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Chronological sequence of the early Pleistocene Gigantopithecus faunas from cave sites in the Chongzuo, Zuojiang River area, South China
Quaternary International xxx (2014) 1e11
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Chronological sequence of the early Pleistocene Gigantopithecus faunas from cave sites in the Chongzuo, Zuojiang River area, South China Changzhu Jin a, *, Yuan Wang a, *, Chenglong Deng b, Terry Harrison c, Dagong Qin d, Wenshi Pan d, Yingqi Zhang a, Min Zhu a, Yaling Yan a a Key Laboratory of Vertebrate Evolution and Human Origins of Chinese Academy of Sciences, Institute of Vertebrate Paleontology and Paleoanthropology, Chinese Academy of Sciences, 142 Xizhimenwai Street, Beijing 100044, China b State Key Laboratory of Lithospheric Evolution, Institute of Geology and Geophysics, Chinese Academy of Sciences, Beijing 100029, China c Center for the Study of Human Origins, Department of Anthropology, New York University, New York, NY 10003, USA d School of Life Sciences, Peking University, Beijing 100871, China
a r t i c l e i n f o
a b s t r a c t
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Since 2004, five new early Pleistocene cave sites with Gigantopithecus blacki and other fossil mammals have been discovered in the Chongzuo, Zuojiang River area, Guangxi Zhuang Autonomous Region (Guangxi ZAR), South China. These sites include Baikong Cave of Liyu Mountain, Juyuan Cave of Boyue Mountain, Sanhe and Queque Caves of Wuming Mountain, and Wuming Cave of Mulan Mountain. The present study provides a preliminary account of these newly discovered Gigantopithecus faunas, and discusses their taxonomic composition, evolutionary trends and chronology based on recent biostratigraphic and geochronological studies. Combining faunal and paleomagnetic data, the Gigantopithecus faunas from the Zuojiang River area can be divided into three temporal stages during the early Pleistocene. The early stage (2.6e1.8 Ma) is represented by the Baikong and Boyue faunas, which include some Neogene relics (e.g., Sinomastodon, Hesperotherium, Dicoryphochoerus, Dorcabune and Cervavitus) and several species that make their first appearance during the Pleistocene (e.g., Sinicuon dubius, Pachycrocuta licenti, Ailuropoda microta, Sinomastodon jiangnanensis, Stegodon huananensis and Tapirus sanyuanensis). The middle stage (1.8e1.2 Ma) is represented by the Sanhe fauna, which is characterized by the appearance of Ailuropoda wulingshanensis, Cuon antiquus, and Tapirus sinensis. The late stage (1.2e0.8 Ma) is represented by the Queque fauna, which is characterized by a marked decrease in Neogene relics and the appearance of a few species that are more typical of the middle Pleistocene (e.g., Ailuropoda baconi and Stegodon orientalis). Evolutionary trends in the early Pleistocene Gigantopithecus faunas include a gradual increase in dental size in some of the typical taxa, such as G. blacki, Ailuropoda and Tapirus, and the successive turnover of representative species, such as A. microta e A. wulingshanensis e A. baconi. Thus far, Eleven G. blacki sites of early Pleistocene age have been recorded in southern China, and these are primarily concentrated in Guangxi ZAR, where more than 70% of early Pleistocene G. blacki cave sites are known. Given this temporal and geographic distribution it is possible to infer that Guangxi ZAR represents an important early evolutionary and zoogeographic center for G. blacki. Ó 2014 Elsevier Ltd and INQUA. All rights reserved.
1. Introduction
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, 1946; Von Koenigswald, 1952; Dart, 1960; Tung, 1962; Pilbeam, 1970; Simons and Ettel, 1970; Eckhardt, 1972, 1973; Frayer, 1973; Robinson and Steudel, 1973; Corruccini, 1975; Szalay and Delson, 1979; Gelvin, 1980). As a consequence, Gigantopithecus and its
Gigantopithecus blacki is an extinct giant ape known from Pleistocene sites in southern China and Southeast Asia. Von
* Corresponding authors. E-mail addresses: [email protected] (C. Jin), [email protected] (Y. Wang). 1040-6182/$ e see front matter Ó 2014 Elsevier Ltd and INQUA. All rights reserved. http://dx.doi.org/10.1016/j.quaint.2013.12.051
Please cite this article in press as: Jin, C., et al., Chronological sequence of the early Pleistocene Gigantopithecus faunas from cave sites in the Chongzuo, Zuojiang River area, South China, Quaternary International (2014), http://dx.doi.org/10.1016/j.quaint.2013.12.051
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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 AiluropodaeStegodon 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 (¼ 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 AiluropodaeStegodon 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 W. et al., 2005, 2007a, 2007b, 2009; Zhao et al., 2006, 2008, 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. 2. 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 (2216.4930 N, 107 30.6630 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 ChinaeVietnam
Fig. 1. The geographical location of Chongzuo Ecological Park (fieldwork area), Zuojiang River area, South China.
Please cite this article in press as: Jin, C., et al., Chronological sequence of the early Pleistocene Gigantopithecus faunas from cave sites in the Chongzuo, Zuojiang River area, South China, Quaternary International (2014), http://dx.doi.org/10.1016/j.quaint.2013.12.051
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Fig. 2. The geographical distribution of the early Pleistocene Gigantopithecus blacki cave sites in the vicinity of the Chongzuo Ecological Park, Zuojiang River area, South China.
border. Meteorological data show that the area is characterized by a subtropical humid climate, differing from that of Malaysia and Indonesia in the relatively lower average annual temperature, occasional frosts in winter, and the influence of the monsoon. The local flora and fauna belong to the north tropical zone. The fauna has more than 140 species that are protected nationally, including the critically endangered white-headed langur (Trachypithecus leucocephalus), and during the late Pleistocene included a number of locally extinct forms, such as the orangutan and Asian elephant (Jin et al., 2009b). The study of the extant and fossil fauna is significant for reconstructing the paleozoogeography and ecological change through time in the region. The Zuojiang River area was part of the Tethys during the early Mesozoic and is associated with the deposition of carbonate and shallow marine sediments. The limestone of the Majiaoling Formation of the Lower Triassic has a thickness of up to 4500 m. Subsequent late Mesozoic tectonic movement led to the deposition of terrestrial red clastic sediments containing fossil plants and dinosaurs. The northern tropical karsts in the area include two types of landforms: peak valleys and peak depressions. The deeply undercut peak depressions reflect strong vertical movement of water infiltration, leading to a high density of tower karsts and numerous polygonal depressions. The height of the mountains is generally about 300 m above sea level (asl) and the river valley floor is about 145 m asl. On the steep faces of the karst slopes, numerous caves and fissures of various dimensions have been formed, most of which contain fossiliferous deposits. As a consequence of the continuous uplift of the area since the Pliocene, multiple horizons of caves have been formed at different elevations. These show a degree of regularity in the relationship between elevation and time of formation (Jin et al., 2009a,b). Generally, the higher the elevation the older the age of the cave. A total of six horizons of karst caves have been recognized in the Zuojiang River area (Fig. 3). The elevation of the highest horizon (the sixth horizon) is about 270 m asl, and that of the lowest (the first horizon) is about 150 m asl, which is situated only a few meters above the valley floor. It is the caves in the fifth horizon, at an elevation of about 200 m asl, that yield the early Pleistocene
Gigantopithecus fauna, including G. blacki, Sinomastodon, Stegodon, Ailuropoda and Dicoryphochoerus (Jin et al., 2009a). 3. 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., this volume; 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 excavation. 3.1. Baikong Cave Baikong Cave (2217.7550 N, 107 29.9070 E) is located on the western slope of Liyu Mountain near Banrong Village, about 3 km north of the Chongzuo Ecological Park (Fig. 2). Baikong Cave is a large tubular karst cave, about 300 m long and up to 8 m wide. The natural entrance of the cave faces south. The elevation of the cave entrance is 212 m asl, and is located 73 m above the floor of the Hejiang River valley. It is the highest cave in the area with G. blacki remains. Baikong Cave was originally filled with sediments comprising grayish-white speleothems, yellowish-brown silts and numerous layers of limestone breccias, calcareous sediments, brown silty clays and red clays. The stratigraphic sequence can be divided into six layers with a thickness of approximately 5 m (Fig. 4). Fossil G. blacki and other mammals were recovered from the third and fifth layers from the top of the sequence. The mammalian assemblage from Baikong Cave contains 10 orders, 33 families, 60 genera and 64 species, in addition to amphibians, reptiles, and birds. The main characteristics of the Baikong fauna are summarized as follows. (1) The large mammals include several relic Neogene genera, such as Sinomastodon, Hesperotherium, Dicoryphochoerus and Dorcabune. Extinct genera
Please cite this article in press as: Jin, C., et al., Chronological sequence of the early Pleistocene Gigantopithecus faunas from cave sites in the Chongzuo, Zuojiang River area, South China, Quaternary International (2014), http://dx.doi.org/10.1016/j.quaint.2013.12.051
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Fig. 3. The vertical distribution (elevation above sea level) of the karst caves on Mulan Mountain near Chongzuo Ecological Park, Zuojiang River area, South China.
