New fossils of Bison palaeosinensis (Artiodactyla, Mammalia) from the steppe mammoth site of Early Pleistocene in Nihewan Basin, China
Introduction
The Nihewan Fauna consists of numerous and diversified ungulates, among which the Bovini fossils are still tied to many open questions (Flerov, 1979, McDonald, 1981). In early studies, Bison palaeosinensis was the only species assigned to the tribe of Bovini. Although quite a number of postcranial bones of diversified bovids were described and measured by Teilhard de Chardin and Piveteau (1930), their classification is far from clear.
Recent excavations at the Shanshenmiaozui site has led to the discovery of more Bovini fossils, including a nearly complete mandible and rich postcranial bones, of which some are still articulated with one another. These fossils can be used to classify the large bovid fossils from Nihewan Basin in northern China. The Nihewan Basin is famous for its fluvio-lacustrine deposits containing rich mammalian fossils of Early Pleistocene age, which was first reported during 1924–1930 (Barbour, 1924, Barbour, 1925, Teilhard de Chardin, 1926, Barbour et al., 1927, Teilhard de Chardin and Piveteau, 1930). More recent study (Qiu, 2000) shows that the classical Nihewan Fauna can be correlated with the Olivola and Senèze faunas, two Late Villafranchian sites in Italy and France.
The SSMZ locality lies at the neighboring hill slope of Xiaochangliang, a well-known Paleolithic site in Nihewan Basin, Hebei Province (Fig. 1). Based on the stratigraphic correlation in the field, the fossil-bearing sand-silt bed at the new locality is a little higher than the cultural layer at Xiaochangliang site (Tong et al., 2011, Liu et al., 2016), whose paleomagnetic age is about 1.36 Ma BP (Zhu et al., 2001). The SSMZ site bears quite rich and well preserved fossils, such as Canis chihliensis (Tong et al., 2012), Coelodonta nihowanensis (Tong and Wang, 2014), Mammuthus trogontherii (Tong, 2012, Tong and Chen, 2016; Chen and Tong, under review) and B. palaeosinensis (this paper), among which the juvenile specimens of Coelodonta nihowanensis and Mammuthus trogontherii are very significant. The mammalian taxa associated with B. palaeosinensis at SSMZ are as follows: Lepus sp., Ochotona sp., Canis chihliensis, Pantherinae gen. et sp. indet., Pachycrocuta sp., Mammuthus trogontherii, Coelodonta nihowanensis, Elasmotherium peii, Proboscidipparion sp., Equus sanmeniensis, Sus sp., Eucladoceros sp., Spirocerus wongi and Gazella sinensis (Tong et al., 2011). During the 2015 field season, a metatarsal bone of Paracamelus was recovered.
Section snippets
Fossil material
The majority of the studied fossils of Bison palaeosinensis were mainly recovered recently from the SSMZ locality. Only one mandible (V 22669) and one foot bone (V 22670.1–7) were collected in the Cenjiawan area by local people and during the biostratigraphic survey. All of the studied specimens are listed in Table 1. In addition, one mandible of B. palaeosinensis (IVPP RV30036) studied previously by Teilhard de Chardin and Piveteau (1930) was reexamined.
Methods, terminology and abbreviations
The osteological terms follow Olsen, 1960, Reshetov and Sukhanov, 1979, Gee, 1993 and Brown and Gustafson (1979). The terms of tooth nomenclature follow Sala, 1986, Duvernois, 1990, Gentry, 1992, Gentry et al., 1999, Gentry, 2010 and Bärmann and Rössner (2011). The upper molars were denoted by uppercase letter “M”; the lower premolars and molars were denoted by lowercase letters “p” and “m” respectively. The specimens were measured according to the methods used by von den Driesch, 1976,
Classification
Class Mammalia Linnaeus, 1758
Order Artiodactyla Owen, 1848
Suborder Ruminantia Scopoli, 1777
Superfamily Bovoidea Gray, 1821
Family Bovidae Gray, 1821
Subfamily Bovinae Gray, 1821
Tribe Bovini Gray, 1821
Genus Bison C.H. Smith, in Griffith, Smith & Pidgeon, 1827
Subgenus Eobison Flerov, 1972
Bison (Eobison) palaeosinensis Teilhard de Chardin and Piveteau, 1930
1947. Bison (Subgenus?) palaeosinensis Teilhard de Chardin and Piveteau, 1930–Skinner and Kaisen, p. 218, text Fig. 2-A & B
1979. Bison (Eobison)
Taxonomic assignment
