Elsevier

Quaternary International

Volume 445, 25 July 2017, Pages 250-268
Quaternary International

New fossils of Bison palaeosinensis (Artiodactyla, Mammalia) from the steppe mammoth site of Early Pleistocene in Nihewan Basin, China

https://doi.org/10.1016/j.quaint.2016.07.033 Get rights and content

Abstract

Since the establishment of the species Bison palaeosinensis by Teilhard de Chardin and Piveteau in 1930, no new fossils have been reported in the last few decades. Recent excavations at the Shanshenmiaozui site in Nihewan Basin of North China led to the discovery of numerous Bovini fossils, including a nearly complete mandible and articulated postcranial bones. The stout axis and the protruded mandibular angle, thick posterior rim of the mandibular ramus, prominently curved coronoid process, thickened mandibular symphysis and short dentition support an assignment to the Bison genus. Moreover, the new specimens have smaller and proportionally narrower cheek teeth as well as a higher premolar to molar ratio than those found in later forms of bison, further supporting their assignment to the species Bison palaeosinensis. The postcranial bones are also considerably smaller and more slender than those of the later forms of bison. In North China, isolated teeth and postcranial bones have contributed to confusion regarding the distinction between early bison and Leptobos, because these specimens have similar dimensions and were temporally concurrent even at the same site. In addition, sexual dimorphism is another potential factor making identification work difficult. In Nihewan Basin, the joint occurrence of Bison palaeosinensis with Mammuthus trogontherii and Coelodonta nihowanensis marks the appearance of the early stage of the Mammuthus-Coelodonta faunal complex.

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.

. Studied fossil specimens of

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;

References (83)

  • S.K. Vasiliev

    Late Pleistocene bison (Bison p. priscus Bojanis, 1827) from the Southeastern part of Western Siberia

    Archaeology Ethnology & Anthropology of Eurasia

    (2008)
  • Archaeological United Team

    Quaternary mammalian fauna from Jinniushan, Yingkou of Liaoning Province

    Vertebrata PalAsiatica

    (1976)
  • D.M. Balkwill et al.

    A guide to the identification of postcranial bones of Bos taurus and Bison bison

    Syllogeus

    (1992)
  • G.B. Barbour

    Preliminary observation in Kalgan area

    Bulletin of the Geological Society of China

    (1924)
  • G.B. Barbour

    The deposits of the Sang Kan Ho valley

    Bulletin of the Geological Society of China

    (1925)
  • G.B. Barbour et al.

    Geological study of the deposits of the San kan ho basin

    Bulletin of the Geological Society of China

    (1927)
  • C.L. Brown et al.

    A Key to Postcranial Skeletal Remains of Cattle/Bison, Elk, and Horse

    (1979)
  • M. Bukhsianidze

    The Fossil Bovidae of Dmanisi

    (2005)
  • G.F. Chen

    Artiodactyla

  • H.X. Chi

    The lower Pleistocene mammalian fossils of Lantian district, Shenxi

    Vertebrata PalAsiatica

    (1975)
  • L.P. Chia et al.

    Hsihoutu—A Culture Site of Early Pleistocene in Shansi Province

    (1978)
  • M.C. Chow et al.

    Notes on Villafranchian mammals of Linyi, Shansi

    Vertebrata PalAsiatica

    (1965)
  • C. De Giuli

    Late Villafranchian faunas of Italy: the Selvella local fauna in the southern Chiana valley (Umbria)

    Paleontographia Italica

    (1987)
  • F.A. Demirel et al.

    A new early Pleistocene mammalian fauna from Burdur basin, SW Turkey

    Russian Journal of Theriology

    (2014)
  • W. Dong et al.

    Artiodactyla

  • M. Drees

    Sexual dimorphism in Pleistocene Bison priscus (Mammalia, Bovidae) with a discussion on the position of Bison schoetensacki

    Senckenbergiana Lethae

    (2005)
  • M.-P. Duvernois

    Les Leptobos (Mammalia, Artiodactyla) du Villafranchien d'Europe Occidentale: Systématique, Évolution, Biostratigraphie, Paléoécologie

    Documents des Laboratoirres de Géologie Lyon

    (1990)
  • C.C. Flerov

    The earliest representatives and the history of genus Bison

  • C.C. Flerov

    Systematics and evolution

  • W.A. Fuller

    The horns and teeth as indicators of age in Bison

    The Journal of Wildlife Management

    (1959)
  • H. Gee

    The distinction between postcranial bones of Bos primigenius Bojanus, 1827 and Bison priscus Bojanus, 1827 from the British Pleistocene and the taxonomic status of Bos and Bison

    Journal of Quaternary Science

    (1993)
  • A.W. Gentry

    The subfamilies and tribes of the family Bovidae

    Mammal Review

    (1992)
  • A.W. Gentry

    Chapter 38: Bovidae

  • A.W. Gentry et al.

    Suborder Ruminantia

  • D. Geraads

    Phylogenetic analysis of the tribe bovini (Mammalia: Artiodactyla)

    Zoological Journal of the Linnean Society

    (1992)
  • C.P. Groves

    Systematic relationships in the bovini (Artiodactyla, Bovidae)

    Zeitschrift für Zoologische Systematik und Evolutionsforschung

    (1981)
  • H.H. Hsieh

    Mammalian fossils from the Pleistocene of Huituipo, Tungchuan, Shensi

    Vertebrata PalAsiatica

    (1964)
  • C.K. Hu et al.

    Gongwangling Pleistocene mammalian fauna of Lantian, Shaanxi. Palaeontologia Sinica

    New Series C

    (1978)
  • W.P. Huang et al.

    The quaternary mammalian fossil localities of Lantian District, Shensi

    Vertebrata PalAsiatica

    (1966)
  • W.P. Huang et al.

    A brief of quaternary mammalian fossils of the Sanmenxia district

    Vertebrata PalAsiatica

    (1984)
  • C.M. Janis et al.

    Cranial and postcranial morphological data in ruminant phylogenetics

    Zitteliana B

    (2014)
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