account for more than 35% of the large mammals. (2) The fauna includes species that make their first appearance during the Pleistocene, such as G. blacki, Ailuropoda microta, Sinomastodon jiangnanensis, Sinocuon dubius, Pachycrocuta licenti, Stegodon huananensis, Tapirus sanyuanensis, Sus peii, Sus xiaozhu, Cervavitus fenqii and Megalovis guangxiensis (Jin et al., 2007; Chen, 2011; Wang Y. et al., 2012; Wang Y. et al., this volume). These comprise about 60% of the large mammal species. (3) The abundance of G. blacki specimens (more than 140 isolated teeth) indicates that it was a relatively common species. The teeth of Gigantopithecus appear to be relatively smaller and more primitive than those from younger sites. (4) The primate fauna contains several isolated teeth that are possibly referable to the undescribed “mystery ape” (Ciochon, 2009). These finds are potentially significant for resolving the long-standing debate about whether these enigmatic early Pleistocene hominoids from southern China are pongins or hominins (Jin et al., 2009a; Wang W, personal communication). Liucheng Juyuan Cave, which is the classic early Pleistocene site with a typical “Gigantopithecus fauna”, is located farther to the north of Baikong Cave. Although several Palaearctic taxa, such as Equus, are represented in the Liucheng Juyuan fauna and are absent from the Baikong fauna, the general composition of the two faunas is similar (Han, 1987; Pei, 1987). Both sites include primitive Quaternary species, such as A. microta, P. licenti and T. sanyuanensis. The similarities between the two faunas confirm their contemporaneity, and imply an earliest Pleistocene age. Paleomagnetic analyses demonstrate that the fossil-bearing sediments in Baikong Cave are dominated by a reversed polarity (Fig. 4). Combining the biochronological and magnetostratigraphic evidence the Baikong sediments can be best correlated with the pre-Olduvai Matuyama reverse chron, which has an estimated minimum age of 2.0 Ma (Sun et al., this volume). The Baikong fauna is therefore contemporary with those from Longgupo, Chongqing (Huang and Fang, 1991) and Renzidong, Anhui (Jin and Liu, 2009). Determination of the lower age limit of the Baikong Cave sediments will require further study.
3.2. Juyuan Cave Located on the eastern slope of Boyue Mountain, near the entrance of the Chongzuo Ecological Park (Fig. 2), Juyuan Cave (2217.3650 N, 107 30.6470 E) is an elongated slit-like karst cave oriented southwest to northeast. The natural entrance of the cave faces east. The maximum width of the cave is 3.6 m. The elevation of the cave entrance is 206 m asl, and is situated 63 m above the valley floor. Juyuan Cave, discovered in 2004, was the first site where G. blacki fossils were discovered in the area. The sediments of the cave can be subdivided into six layers with a thickness of approximately 5 m. They are composed of yellowishbrown silty conglomerates and breccias, brown calcareous silts, grayish-white calcareous speleothems, maroon argillaceous silts and red clays. The G. blacki and associated mammalian remains are concentrated in the first and fifth layers from the top of the sequence. After preliminary identification, the mammalian assemblage from Juyuan Cave includes 18 species of large mammals and 26 species of small mammals. The distinctive features of the Juyuan fauna can be listed as follows: (1) The large mammals include several Neogene relics, such as Sinomastodon, Hesperotherium and Dicoryphochoerus. Extinct genera account for about 25% of the large mammals. (2) There is a low proportion of extant species (only 10% of large mammals). (3) Several primitive species (e.g., P. licenti and S. dubius) disappear and S. jiangnanensis is replaced by the more derived Sinomastodon yangziensis. The composition of the Juyuan fauna resembles that from Baikong Cave, with up to 60% of the species shared in common. However, the disappearance of primitive carnivores and the occurrence of the progressive S. yangziensis in the Juyuan fauna indicate a slightly younger age. The paleomagnetic data show that the fossil-bearing sediments in Juyuan Cave are of normal polarity (Fig. 4). In conjunction with the biochronological evidence, this indicates that the Juyuan Cave sediments should be correlated with the Olduvai normal subchron, giving an estimated age of 1.8 Ma (Sun et al., this volume).
Please cite this article in press as: Jin, C., et al., Chronological sequence of the early Pleistocene Gigantopithecus faunas from cave sites in the Chongzuo, Zuojiang River area, South China, Quaternary International (2014), http://dx.doi.org/10.1016/j.quaint.2013.12.051
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Fig. 4. The stratigraphic sections and magnetostratigraphy of the Juyuan Cave of Boyue Mountain and Baikong Cave of Liyu Mountain, and their correlation with the geomagnetic polarity time scale (GPTS).
3.3. Sanhe Cave Sanhe Cave (2216.4930 N, 107 30.6630 E), located on the southern slope of Wuming Mountain in Chongzuo Ecological Park (Fig. 2), is the largest tubular karst cave in the area. The natural entrance of the cave faces south. The cave is about 156 m long from north to south, with a maximum width of 100 m. The elevation of the cave entrance is 203 m asl, and is located 60 m above the valley floor. The sediments are composed of brown sandy clays, grayishwhite speleothems, brown sands, yellow sandy silts with
breccias, yellow lutaceous silts and yellow sandy clays. The stratigraphic sequence can be subdivided into seven layers with a thickness of up to 14 m (Fig. 5). Fossil G. blacki and other mammals were recovered from the fourth and fifth layers from the top (Jin et al., 2009a). The Sanhe fauna includes 32 species of large mammals and 52 species of small mammals. Detailed findings have already been published on the biochronology (Jin et al., 2009a), Gigantopithecus (Zhao et al., 2008), Pongo (Zhao et al., this volume), artiodactyls (Dong et al., 2011, 2013), proboscideans (Wang Y. et al., 2013), small
Please cite this article in press as: Jin, C., et al., Chronological sequence of the early Pleistocene Gigantopithecus faunas from cave sites in the Chongzuo, Zuojiang River area, South China, Quaternary International (2014), http://dx.doi.org/10.1016/j.quaint.2013.12.051
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Fig. 5. The stratigraphic sections and magnetostratigraphy of the Sanhe and Queque Caves of Wuming Mountain, and their correlation with the geomagnetic polarity time scale (GPTS).