The validity and taxonomic position of the species B. palaeosinensis has been controversial since its establishment. Despite long debates in the last few decades, the taxonomic problem is still not resolved. However, it is becoming clear that the fossils of large bovini from the Nihewan beds definitely belong to Bison, rather than to Leptobos or another large bovini genus. The new fossils from this study can also be assigned to the species B. palaeosinensis due to its following characters:
Conclusion
The protruded mandibular angle, stout mandibular condyle, strongly bent coronoid process, thickened mandibular symphysis, short dentition and stout axis recently recovered from SSMZ site in Nihewan Basin support an assignment of Bison. Moreover, the new specimens are prominently smaller than later forms of bison, and they can be included in the species B. palaeosinensis, which has narrower upper and lower teeth. Furthermore, B. palaeosinensis also has the following dental characters: hypsodonty
Acknowledgements
The authors wish to express their thanks to the following people and organizations for their help: F. Han, Z. J. Xu, C. Yin, N. Hu and X. M. Wang for field work; Q. Wei for providing the site information; V. Titov and M. Bukhsianidze for providing bibliographies; B. Bai for providing photos of the specimen of Leptobos falconeri in the American Museum of Natural History; Y. Xu for finishing the line drawings in Fig. 7; G. D. Wang for arrangement of collection visit at Qinghai Provincial Museum;
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2019, Quaternary InternationalCitation Excerpt :In addition, both horn cores and dentitions are evidently smaller than those from Shuichongkou (Tables 2, 4). Compared with the left metatarsal (V22667) of Bison palaeosinensis from the Early Pleistocene deposits at SSMZ (Shanshenmiaozui) in Nihewan basin (Tong et al., 2017), the SSMZ specimen is evidently shorter but slightly thicker than the Shuichongkou specimen (Fig. 4, Table 3). In dorsal view, the vascular groove of SSMZ specimen is deeper than that of Shuichongkou specimen, the proximal nutrient foramen is present in SSMZ specimen but absent in Shuichongkou specimen.
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2018, Quaternary Science ReviewsCitation Excerpt :These sites include Xiaochangliang (Zhu et al., 2001), Majuangou and Banshan (Zhu et al., 2004), Donggutuo (Li and Wang, 1982; Wang et al., 2005), Maliang (Wang et al., 2005), Dachangliang (or Xiantai) (Deng et al., 2006a), Cenjiawan (Wang et al., 2006), Feiliang (Deng et al., 2007) and Huojiadi (Liu et al., 2010). The Nihewan Fauna (sensu lato) comprises a series of mammalian faunas from Nihewan Basin, such as the Donggutuo (Li and Wang, 1982; Wei, 1985, 1991; Wang et al., 2005), Maliang (Wei, 1991; Wang et al., 2005), Xiaochangliang (You et al., 1980; Tang et al., 1995; Zhu et al., 2001), Banshan (Wei, 1994; Zhu et al., 2004), Majuangou-I, Majuangou-II, Majuangou-III (Zhu et al., 2004), Hongya (Huang and Tang, 1974), Huabaogou-I (Wang, 1982), Huabaogou-II (Wang, 1982; Zhou et al., 1991), Yangshuizhan (Ao et al., 2013b; Liu et al., 2017), Shanshenmiaozui (Liu et al., 2016; Chen and Tong, 2017; Tong et al., 2017), Danangou (Li, 1984), Daodi-Dongyaozitou (Tang, 1980; Tang and Ji, 1983) and Xiashagou (Teilhard de Chardin and Piveteau, 1930; Liu et al., 2012, 2016) faunas, some of which have been well dated by comprehensive stratigraphic investigations. Nihewan Basin has been called “the Chinese Olduvai Gorge” due to its abundant Paleolithic artifacts (Yuan et al., 2009).
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Morphometric-palaeoecological discrimination between Bison populations of the western Palaearctic
2017, GeobiosCitation Excerpt :All the data were analysed with the software IBM SPSS (Statistical Package for Social Sciences) 20 and PAST (Hammer et al., 2001). Selected measured dimensions of metapodial and dental variables of Bison degiulii (from the Italian locality Pirro Nord, ca. 1.5 Ma; De Giuli et al., 1987), Leptobos etruscus (Falconer, 1859) and Bison palaeosinensis included in the analysis are taken from Masini (1989) and Tong et al. (2016), and those of Bison georgicus from Bukhsianidze (2005). Comparative material of the latest early Pleistocene (∼0.9 Ma) and early middle Pleistocene (∼0.7 Ma) Bison schoetensacki from Durfort (southern France) and West Runton Freshwater Bed in Norfolk, were taken from Brugal (1995) and Sala (1986), respectively.