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Holocene Survival of Late Pleistocene Megafauna in China: a Critical Review of the Evidence

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Holocene Survival of Late Pleistocene Megafauna in China: a Critical Review of the Evidence Quaternary Science Reviews 76 (2013) 156e166 Contents lists available at SciVerse ScienceDirect Quaternary Science Reviews journal homepage: www.elsevier.com/locate/quascirev Holocene survival of Late Pleistocene megafauna in China: a critical review of the evidence Samuel T. Turvey a,*, Haowen Tong b, Anthony J. Stuart c, Adrian M. Lister d a Institute of Zoology, Zoological Society of London, Regent’s Park, London NW1 4RY, UK b Key Laboratory of Vertebrate Evolution and Human Origin of Chinese Academy of Sciences, Institute of Vertebrate Paleontology and Paleoanthropology, Chinese Academy of Sciences, Beijing 100044, China c School of Biological and Biomedical Sciences, Durham University, South Road, Durham DH1 3LE, UK d Department of Earth Sciences, Natural History Museum, London SW7 5BD, UK article info abstract Article history: Late Quaternary megafaunal extinction chronologies are poorly understood across eastern and south- Received 24 November 2012 east Asia. Previous radiometric studies suggested that surprisingly many extinct Late Pleistocene large Received in revised form mammal species survived into the Holocene in northern China (Bos primigenius, Coelodonta antiquitatis, 22 April 2013 Mammuthus primigenius) and southern China (Ailuropoda baconi, Crocuta [crocuta] ultima, Megatapirus Accepted 29 June 2013 augustus, Stegodon orientalis, Sus cf. xiaozhu), indicating that Chinese megafaunal extinctions may have Available online been “staggered” across the Late Quaternary. We critically re-examined all radiometric evidence sug- gesting Holocene survival of Chinese Late Quaternary megafauna, and conducted new dating of mammal Keywords: 14C Dates material from reportedly Holocene sites containing characteristically Late Pleistocene faunas. Evidence Extinct mammal for Holocene survival of any Chinese Late Pleistocene megafaunal species is weak or untenable. No Late Quaternary extinction previous radiometric dates used to support Holocene megafaunal survival represent direct bone dates for Mammoth species of interest, and stratigraphic association between material yielding Holocene dates and mega- Palaeoloxodon faunal remains is dubious at most sites. Concerns over accurate identification of faunal material further Radiocarbon dating confuse claims for Holocene survival of many species. Robust radiometric last-occurrence dates for Stegodon extinct Chinese megafauna are all restricted to the Late Pleistocene, similar to the timing of many other Late Quaternary megafaunal species extinctions elsewhere in Eurasia and the Americas. Ó 2013 Elsevier Ltd. All rights reserved. 1. Introduction poorly constrained elsewhere, notably across much of eastern and south-east Asia (Louys et al., 2007; Turvey, 2009). Clarifying the The disappearance of a large proportion of the world’s mega- magnitude, timing and causation of Late Quaternary extinctions in faunal mammal taxa during the Late Quaternary (two-thirds of all this region is of particular significance because, apart from Africa, it mammal genera, and half of all species, with body masses >44 kg) is the only part of the world with surviving representatives of is the subject of considerable scientific interest (Martin and Klein, “truly” megafaunal (sensu Owen-Smith, 1988, i.e. >1000 kg) pro- 1984; MacPhee, 1999; Barnosky et al., 2004). The great majority boscidean and rhinocerotid taxa (Dicerorhinus, Elephas, Rhinoceros). of research into Late Quaternary extinctions has focused on It is also recognized as having the world’s highest number of megafaunal species losses in the continental regions of North currently threatened large land mammal species, making attempts America, northern Eurasia, and Australia. Although many questions to understand the dynamics and drivers of past regional mammal remain, this research has led to an increasingly sophisticated un- extinction of considerable conservation relevance (Schipper et al., derstanding of the timing and dynamics of megafaunal extinction 2008). in response to both anthropogenic impacts and environmental China possesses the richest known Late Quaternary palae- change in each of these regions (e.g. Stuart et al., 2004; Turney et al., ontological and zooarchaeological record in the eastern/south-east 2008; Gill et al., 2009; Rule et al., 2012; Stuart and Lister, 2012). In Asian region, and spatial and temporal variation in the species contrast, megafaunal extinction chronologies remain far more composition of Chinese Late Quaternary mammal faunas has long been recognized. This vast country spans substantial portions of both the Palaearctic and Oriental zoogeographic realms. The * Corresponding author. Tel.: 