mammals (Jin et al., 2008, 2010; Wang Y. et al., 2009), and isotopes analysis of tooth enamel (Qu et al., this volume). The essential features of the fauna can be summarized as follows. (1) The large mammals include a combination of Neogene remnant genera (e.g., Sinomastodon, Dicoryphochoerus and Dorcabune) which account for about 18% of the large mammals and some typical Pleistocene taxa
(e.g., Gigantopithecus, Macaca, Erictis, Stegodon, Megalovis and Bibos). (2) There are 23 extinct species, which account for about 70% of the large mammal species. The proportion is slightly lower than in the Baikong and Juyuan faunas. (3) Several species that make their first appearance in the Pleistocene, such as A. microta, S. dubius and T. sanyuanensis, are replaced by more derived species, such as
Please cite this article in press as: Jin, C., et al., Chronological sequence of the early Pleistocene Gigantopithecus faunas from cave sites in the Chongzuo, Zuojiang River area, South China, Quaternary International (2014), http://dx.doi.org/10.1016/j.quaint.2013.12.051
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Ailuropoda wulingshanensis, Cuon antiquus and Tapirus sinensis. (4) The fauna is composed almost entirely of Oriental taxa, and lacks the typical Palaearctic elements, such as Equus and P. licenti. (5) There is a tendency for an increase in dental size in Gigantopithecus, Ailuropoda and Tapirus (Fig. 6). Paleomagnetic analyses show that the lower and upper parts of the stratigraphic section have a normal polarity and can be correlated with the Olduvai and Jaramillo subchrons respectively, while the fossil-bearing sediments have a reversed polarity and can be correlated with the pre-Jaramillo Matuyama reverse chron (Fig. 5). This correlation provides an estimated age of about 1.2 Ma (Sun et al., this volume). 3.4. Queque Cave Queque Cave (2216.3670 N, 107 30.3670 E) is also located on the southern slope of Wuming Mountain, about 6 km south of Sanhe Cave (Fig. 2). It is a tubular karst cave, with a south facing entrance. The cave is about 26 m long from northeast to southwest, with a maximum width of 8.2 m. The cave entrance is located 196 m asl, about 7 m lower than that of Sanhe Cave. The sediments in the cave can be divided into nine layers with a thickness of approximately 10.8 m. These are composed of three layers of speleothems, grayishbrown breccias, red-brown sands, brown silty conglomerates, grayish-yellow silts and red clays (Fig. 5). The G. blacki specimens and other mammalian remains were recovered from the third to fifth layers from the top of the sequence. Based on preliminary identification, the Queque fauna contains 26 species of large mammals and 38 species of small mammals. The
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characteristic features of the fauna are as follows: (1) There is a decline in the proportion of Neogene relics which account for only about 15% of the large mammals. (2) There is a higher proportion of extant species (30%), which include Arctonyx collaris, Paguma larvata and Cervus (Rusa) yunnanensis. (3) Several typical early Pleistocene species continue to occur, such as S. yangziensis, T. sinensis and M. guangxiensis. (4) Typical middle Pleistocene species appear in the fauna, including Stegodon orientalias, Ailuropoda baconi and Bubalus brevicornis (Dong et al., this volume). (5) There is a marked increase in the dental size of Gigantopithecus, Ailuropoda and Tapirus (Fig. 6). Given the relatively higher proportion of extinct species of large mammals, the Queque fauna can be inferred to be older than the Yanjinggou fauna (Matthew and Granger, 1923; Colbert and Hooijer, 1953; Chen et al., 2013), but younger than the Sanhe fauna. The Queque fauna represents a transitional fauna between the early Pleistocene and middle Pleistocene. The paleomagnetic analysis shows that the fossil-bearing sediments have a normal polarity, which is best correlated to the Jaramillo subchron (Fig. 5). The age of the Queque fauna is, therefore, estimated at about 1.0 Ma, during the late Early Pleistocene (Sun et al., this volume). In summary, the chronological sequence of the new early Pleistocene Gigantopithecus faunas from the Zuojiang River area can be divided into three chronological stages based on a combination of biostratigraphy, geochronology and geomorphology. The early stage, with a correlation between the Gauss-Matuyama boundary (2.58 Ma) and the upper boundary of the Olduvai normal subchron, 1.78 Ma, is represented by the Baikong and Boyue faunas. These faunas include G. blacki with relatively small teeth, as well as A.
Fig. 6. Teeth of Gigantopithecus, Ailuropoda and Tapirus from the early Pleistocene Gigantopithecus faunas in Chongzuo, Zuojiang River area, South China. A, Baikong Cave; B, Juyuan Cave; C, Sanhe Cave; D, Queque Cave. 1e3, Gigantopithecus blacki (M1eM3). A1eA3 (CLBBD201011-833, 834, 837), B1eB3 (CBJ0704-P16, P17, P14), C1eC3 (CSD0410-251, 258, 256), D1eD3 (CSD0704-280, 29, 5). A4 (m3, CLBBD201011-13), A5 (M1, CLBBD201011-5), B4 (m3, CBJ0704-1), B5 (m2, CBJ0704-11), Ailuropoda microta; C4 (m3, CSD0410-568), C5 (P3, CSD0410-561), Ailuropoda wulingshanensis; D4 (m2, CSQ0811a), Ailuropoda baconi. A6 (m2, CLBBD201011-2442), A7 (P4, CLBBD201011-2441), B6 (m3, CBJ0704-17), Tapirus sanyuanensis; C6 (dp4, CSD0704-103a), C7 (m2, CSD0704-103b), D5 (dp3, CSQ0811b), Tapirus sinensis. CLBBD, CBJ, CSD, CSQ, field number.
Please cite this article in press as: Jin, C., et al., Chronological sequence of the early Pleistocene Gigantopithecus faunas from cave sites in the Chongzuo, Zuojiang River area, South China, Quaternary International (2014), http://dx.doi.org/10.1016/j.quaint.2013.12.051
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microta, P. licenti, S. dubius, S. huananensis, and T. sanyuanensis. The middle stage, correlated between the upper boundary of the Olduvai subchron (1.78 Ma) and the Cobb Mountain subchron (w1.2 Ma), is represented by the Sanhe fauna, which includes A. wulingshanensis, S. yangziensis, Cuon antiquus, and T. sinensis. The late stage, correlated between the Cobb Mountain subchron (w1.2 Ma) and the MatuyamaeBrunhes boundary (0.78 Ma), is represented by the Queque fauna that includes G. blacki with relatively large teeth, Cuon antiquus, A. baconi, S. yangziensis, Stegodon orientalis and B. brevicornis. The new Gigantopithecus faunas extend through much of the early Pleistocene (from 2 Ma to 1 Ma), and it possible to discern a number of evolutionary trends, including a gradual increase in the dental size of Gigantopithecus (Zhang, 1982), Ailuropoda and Tapirus (Fig. 6), the successive replacement of key species (e.g., A. microta e A. wulingshanensis e A. baconi), and the occurrence of hominins. 4. Discussion 4.1. 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 1.95 Ma using
magnetostratigraphy and electron spin resonance (ESR) (Huang and Fang, 1991; Huang et al., 1995) are contemporaneous with those from Baikong Cave. The classic early Pleistocene “Gigantopithecus fauna” from Liucheng Juyuan Cave (Han, 1987; Pei, 1987) is also similar to those from Baikong and Longgupo, although a later date of 0.94e1.21 Ma has been inferred based on ESR dating of mammalian teeth from the upper part of the Liucheng Cave sediments (Rink et al., 2008). All three faunas contain primitive Quaternary species, such as A. microta, P. licenti and T. sanyuanensis. The Mohui fauna from Tiandong County, Guangxi ZAR, composed of 19 mammalian species (e.g., G. blacki, A. microta, T. sanyuanensis and S. peii) (Wang W. et al., 2005, 2007a, 2007b), closely resembles the Juyuan fauna on Boyue Mountain. The age of the Mohui fauna is provisionally estimated to around 1.8 Ma based on a combination of paleomagnetic and biostratigraphic data (Wang W. et al., 2005, 2007a). Moreover, the Mohui fauna appears to be slightly younger than the Liucheng Juyuan fauna due to the absence in the Mohui fauna of several common Neogene remnant genera, such as Dicoryphochoerus and Dorcabune. The Chuifeng fauna, also from Tiandong County, Guangxi ZAR is considered to be slightly older than the Mohui fauna because it includes more primitive species and the elevation of the Chuifeng Cave is 11 m higher than that of the Mohui Cave (Wang W. et al., 2009). The Chuifeng fauna is probably contemporaneous with the Liucheng Juyuan fauna. The Baeryan fauna in Bijie, Guizhou includes 10 mammalian taxa, of which only four can be definitively identified at the species level (Zhao et al., 2006; Zhao and Zhang, 2013). Comparisons of Gigantopithecus, Ailuropoda and Tapirus from Baeryan indicate that the fauna is equivalent in age to the Liucheng Juyuan fauna.
