Chronological sequence of the early Pleistocene Gigantopithecus faunas from cave sites in the Chongzuo, Zuojiang River area, South China
Introduction
Gigantopithecus blacki is an extinct giant ape known from Pleistocene sites in southern China and Southeast Asia. Von Koenigswald (1935) presented the initial description of the species, based on a lower molar found among “dragon bones” obtained from a Hong Kong drugstore. The ensuing decades saw a heated debate about whether Gigantopithecus was an ape or was more closely related to humans (Weidenreich, 1945, Weidenreich, 1946, Von Koenigswald, 1952, Dart, 1960, Tung, 1962, Pilbeam, 1970, Simons and Ettel, 1970, Eckhardt, 1972, Eckhardt, 1973, Frayer, 1973, Robinson and Steudel, 1973, Corruccini, 1975, Szalay and Delson, 1979, Gelvin, 1980). As a consequence, Gigantopithecus and its associated mammalian fauna attracted much attention. However, the consensus among current researchers is that G. blacki represents a specialized pongine hominoid (Kelley, 2002, Cameron, 2003, Harrison, 2010, Wood and Harrison, 2011, Begun, 2013, Fleagle, 2013).
The typical fauna of the middle Pleistocene of southern China (and more widely distributed in Southeast Asia) was initially described as the “Sino-Malayan fauna” (Von Koenigswald, 1939), but later commonly referred to as the Ailuropoda–Stegodon faunal complex (Teilhard de Chardin et al., 1935, Bien and Chia, 1938, Kahlke, 1961, Pei, 1962). This fauna was considered contemporaneous with the classic Yanjinggou (= Yenchingkou) fauna in Chongqing (formerly Sichuan) (Matthew and Granger, 1923, Colbert and Hooijer, 1953, Chen et al., 2013). Gigantopithecus teeth were commonly found in traditional drug stores associated with other “dragon bones”, including those of Pongo, Ailuropoda, Sinomastodon, and Stegodon, but lacking information on provenance. Scholars referred to this assemblage as the “Drugstore fauna”.
In 1956, Pei and his colleagues carried out large-scale paleontological investigations at Pleistocene cave sites in Tahsin and Liucheng in Guangxi ZAR, southern China, and discovered the first Gigantopithecus remains with well-documented provenance and stratigraphic control (Pei and Woo, 1956, Pei, 1957, Pei and Li, 1958, Han, 1982). Since the 1950s, abundant fossil mammals have been recovered from Quaternary cave sites and fissure deposits in southern China. The fauna from Liucheng, which includes G. blacki, Sinomastodon (formerly gomphotheres) and small-sized tapirs, was considered to be more primitive than the typical middle Pleistocene Ailuropoda–Stegodon fauna (sensu stricto) and was referred to as the Gigantopithecus fauna with an estimated age of early Pleistocene (or latest Pliocene) (Chow, 1957, Pei, 1962).
Today, G. blacki is known from more than fifteen early and middle Pleistocene localities in southern China and northern Vietnam, some in association with hominin fossils and their cultural remains (Woo, 1962, Han, 1982, Huang and Fang, 1991, Ciochon et al., 1996, Zheng, 2004, Wang et al., 2005, Wang et al., 2007a, Wang et al., 2007b, Wang, 2009, Zhao et al., 2006, Zhao et al., 2008, Zhao et al., 2011, Rink et al., 2008, Jin et al., 2009a, Zhao and Zhang, 2013). New discoveries of G. blacki have contributed important information on its anatomy, paleobiology, age, and paleoenvironmental context, as well as on its phylogenetic relationships.
Since 2004, a research team co-organized by the Chinese Academy of Sciences and Peking University has carried out paleontological investigations in the Chongzuo, Zuojiang River area, Guangxi ZAR (Fig. 1). We have succeeded in recovering a diverse vertebrate fauna, including G. blacki and other fossil primates, from five early Pleistocene cave sites (Baikong Cave of Liyu Mountain, Juyuan Cave of Boyue Mountain, Sanhe Cave and Queque Cave of Wuming Mountain, and Wuming Cave of Mulan Mountain) (Fig. 2). These important new finds provide significant information that helps further document the composition, faunal succession, evolutionary history and paleoecology of the Gigantopithecus fauna in southern China.
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Geographical and geological context
The newly discovered early Pleistocene Gigantopithecus cave sites (i.e., Baikong, Boyue, Sanhe and Queque, Fig. 2) are all located in the vicinity of the Chongzuo Ecological Park (22°16.493′N, 107°30.663′E, Figs. 1 and 2) in the Zuojiang River area, Guangxi ZAR, southern China. The park, 16 km southeast of the Chongzuo urban district, is located about 120 km northeast of the China–Vietnam border. Meteorological data show that the area is characterized by a subtropical humid climate, differing
The new Gigantopithecus faunas from Zuojiang River area and their ages
Since 2004, five new cave sites with G. blacki and other vertebrate remains (Harrison et al., 2014, Mead et al., this volume) have been discovered in the vicinity of the Chongzuo Ecological Park (within a radius of less than 3 km). The fossils are derived from early Pleistocene deposits of different ages. The current study presents a preliminary report on four of these sites (i.e., Baikong, Juyuan, Sanhe and Queque Caves). The fifth cave site, Wuming Cave on Mulan Mountain, awaits systematic
Sequence of early Pleistocene Gigantopithecus faunas in southern China
The discovery of new early Pleistocene sites and faunas in southern China (Fig. 7) has led to an improved understanding of G. blacki and its associated fauna, but the absence of comprehensive analyses of the faunas and reliable dating have previously limited our appreciation of the detailed chronology and evolutionary trends.
Based on biostratigraphic and paleomagnetic evidence, the Baikong fauna is estimated to be at least 2.0 Ma. Remains of G. blacki from Longgupo Cave, Chongqing, dated to
Conclusion
Abundant remains of G. blacki and other fossil mammals have recently been recovered from a number of cave sites in the Chongzuo, Zuojiang River area, Guangxi ZAR, South China. These newly discovered Gigantopithecus faunas have estimated dates that span the early Pleistocene (at least from 2 to 1 Ma). The present study divides the early Pleistocene Gigantopithecus faunas into three chronological stages. The early stage (2.6–1.8 Ma) is represented by the Baikong and Boyue faunas, which include G.
Acknowledgements
We are grateful for the discussions or field assistance from Riwan Xie, Lixia Wang, Wei Wang, Jiajian Zheng, Qinqi Xu, Jinyi Liu, Haowen Tong, Yihong Liu, Zhilu Tang, Qiuyuan Wang and Weimin Zheng. We are also thankful for the hard work of the guest editor Dr. Wei Dong and the anonymous reviewers for their constructive comments to improve the manuscript. This work was supported by the Program of Chinese Academy of Sciences (KZZD-EW-03), National Natural Science Foundation of China (41072013,
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