st terrace within Bubing Basin.
Three areas east to west inside the cave were designated A, B, and C based on the composition of the deposits. A square 1.4 m × 1 m was selected in Area B, and excavated in 5 cm intervals; four squares of 2 m × 2 m were chosen in Area A, and excavated in 10 cm interval;
Fauna assemblage
We define a large-mammal as one in which average adult weight equals or exceeds 0.5 kg. The smaller mammals, mostly small rodents and insectivores, are not included in this study. The Mohui large-mammal fauna consists of 28 genus/species, including Macaca sp., Trachypithecus sp., Hylobates sp., G. blacki, H. magna, Hystrix kiangsenensis, Sinomastodon yangziensis, Stegodon huananensis, Cuon dubius, Arctonyx collaris, Ursus thibetanus, Ailuropoda microta, Pachycrocuta licenti, Panthera sp., Felis
Comparison to local faunas
Table 1 lists the large-mammal taxa in the Mohui fauna and other progressively younger regional faunas. The small Bubing Basin has furnished numerous other Quaternary faunas. Although local faunal comparison was provided in our previous work (Wang et al., 2007a), chronological research and further excavations have recently been carried out by our team. In this comparison, we add chronologically-secured data from three supplemental faunas: late Pleistocene (Luna Cave), middle Pleistocene
Conclusion
The large-mammal assemblage in the Mohui Cave contains a typical early Pleistocene component, including Hystrix magna, Sinomastodon yangziensis, Stegodon huananensis, Ailuropoda microta, Tapirus sanyuanensis, Hespertherium sp., Sus peii, Dorcabune liuchengense, and Cervus fengqii. Biostratigraphically, the Mohui large-mammal fauna should be allocated to the early Pleistocene. Paleomagnetic, ESR, and U-series analysis of this cave indicate the age of the Mohui fauna can be reasonably estimated
Acknowledgments
The authors would like to thank colleagues from the Natural History Museum of Guangxi Zhuang Autonomous Region and the Tiandong County Museum for their careful field excavations, and thank Mr. Hu Pengcheng in Guangxi Museum of Nationalities for his drawing of large-mammal figures. We are grateful for the comments from Jin Changzhu, Jim I. Mead, and three anonymous reviewers. This research is supported by the National Natural Science Foundation of China (Grant Nos. 40163001, 40772011 and 41202017
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