Fig. 7. Geographical distribution of early Pleistocene Gigantopithecus blacki sites in southern China. 1e5, Baikong, Juyuan, Sanhe, Queque and Wuming Caves in Chongzuo, Zuojiang River area, Guangxi; 6, Liucheng Juyuan Gigantopithecus Cave, Guangxi; 7e8, Chuifeng and Mohui Caves in Tiandong, Guangxi; 9, Baeryan Cave in Bijie, Guizhou; 10, Longgupo Cave in Wushan, Chongqing; 11, Longgudong Cave in Jianshi, Hubei.
Please cite this article in press as: Jin, C., et al., Chronological sequence of the early Pleistocene Gigantopithecus faunas from cave sites in the Chongzuo, Zuojiang River area, South China, Quaternary International (2014), http://dx.doi.org/10.1016/j.quaint.2013.12.051
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The Longgudong fauna in Jianshi County, Hubei and the Sanhe fauna both contain S. yangziensis, S. huananensis, A. wulingshanensis and this implies that the two faunas are contemporaneous (Zheng, 2004; Jin et al., 2009a). Paleomagnetic studies indicate that the Longgudong fauna is at least 1.2 Ma (Zheng, 2004) or 1.1e1.4 Ma (Shi, 2006). However, the co-occurrence of primitive T. sanyuanensis with T. sinensis, and the slightly lower proportion of extant species of small mammals in the Longgudong fauna (Zheng, 2004) compared to the Sanhe fauna, indicates that the Longgudong fauna is slightly older than the latter (Jin et al., 2009a). To sum up, the chronological sequence of the early Pleistocene Gigantopithecus faunas in southern China (from oldest to youngest), based on a combination of biostratigraphic and geochronological evidence, is as follows: Baikong, Longgupo, Liucheng, Baeryan, Chuifeng, Mohui, Juyuan, Longgudong, Sanhe, and Queque. 4.2. Geographic distribution and paleoecology of the early Pleistocene Gigantopithecus faunas in southern China Eleven G. blacki sites have so far been discovered at early Pleistocene sites in southern China (Fig. 7). The sites are all karst caves or fissures in the Oriental zoogeographic realm associated with mesic tropical and subtropical climatic zones (south of the Yangtze River). The northernmost limit is 30 380 N (i.e., Longgudong Cave in Jianshi, Hubei) and the southern boundary is located at 22160 N (i.e., Queque Cave in Chongzuo, Guangxi). The early Pleistocene G. blacki sites are primarily concentrated in Guangxi ZAR (sites 1e8 in Fig. 7), accounting for more than 70% of the total number of G. blacki fossil sites from this time period. The density of sites in Guangxi ZAR possibly implies that this area was the evolutionary center for G. blacki during the early Pleistocene. Generally, the elevation of the cave entrance in the local setting can be used as a guide to the relative age of the deposits. Cave sites with entrances that have a higher elevation are older than those at lower elevations. This allows geomorphological evidence to support biochronological studies in determining the relative age of cave sites. In the Zuojiang River area, especially in the vicinity of the Chongzuo Ecological Park, the elevations of early Pleistocene G. blacki-bearing caves are located at about 200 m asl, but the elevation varies in different areas. For example, Baeryan Cave in Bijie, Guizhou, on the YunnaneGuizhou Plateau, has the highest elevation at 1600 m asl, and Longgupo Cave in Wushan, Chongqing has the next highest elevation at about 800 m asl. The elevations of Chuifeng and Mohui Caves in the Youjiang River area, Guangxi ZAR are lower than 600 m, but are still substantially higher than those of Zuojiang River area. These major altitudinal differences are presumably related to the late Cenozoic uplift of the Tibetan Plateau. The elevation of the early Pleistocene cave sites declines with greater distance from the YunnaneGuizhou Plateau. Based on the distribution of modern terrestrial vertebrates, China can be divided into two zoogeographical realms, with the approximate eastewest boundary being delimited by the Huaihe River, the Qinling Mountains and the Hengduan Mountains. The Palearctic and Oriental realms are distributed to the north and south of this boundary, respectively. Studies of the faunas demonstrate that the early Pleistocene Gigantopithecus faunas in southern China are representative of diverse ecologies and environments. For example, the composition of the fauna from Sanhe Cave, one of the southernmost locations of the Gigantopithecus faunas in China, is distinctive. The family level composition of the mammalian fauna shows that 56% occur today in the Oriental realm, including Tupaiidae, Ailuridae, Pongidae and Platacanthomyidae. At the genus level, 56 genera (more than 87% of the total) belong to the Oriental realm, which include Tupaia, Ia, Ailuropoda, Pongo, Nomascus and Trachypithecus. Comparisons at the species
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level show that typical Palaearctic species are absent, except widespread taxa. The mammalian fauna from Sanhe Cave can be considered the most typical tropical rainforest fauna from the Quaternary of southern China (Jin et al., 2009a). By contrast, Longgudong Cave, which is over 1300 km north of Sanhe, has a mammalian fauna characterized by a combination of Palaearctic taxa (e.g., Erinaceus, Pachycrocuta, Equus, Leptobos and Ochotona) and Oriental taxa (e.g., Ailuropoda, Stegodon and Rhizomys), and demonstrates its distinctiveness in the transitional zone between the two major zoogeographic realms. This transitional zone was located at approximately 30 N during the early Pleistocene, but today is situated further to the south, as it was during the Neogene. 5. 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 (2.6e1.8 Ma) is represented by the Baikong and Boyue faunas, which include G. blacki with relatively small teeth, some Neogene relics (e.g., Sinomastodon, Hesperotherium and Dicoryphochoerus), and taxa that make their first appearance during the Pleistocene (e.g., A. microta, S. dubius, S. huananensis, and T. sanyuanensis). The middle stage (1.8e 1.2 Ma) is represented by the Sanhe fauna, which is characterized by the appearance of A. wulingshanensis, C. antiquus, and T. sinensis. The late stage (1.2e0.8 Ma) is represented by the Queque fauna, which exhibits a distinct decrease in Neogene relic taxa and the appearance of several typical middle Pleistocene taxa, such as A. baconi, S. orientalis and B. brevicornis. The early Pleistocene Gigantopithecus faunas are also characterized by a gradual increase in dental size of some typical faunal members, such as Gigantopithecus, Ailuropoda and Tapirus, and the successive replacement of representative species, such as A. microta e A. wulingshanensis e A. baconi. The geographic distribution of G. blacki in the early Pleistocene is limited to the Oriental realm of southern China (south of the Yangtze River). Eleven G. blacki fossil sites, representing different periods of the early Pleistocene, have been recorded in southern China. The following chronological sequence is proposed for these sites (from oldest to youngest): Baikong, Longgupo, Liucheng, Baeryan, Chuifeng, Mohui and Juyuan (early Early Pleistocene), Longgudong and Sanhe (middle Early Pleistocene), and Queque (late Early Pleistocene). Early Pleistocene Gigantopithecus sites occur predominantly in Guangxi ZAR in southern China, accounting for more than 70% of all known G. blacki caves from this time period. This may imply that Guangxi ZAR represents a possible center of evolution and dispersal for G. blacki. Preliminary analysis of the dental metrics of G. blacki indicates that the teeth (especially the molars) increase in average size during the early Pleistocene from the early stage to the late stage. However, the significance of this observed tendency will require further study and the recovery of additional material of Gigantopithecus. 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
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Chronological sequence of the early Pleistocene Gigantopithecus faunas from cave sites in the Chongzuo, Zuojiang River area, South China Changzhu Jin a, *, Yuan Wang a, *, Chenglong Deng b, Terry Harrison c, Dagong Qin d, Wenshi Pan d, Yingqi Zhang a, Min Zhu a, Yaling Yan a a Key Laboratory of Vertebrate Evolution and Human Origins of Chinese Academy of Sciences, Institute of Vertebrate Paleontology and Paleoanthropology, Chinese Academy of Sciences, 142 Xizhimenwai Street, Beijing 100044, China b State Key Laboratory of Lithospheric Evolution, Institute of Geology and Geophysics, Chinese Academy of Sciences, Beijing 100029, China c Center for the Study of Human Origins, Department of Anthropology, New York University, New York, NY 10003, USA d School of Life Sciences, Peking University, Beijing 100871, China