44 207 449 6326; fax: 44 207 586 2870. northeastern Chinese Late Pleistocene mammal fauna is interpreted þ þ E-mail address: [email protected] (S.T. Turvey). as an extension of the northern Eurasian fauna that was also 0277-3791/$ e see front matter Ó 2013 Elsevier Ltd. All rights reserved. http://dx.doi.org/10.1016/j.quascirev.2013.06.030 S.T. Turvey et al. / Quaternary Science Reviews 76 (2013) 156e166 157 distributed in Europe, the Russian Federation, Kazakhstan, extirpations and global extinctions of Chinese large mammals, Mongolia and Japan, and contained several characteristic repre- including elephants (Xu, 2000; Elvin, 2004), ungulates (Yang et al., sentatives of this fauna such as woolly mammoth Mammuthus 2005; Wen, 2009), carnivores (Tilson et al., 2004; Yang et al., 2005; primigenius and woolly rhinoceros Coelodonta antiquitatis (Dong Jablonski et al., 2012), primates (Zhang et al., 1989; Li et al., 2002; et al., 1996; Tong and Patou-Mathis, 2003; Stuart and Lister, 2012), Grueter et al., 2009; Chatterjee et al., 2012), and endemic fresh- with mammoths occurring as far south as Shandong Province dur- water cetaceans (Turvey et al., 2007), associated with extreme ing Marine Oxygen Isotope Stage 3a (Takahashi et al., 2007). The levels of human overpopulation, natural resource overexploitation more warm-adapted straight-tusked elephant Palaeoloxodon nau- and habitat modification throughout the development of Chinese manni was also distributed in northeastern China during the Late civilization. Pleistocene, where it is known from more than 30 localities, often in Only one globally extinct and taxonomically valid Chinese apparent stratigraphic association with Coelodonta (Xue et al., 2000; mammal species, the short-horned water buffalo Bubalus mephis- Tong and Patou-Mathis, 2003). Other extinct megafaunal taxa topheles, is known with certainty to have disappeared during the associated with the Eurasian Palaeoloxodon assemblage (e.g. Bos Holocene before the recent historical era. Although no direct primigenius, Stephanorhinus, megacerine deer) are also known from radiometric dates are available, this species is present in a series of northeastern China during the Late Pleistocene (Dong et al., 1996; well-dated earlyemiddle Holocene (NeolithiceBronze Age) Pushkina, 2007; Tong and Wu, 2010). The subtropical area of zooarchaeological deposits across southern, central and eastern China and neighbouring south-east Asia south of the Yellow Rivere China (from Yunnan to northern Henan), in association with rep- Qinling Mountains, the traditional biogeographic boundary be- resentatives of the modern Holocene Chinese large mammal fauna tween the Palaearctic and Oriental Realms (Tong, 2007), was (Teilhard de Chardin and Young, 1936; Wei et al., 1990; Zhang et al., defined during much of the Pleistocene by the regionally distinctive 1992; Ji et al., 2004; Liu et al., 2004). Ancient DNA analysis demon- StegodoneAiluropoda fauna (Colbert and Hooijer, 1953; Kahlke, strates that it is phylogenetically distinct from domesticated water 1961; Wang and Ouyang, 1982; Yang et al., 1995; Long et al., 1996; buffalo Bubalus bubalis present in China today (Yang et al., 2008). Louys et al., 2007). This fauna was characterized by the extinct However, it has also been suggested that a surprisingly large proboscidean Stegodon orientalis and the “giant” giant panda number of globally extinct representatives of China’s Late Pleisto- Ailuropoda baconi, formerly considered to be a subspecies of the cene megafauna survived well beyond the PleistoceneeHolocene extant Ailuropoda melanoleuca but now often interpreted as a transition in both northern and southern China (Table 1; Figs. 1 distinct chronospecies (Jin et al., 2007). It also contained a relatively and 2). A series of radiometric dates was put forward by Ma and high diversity of anthropoid primates (including archaic hominins Tang (1992) and Tong and Liu (2004) as evidence for the earlye that may be specifically distinct from Homo sapiens; Curnoe et al., middle Holocene persistence of Bos primigenius, Coelodonta anti- 2012), as well as several other regionally endemic extinct large quitatis and Mammuthus primigenius in northern China, and Ailur- mammals such as Megatapirus augustus, Rhinoceros sinensis, Sus opoda baconi, Crocuta [crocuta] ultima, Megatapirus augustus, xiaozhu, and Crocuta [crocuta] ultima (which may also represent a Stegodon orientalis and Sus cf. xiaozhu in southern China. More distinct species; Rohland et al., 2005) and also extant taxa such as recently, Li et al. (2012) reassigned two allegedly mid-Holocene the Asian elephant Elephas maximus. elephant molars from northern China from Elephas maximus to The StegodoneAiluropoda fauna has been interpreted as a gen- Palaeoloxodon sp., and proposed that this extinct proboscidean also eral category of fossil assemblages rather than an
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