a r t i c l e i n f o
a b s t r a c t
Article history: Available online xxx
Since 2004, five new early Pleistocene cave sites with Gigantopithecus blacki and other fossil mammals have been discovered in the Chongzuo, Zuojiang River area, Guangxi Zhuang Autonomous Region (Guangxi ZAR), South China. These sites include Baikong Cave of Liyu Mountain, Juyuan Cave of Boyue Mountain, Sanhe and Queque Caves of Wuming Mountain, and Wuming Cave of Mulan Mountain. The present study provides a preliminary account of these newly discovered Gigantopithecus faunas, and discusses their taxonomic composition, evolutionary trends and chronology based on recent biostratigraphic and geochronological studies. Combining faunal and paleomagnetic data, the Gigantopithecus faunas from the Zuojiang River area can be divided into three temporal stages during the early Pleistocene. The early stage (2.6e1.8 Ma) is represented by the Baikong and Boyue faunas, which include some Neogene relics (e.g., Sinomastodon, Hesperotherium, Dicoryphochoerus, Dorcabune and Cervavitus) and several species that make their first appearance during the Pleistocene (e.g., Sinicuon dubius, Pachycrocuta licenti, Ailuropoda microta, Sinomastodon jiangnanensis, Stegodon huananensis and Tapirus sanyuanensis). The middle stage (1.8e1.2 Ma) is represented by the Sanhe fauna, which is characterized by the appearance of Ailuropoda wulingshanensis, Cuon antiquus, and Tapirus sinensis. The late stage (1.2e0.8 Ma) is represented by the Queque fauna, which is characterized by a marked decrease in Neogene relics and the appearance of a few species that are more typical of the middle Pleistocene (e.g., Ailuropoda baconi and Stegodon orientalis). Evolutionary trends in the early Pleistocene Gigantopithecus faunas include a gradual increase in dental size in some of the typical taxa, such as G. blacki, Ailuropoda and Tapirus, and the successive turnover of representative species, such as A. microta e A. wulingshanensis e A. baconi. Thus far, Eleven G. blacki sites of early Pleistocene age have been recorded in southern China, and these are primarily concentrated in Guangxi ZAR, where more than 70% of early Pleistocene G. blacki cave sites are known. Given this temporal and geographic distribution it is possible to infer that Guangxi ZAR represents an important early evolutionary and zoogeographic center for G. blacki. Ó 2014 Elsevier Ltd and INQUA. All rights reserved.
1. Introduction
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, 1946; Von Koenigswald, 1952; Dart, 1960; Tung, 1962; Pilbeam, 1970; Simons and Ettel, 1970; Eckhardt, 1972, 1973; Frayer, 1973; Robinson and Steudel, 1973; Corruccini, 1975; Szalay and Delson, 1979; Gelvin, 1980). As a consequence, Gigantopithecus and its
Gigantopithecus blacki is an extinct giant ape known from Pleistocene sites in southern China and Southeast Asia. Von
* Corresponding authors. E-mail addresses: [email protected] (C. Jin), [email protected] (Y. Wang). 1040-6182/$ e see front matter Ó 2014 Elsevier Ltd and INQUA. All rights reserved. http://dx.doi.org/10.1016/j.quaint.2013.12.051
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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 AiluropodaeStegodon 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 (¼ 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 AiluropodaeStegodon 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 W. et al., 2005, 2007a, 2007b, 2009; Zhao et al., 2006, 2008, 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. 2. 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 (2216.4930 N, 107 30.6630 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 ChinaeVietnam
Fig. 1. The geographical location of Chongzuo Ecological Park (fieldwork area), Zuojiang River area, South China.
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Fig. 2. The geographical distribution of the early Pleistocene Gigantopithecus blacki cave sites in the vicinity of the Chongzuo Ecological Park, Zuojiang River area, South China.
border. Meteorological data show that the area is characterized by a subtropical humid climate, differing from that of Malaysia and Indonesia in the relatively lower average annual temperature, occasional frosts in winter, and the influence of the monsoon. The local flora and fauna belong to the north tropical zone. The fauna has more than 140 species that are protected nationally, including the critically endangered white-headed langur (Trachypithecus leucocephalus), and during the late Pleistocene included a number of locally extinct forms, such as the orangutan and Asian elephant (Jin et al., 2009b). The study of the extant and fossil fauna is significant for reconstructing the paleozoogeography and ecological change through time in the region. The Zuojiang River area was part of the Tethys during the early Mesozoic and is associated with the deposition of carbonate and shallow marine sediments. The limestone of the Majiaoling Formation of the Lower Triassic has a thickness of up to 4500 m. Subsequent late Mesozoic tectonic movement led to the deposition of terrestrial red clastic sediments containing fossil plants and dinosaurs. The northern tropical karsts in the area include two types of landforms: peak valleys and peak depressions. The deeply undercut peak depressions reflect strong vertical movement of water infiltration, leading to a high density of tower karsts and numerous polygonal depressions. The height of the mountains is generally about 300 m above sea level (asl) and the river valley floor is about 145 m asl. On the steep faces of the karst slopes, numerous caves and fissures of various dimensions have been formed, most of which contain fossiliferous deposits. As a consequence of the continuous uplift of the area since the Pliocene, multiple horizons of caves have been formed at different elevations. These show a degree of regularity in the relationship between elevation and time of formation (Jin et al., 2009a,b). Generally, the higher the elevation the older the age of the cave. A total of six horizons of karst caves have been recognized in the Zuojiang River area (Fig. 3). The elevation of the highest horizon (the sixth horizon) is about 270 m asl, and that of the lowest (the first horizon) is about 150 m asl, which is situated only a few meters above the valley floor. It is the caves in the fifth horizon, at an elevation of about 200 m asl, that yield the early Pleistocene
Gigantopithecus fauna, including G. blacki, Sinomastodon, Stegodon, Ailuropoda and Dicoryphochoerus (Jin et al., 2009a). 3. 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., this volume; 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 excavation. 3.1. Baikong Cave Baikong Cave (2217.7550 N, 107 29.9070 E) is located on the western slope of Liyu Mountain near Banrong Village, about 3 km north of the Chongzuo Ecological Park (Fig. 2). Baikong Cave is a large tubular karst cave, about 300 m long and up to 8 m wide. The natural entrance of the cave faces south. The elevation of the cave entrance is 212 m asl, and is located 73 m above the floor of the Hejiang River valley. It is the highest cave in the area with G. blacki remains. Baikong Cave was originally filled with sediments comprising grayish-white speleothems, yellowish-brown silts and numerous layers of limestone breccias, calcareous sediments, brown silty clays and red clays. The stratigraphic sequence can be divided into six layers with a thickness of approximately 5 m (Fig. 4). Fossil G. blacki and other mammals were recovered from the third and fifth layers from the top of the sequence. The mammalian assemblage from Baikong Cave contains 10 orders, 33 families, 60 genera and 64 species, in addition to amphibians, reptiles, and birds. The main characteristics of the Baikong fauna are summarized as follows. (1) The large mammals include several relic Neogene genera, such as Sinomastodon, Hesperotherium, Dicoryphochoerus and Dorcabune. Extinct genera
Please cite this article in press as: Jin, C., et al., Chronological sequence of the early Pleistocene Gigantopithecus faunas from cave sites in the Chongzuo, Zuojiang River area, South China, Quaternary International (2014), http://dx.doi.org/10.1016/j.quaint.2013.12.051
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Fig. 3. The vertical distribution (elevation above sea level) of the karst caves on Mulan Mountain near Chongzuo Ecological Park, Zuojiang River area, South China.
account for more than 35% of the large mammals. (2) The fauna includes species that make their first appearance during the Pleistocene, such as G. blacki, Ailuropoda microta, Sinomastodon jiangnanensis, Sinocuon dubius, Pachycrocuta licenti, Stegodon huananensis, Tapirus sanyuanensis, Sus peii, Sus xiaozhu, Cervavitus fenqii and Megalovis guangxiensis (Jin et al., 2007; Chen, 2011; Wang Y. et al., 2012; Wang Y. et al., this volume). These comprise about 60% of the large mammal species. (3) The abundance of G. blacki specimens (more than 140 isolated teeth) indicates that it was a relatively common species. The teeth of Gigantopithecus appear to be relatively smaller and more primitive than those from younger sites. (4) The primate fauna contains several isolated teeth that are possibly referable to the undescribed “mystery ape” (Ciochon, 2009). These finds are potentially significant for resolving the long-standing debate about whether these enigmatic early Pleistocene hominoids from southern China are pongins or hominins (Jin et al., 2009a; Wang W, personal communication). Liucheng Juyuan Cave, which is the classic early Pleistocene site with a typical “Gigantopithecus fauna”, is located farther to the north of Baikong Cave. Although several Palaearctic taxa, such as Equus, are represented in the Liucheng Juyuan fauna and are absent from the Baikong fauna, the general composition of the two faunas is similar (Han, 1987; Pei, 1987). Both sites include primitive Quaternary species, such as A. microta, P. licenti and T. sanyuanensis. The similarities between the two faunas confirm their contemporaneity, and imply an earliest Pleistocene age. Paleomagnetic analyses demonstrate that the fossil-bearing sediments in Baikong Cave are dominated by a reversed polarity (Fig. 4). Combining the biochronological and magnetostratigraphic evidence the Baikong sediments can be best correlated with the pre-Olduvai Matuyama reverse chron, which has an estimated minimum age of 2.0 Ma (Sun et al., this volume). The Baikong fauna is therefore contemporary with those from Longgupo, Chongqing (Huang and Fang, 1991) and Renzidong, Anhui (Jin and Liu, 2009). Determination of the lower age limit of the Baikong Cave sediments will require further study.
3.2. Juyuan Cave Located on the eastern slope of Boyue Mountain, near the entrance of the Chongzuo Ecological Park (Fig. 2), Juyuan Cave (2217.3650 N, 107 30.6470 E) is an elongated slit-like karst cave oriented southwest to northeast. The natural entrance of the cave faces east. The maximum width of the cave is 3.6 m. The elevation of the cave entrance is 206 m asl, and is situated 63 m above the valley floor. Juyuan Cave, discovered in 2004, was the first site where G. blacki fossils were discovered in the area. The sediments of the cave can be subdivided into six layers with a thickness of approximately 5 m. They are composed of yellowishbrown silty conglomerates and breccias, brown calcareous silts, grayish-white calcareous speleothems, maroon argillaceous silts and red clays. The G. blacki and associated mammalian remains are concentrated in the first and fifth layers from the top of the sequence. After preliminary identification, the mammalian assemblage from Juyuan Cave includes 18 species of large mammals and 26 species of small mammals. The distinctive features of the Juyuan fauna can be listed as follows: (1) The large mammals include several Neogene relics, such as Sinomastodon, Hesperotherium and Dicoryphochoerus. Extinct genera account for about 25% of the large mammals. (2) There is a low proportion of extant species (only 10% of large mammals). (3) Several primitive species (e.g., P. licenti and S. dubius) disappear and S. jiangnanensis is replaced by the more derived Sinomastodon yangziensis. The composition of the Juyuan fauna resembles that from Baikong Cave, with up to 60% of the species shared in common. However, the disappearance of primitive carnivores and the occurrence of the progressive S. yangziensis in the Juyuan fauna indicate a slightly younger age. The paleomagnetic data show that the fossil-bearing sediments in Juyuan Cave are of normal polarity (Fig. 4). In conjunction with the biochronological evidence, this indicates that the Juyuan Cave sediments should be correlated with the Olduvai normal subchron, giving an estimated age of 1.8 Ma (Sun et al., this volume).
Please cite this article in press as: Jin, C., et al., Chronological sequence of the early Pleistocene Gigantopithecus faunas from cave sites in the Chongzuo, Zuojiang River area, South China, Quaternary International (2014), http://dx.doi.org/10.1016/j.quaint.2013.12.051
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Fig. 4. The stratigraphic sections and magnetostratigraphy of the Juyuan Cave of Boyue Mountain and Baikong Cave of Liyu Mountain, and their correlation with the geomagnetic polarity time scale (GPTS).
3.3. Sanhe Cave Sanhe Cave (2216.4930 N, 107 30.6630 E), located on the southern slope of Wuming Mountain in Chongzuo Ecological Park (Fig. 2), is the largest tubular karst cave in the area. The natural entrance of the cave faces south. The cave is about 156 m long from north to south, with a maximum width of 100 m. The elevation of the cave entrance is 203 m asl, and is located 60 m above the valley floor. The sediments are composed of brown sandy clays, grayishwhite speleothems, brown sands, yellow sandy silts with
breccias, yellow lutaceous silts and yellow sandy clays. The stratigraphic sequence can be subdivided into seven layers with a thickness of up to 14 m (Fig. 5). Fossil G. blacki and other mammals were recovered from the fourth and fifth layers from the top (Jin et al., 2009a). The Sanhe fauna includes 32 species of large mammals and 52 species of small mammals. Detailed findings have already been published on the biochronology (Jin et al., 2009a), Gigantopithecus (Zhao et al., 2008), Pongo (Zhao et al., this volume), artiodactyls (Dong et al., 2011, 2013), proboscideans (Wang Y. et al., 2013), small
Please cite this article in press as: Jin, C., et al., Chronological sequence of the early Pleistocene Gigantopithecus faunas from cave sites in the Chongzuo, Zuojiang River area, South China, Quaternary International (2014), http://dx.doi.org/10.1016/j.quaint.2013.12.051
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Fig. 5. The stratigraphic sections and magnetostratigraphy of the Sanhe and Queque Caves of Wuming Mountain, and their correlation with the geomagnetic polarity time scale (GPTS).
mammals (Jin et al., 2008, 2010; Wang Y. et al., 2009), and isotopes analysis of tooth enamel (Qu et al., this volume). The essential features of the fauna can be summarized as follows. (1) The large mammals include a combination of Neogene remnant genera (e.g., Sinomastodon, Dicoryphochoerus and Dorcabune) which account for about 18% of the large mammals and some typical Pleistocene taxa
(e.g., Gigantopithecus, Macaca, Erictis, Stegodon, Megalovis and Bibos). (2) There are 23 extinct species, which account for about 70% of the large mammal species. The proportion is slightly lower than in the Baikong and Juyuan faunas. (3) Several species that make their first appearance in the Pleistocene, such as A. microta, S. dubius and T. sanyuanensis, are replaced by more derived species, such as
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Ailuropoda wulingshanensis, Cuon antiquus and Tapirus sinensis. (4) The fauna is composed almost entirely of Oriental taxa, and lacks the typical Palaearctic elements, such as Equus and P. licenti. (5) There is a tendency for an increase in dental size in Gigantopithecus, Ailuropoda and Tapirus (Fig. 6). Paleomagnetic analyses show that the lower and upper parts of the stratigraphic section have a normal polarity and can be correlated with the Olduvai and Jaramillo subchrons respectively, while the fossil-bearing sediments have a reversed polarity and can be correlated with the pre-Jaramillo Matuyama reverse chron (Fig. 5). This correlation provides an estimated age of about 1.2 Ma (Sun et al., this volume). 3.4. Queque Cave Queque Cave (2216.3670 N, 107 30.3670 E) is also located on the southern slope of Wuming Mountain, about 6 km south of Sanhe Cave (Fig. 2). It is a tubular karst cave, with a south facing entrance. The cave is about 26 m long from northeast to southwest, with a maximum width of 8.2 m. The cave entrance is located 196 m asl, about 7 m lower than that of Sanhe Cave. The sediments in the cave can be divided into nine layers with a thickness of approximately 10.8 m. These are composed of three layers of speleothems, grayishbrown breccias, red-brown sands, brown silty conglomerates, grayish-yellow silts and red clays (Fig. 5). The G. blacki specimens and other mammalian remains were recovered from the third to fifth layers from the top of the sequence. Based on preliminary identification, the Queque fauna contains 26 species of large mammals and 38 species of small mammals. The
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characteristic features of the fauna are as follows: (1) There is a decline in the proportion of Neogene relics which account for only about 15% of the large mammals. (2) There is a higher proportion of extant species (30%), which include Arctonyx collaris, Paguma larvata and Cervus (Rusa) yunnanensis. (3) Several typical early Pleistocene species continue to occur, such as S. yangziensis, T. sinensis and M. guangxiensis. (4) Typical middle Pleistocene species appear in the fauna, including Stegodon orientalias, Ailuropoda baconi and Bubalus brevicornis (Dong et al., this volume). (5) There is a marked increase in the dental size of Gigantopithecus, Ailuropoda and Tapirus (Fig. 6). Given the relatively higher proportion of extinct species of large mammals, the Queque fauna can be inferred to be older than the Yanjinggou fauna (Matthew and Granger, 1923; Colbert and Hooijer, 1953; Chen et al., 2013), but younger than the Sanhe fauna. The Queque fauna represents a transitional fauna between the early Pleistocene and middle Pleistocene. The paleomagnetic analysis shows that the fossil-bearing sediments have a normal polarity, which is best correlated to the Jaramillo subchron (Fig. 5). The age of the Queque fauna is, therefore, estimated at about 1.0 Ma, during the late Early Pleistocene (Sun et al., this volume). In summary, the chronological sequence of the new early Pleistocene Gigantopithecus faunas from the Zuojiang River area can be divided into three chronological stages based on a combination of biostratigraphy, geochronology and geomorphology. The early stage, with a correlation between the Gauss-Matuyama boundary (2.58 Ma) and the upper boundary of the Olduvai normal subchron, 1.78 Ma, is represented by the Baikong and Boyue faunas. These faunas include G. blacki with relatively small teeth, as well as A.
Fig. 6. Teeth of Gigantopithecus, Ailuropoda and Tapirus from the early Pleistocene Gigantopithecus faunas in Chongzuo, Zuojiang River area, South China. A, Baikong Cave; B, Juyuan Cave; C, Sanhe Cave; D, Queque Cave. 1e3, Gigantopithecus blacki (M1eM3). A1eA3 (CLBBD201011-833, 834, 837), B1eB3 (CBJ0704-P16, P17, P14), C1eC3 (CSD0410-251, 258, 256), D1eD3 (CSD0704-280, 29, 5). A4 (m3, CLBBD201011-13), A5 (M1, CLBBD201011-5), B4 (m3, CBJ0704-1), B5 (m2, CBJ0704-11), Ailuropoda microta; C4 (m3, CSD0410-568), C5 (P3, CSD0410-561), Ailuropoda wulingshanensis; D4 (m2, CSQ0811a), Ailuropoda baconi. A6 (m2, CLBBD201011-2442), A7 (P4, CLBBD201011-2441), B6 (m3, CBJ0704-17), Tapirus sanyuanensis; C6 (dp4, CSD0704-103a), C7 (m2, CSD0704-103b), D5 (dp3, CSQ0811b), Tapirus sinensis. CLBBD, CBJ, CSD, CSQ, field number.
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microta, P. licenti, S. dubius, S. huananensis, and T. sanyuanensis. The middle stage, correlated between the upper boundary of the Olduvai subchron (1.78 Ma) and the Cobb Mountain subchron (w1.2 Ma), is represented by the Sanhe fauna, which includes A. wulingshanensis, S. yangziensis, Cuon antiquus, and T. sinensis. The late stage, correlated between the Cobb Mountain subchron (w1.2 Ma) and the MatuyamaeBrunhes boundary (0.78 Ma), is represented by the Queque fauna that includes G. blacki with relatively large teeth, Cuon antiquus, A. baconi, S. yangziensis, Stegodon orientalis and B. brevicornis. The new Gigantopithecus faunas extend through much of the early Pleistocene (from 2 Ma to 1 Ma), and it possible to discern a number of evolutionary trends, including a gradual increase in the dental size of Gigantopithecus (Zhang, 1982), Ailuropoda and Tapirus (Fig. 6), the successive replacement of key species (e.g., A. microta e A. wulingshanensis e A. baconi), and the occurrence of hominins. 4. Discussion 4.1. 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 1.95 Ma using
magnetostratigraphy and electron spin resonance (ESR) (Huang and Fang, 1991; Huang et al., 1995) are contemporaneous with those from Baikong Cave. The classic early Pleistocene “Gigantopithecus fauna” from Liucheng Juyuan Cave (Han, 1987; Pei, 1987) is also similar to those from Baikong and Longgupo, although a later date of 0.94e1.21 Ma has been inferred based on ESR dating of mammalian teeth from the upper part of the Liucheng Cave sediments (Rink et al., 2008). All three faunas contain primitive Quaternary species, such as A. microta, P. licenti and T. sanyuanensis. The Mohui fauna from Tiandong County, Guangxi ZAR, composed of 19 mammalian species (e.g., G. blacki, A. microta, T. sanyuanensis and S. peii) (Wang W. et al., 2005, 2007a, 2007b), closely resembles the Juyuan fauna on Boyue Mountain. The age of the Mohui fauna is provisionally estimated to around 1.8 Ma based on a combination of paleomagnetic and biostratigraphic data (Wang W. et al., 2005, 2007a). Moreover, the Mohui fauna appears to be slightly younger than the Liucheng Juyuan fauna due to the absence in the Mohui fauna of several common Neogene remnant genera, such as Dicoryphochoerus and Dorcabune. The Chuifeng fauna, also from Tiandong County, Guangxi ZAR is considered to be slightly older than the Mohui fauna because it includes more primitive species and the elevation of the Chuifeng Cave is 11 m higher than that of the Mohui Cave (Wang W. et al., 2009). The Chuifeng fauna is probably contemporaneous with the Liucheng Juyuan fauna. The Baeryan fauna in Bijie, Guizhou includes 10 mammalian taxa, of which only four can be definitively identified at the species level (Zhao et al., 2006; Zhao and Zhang, 2013). Comparisons of Gigantopithecus, Ailuropoda and Tapirus from Baeryan indicate that the fauna is equivalent in age to the Liucheng Juyuan fauna.
Fig. 7. Geographical distribution of early Pleistocene Gigantopithecus blacki sites in southern China. 1e5, Baikong, Juyuan, Sanhe, Queque and Wuming Caves in Chongzuo, Zuojiang River area, Guangxi; 6, Liucheng Juyuan Gigantopithecus Cave, Guangxi; 7e8, Chuifeng and Mohui Caves in Tiandong, Guangxi; 9, Baeryan Cave in Bijie, Guizhou; 10, Longgupo Cave in Wushan, Chongqing; 11, Longgudong Cave in Jianshi, Hubei.
Please cite this article in press as: Jin, C., et al., Chronological sequence of the early Pleistocene Gigantopithecus faunas from cave sites in the Chongzuo, Zuojiang River area, South China, Quaternary International (2014), http://dx.doi.org/10.1016/j.quaint.2013.12.051
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The Longgudong fauna in Jianshi County, Hubei and the Sanhe fauna both contain S. yangziensis, S. huananensis, A. wulingshanensis and this implies that the two faunas are contemporaneous (Zheng, 2004; Jin et al., 2009a). Paleomagnetic studies indicate that the Longgudong fauna is at least 1.2 Ma (Zheng, 2004) or 1.1e1.4 Ma (Shi, 2006). However, the co-occurrence of primitive T. sanyuanensis with T. sinensis, and the slightly lower proportion of extant species of small mammals in the Longgudong fauna (Zheng, 2004) compared to the Sanhe fauna, indicates that the Longgudong fauna is slightly older than the latter (Jin et al., 2009a). To sum up, the chronological sequence of the early Pleistocene Gigantopithecus faunas in southern China (from oldest to youngest), based on a combination of biostratigraphic and geochronological evidence, is as follows: Baikong, Longgupo, Liucheng, Baeryan, Chuifeng, Mohui, Juyuan, Longgudong, Sanhe, and Queque. 4.2. Geographic distribution and paleoecology of the early Pleistocene Gigantopithecus faunas in southern China Eleven G. blacki sites have so far been discovered at early Pleistocene sites in southern China (Fig. 7). The sites are all karst caves or fissures in the Oriental zoogeographic realm associated with mesic tropical and subtropical climatic zones (south of the Yangtze River). The northernmost limit is 30 380 N (i.e., Longgudong Cave in Jianshi, Hubei) and the southern boundary is located at 22160 N (i.e., Queque Cave in Chongzuo, Guangxi). The early Pleistocene G. blacki sites are primarily concentrated in Guangxi ZAR (sites 1e8 in Fig. 7), accounting for more than 70% of the total number of G. blacki fossil sites from this time period. The density of sites in Guangxi ZAR possibly implies that this area was the evolutionary center for G. blacki during the early Pleistocene. Generally, the elevation of the cave entrance in the local setting can be used as a guide to the relative age of the deposits. Cave sites with entrances that have a higher elevation are older than those at lower elevations. This allows geomorphological evidence to support biochronological studies in determining the relative age of cave sites. In the Zuojiang River area, especially in the vicinity of the Chongzuo Ecological Park, the elevations of early Pleistocene G. blacki-bearing caves are located at about 200 m asl, but the elevation varies in different areas. For example, Baeryan Cave in Bijie, Guizhou, on the YunnaneGuizhou Plateau, has the highest elevation at 1600 m asl, and Longgupo Cave in Wushan, Chongqing has the next highest elevation at about 800 m asl. The elevations of Chuifeng and Mohui Caves in the Youjiang River area, Guangxi ZAR are lower than 600 m, but are still substantially higher than those of Zuojiang River area. These major altitudinal differences are presumably related to the late Cenozoic uplift of the Tibetan Plateau. The elevation of the early Pleistocene cave sites declines with greater distance from the YunnaneGuizhou Plateau. Based on the distribution of modern terrestrial vertebrates, China can be divided into two zoogeographical realms, with the approximate eastewest boundary being delimited by the Huaihe River, the Qinling Mountains and the Hengduan Mountains. The Palearctic and Oriental realms are distributed to the north and south of this boundary, respectively. Studies of the faunas demonstrate that the early Pleistocene Gigantopithecus faunas in southern China are representative of diverse ecologies and environments. For example, the composition of the fauna from Sanhe Cave, one of the southernmost locations of the Gigantopithecus faunas in China, is distinctive. The family level composition of the mammalian fauna shows that 56% occur today in the Oriental realm, including Tupaiidae, Ailuridae, Pongidae and Platacanthomyidae. At the genus level, 56 genera (more than 87% of the total) belong to the Oriental realm, which include Tupaia, Ia, Ailuropoda, Pongo, Nomascus and Trachypithecus. Comparisons at the species
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level show that typical Palaearctic species are absent, except widespread taxa. The mammalian fauna from Sanhe Cave can be considered the most typical tropical rainforest fauna from the Quaternary of southern China (Jin et al., 2009a). By contrast, Longgudong Cave, which is over 1300 km north of Sanhe, has a mammalian fauna characterized by a combination of Palaearctic taxa (e.g., Erinaceus, Pachycrocuta, Equus, Leptobos and Ochotona) and Oriental taxa (e.g., Ailuropoda, Stegodon and Rhizomys), and demonstrates its distinctiveness in the transitional zone between the two major zoogeographic realms. This transitional zone was located at approximately 30 N during the early Pleistocene, but today is situated further to the south, as it was during the Neogene. 5. 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 (2.6e1.8 Ma) is represented by the Baikong and Boyue faunas, which include G. blacki with relatively small teeth, some Neogene relics (e.g., Sinomastodon, Hesperotherium and Dicoryphochoerus), and taxa that make their first appearance during the Pleistocene (e.g., A. microta, S. dubius, S. huananensis, and T. sanyuanensis). The middle stage (1.8e 1.2 Ma) is represented by the Sanhe fauna, which is characterized by the appearance of A. wulingshanensis, C. antiquus, and T. sinensis. The late stage (1.2e0.8 Ma) is represented by the Queque fauna, which exhibits a distinct decrease in Neogene relic taxa and the appearance of several typical middle Pleistocene taxa, such as A. baconi, S. orientalis and B. brevicornis. The early Pleistocene Gigantopithecus faunas are also characterized by a gradual increase in dental size of some typical faunal members, such as Gigantopithecus, Ailuropoda and Tapirus, and the successive replacement of representative species, such as A. microta e A. wulingshanensis e A. baconi. The geographic distribution of G. blacki in the early Pleistocene is limited to the Oriental realm of southern China (south of the Yangtze River). Eleven G. blacki fossil sites, representing different periods of the early Pleistocene, have been recorded in southern China. The following chronological sequence is proposed for these sites (from oldest to youngest): Baikong, Longgupo, Liucheng, Baeryan, Chuifeng, Mohui and Juyuan (early Early Pleistocene), Longgudong and Sanhe (middle Early Pleistocene), and Queque (late Early Pleistocene). Early Pleistocene Gigantopithecus sites occur predominantly in Guangxi ZAR in southern China, accounting for more than 70% of all known G. blacki caves from this time period. This may imply that Guangxi ZAR represents a possible center of evolution and dispersal for G. blacki. Preliminary analysis of the dental metrics of G. blacki indicates that the teeth (especially the molars) increase in average size during the early Pleistocene from the early stage to the late stage. However, the significance of this observed tendency will require further study and the recovery of additional material of Gigantopithecus. 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
Please cite this article in press as: Jin, C., et al., Chronological sequence of the early Pleistocene Gigantopithecus faunas from cave sites in the Chongzuo, Zuojiang River area, South China, Quaternary International (2014), http://dx.doi.org/10.1016/j.quaint.2013.12.051
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Please cite this article in press as: Jin, C., et al., Chronological sequence of the early Pleistocene Gigantopithecus faunas from cave sites in the Chongzuo, Zuojiang River area, South China, Quaternary International (2014), http://dx.doi.org/10.1016/j.quaint.2013.12.051