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Two New Pliocene Hamsters (Cricetidae, Rodentia) from Southwestern Tibet (China), and Their Implications for Rodent Dispersal ‘Into Tibet’

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Two New Pliocene Hamsters (Cricetidae, Rodentia) from Southwestern Tibet (China), and Their Implications for Rodent Dispersal ‘Into Tibet’ Journal of Vertebrate Paleontology ISSN: 0272-4634 (Print) 1937-2809 (Online) Journal homepage: http://www.tandfonline.com/loi/ujvp20 Two new Pliocene hamsters (Cricetidae, Rodentia) from southwestern Tibet (China), and their implications for rodent dispersal ‘into Tibet’ Qiang Li, Thomas A. Stidham, Xijun Ni & Lüzhou Li To cite this article: Qiang Li, Thomas A. Stidham, Xijun Ni & Lüzhou Li (2017) Two new Pliocene hamsters (Cricetidae, Rodentia) from southwestern Tibet (China), and their implications for rodent dispersal ‘into Tibet’, Journal of Vertebrate Paleontology, 37:6, e1403443, DOI: 10.1080/02724634.2017.1403443 To link to this article: https://doi.org/10.1080/02724634.2017.1403443 View supplementary material Published online: 16 Feb 2018. Submit your article to this journal Article views: 30 View related articles View Crossmark data Full Terms & Conditions of access and use can be found at http://www.tandfonline.com/action/journalInformation?journalCode=ujvp20 Journal of Vertebrate Paleontology e1403443 (10 pages) Ó by the Society of Vertebrate Paleontology DOI: 10.1080/02724634.2017.1403443 ARTICLE TWO NEW PLIOCENE HAMSTERS (CRICETIDAE, RODENTIA) FROM SOUTHWESTERN TIBET (CHINA), AND THEIR IMPLICATIONS FOR RODENT DISPERSAL ‘INTO TIBET’ QIANG LI, *,1,2,3 THOMAS A. STIDHAM,1,3 XIJUN NI, 1,2,3 and LUZHOU€ LI 1,3 1Key Laboratory of Vertebrate Evolution and Human Origins of Chinese Academy of Sciences, Institute of Vertebrate Paleontology and Paleoanthropology, Chinese Academy of Sciences, Beijing 100044, China, [email protected]; 2Chinese Academy of Sciences Center for Excellence in Tibetan Plateau Earth Sciences, Beijing 100101, China, [email protected]; 3University of Chinese Academy of Sciences, Beijing 100049, China, [email protected]; [email protected] ABSTRACT—Two new species of fossil hamsters (Cricetinae, Cricetidae) collected from early Pliocene sediments (»4.4 Ma) in the Zanda Basin, southwestern Tibet (China), demonstrate greater past diversity among cricetines in the hinterland of the Tibetan Plateau within the Himalayan Range (beyond the previously known ‘Plesiodipus’ thibetensis from the late Miocene of the Gyirong Basin). The occurrence of Nannocricetus qiui, sp. nov., in the Zanda Basin indicates a dispersal of Nannocricetus from its center of origin in northern and northwestern China and the Mongolian Plateau, into the hinterland of the high-elevation Tibetan Plateau and subsequently into the Himalayan Range. The new taxon Aepyocricetus liuae, gen. et sp. nov., possibly represents a specialized (and endemic) Neogene hamster from the Tibetan Plateau. The dispersal of these hamsters into the high-elevation portions of Tibet during the early Pliocene contrasts with the hypothesized biogeographic shift of several large mammal lineages out of Tibet. The absence of Aepyocricetus and Nannocricetus from adjacent portions of the south slope of the Himalayans (and the Siwalik Hills in India and Pakistan) further implies that the Himalayan range functioned as a dispersal barrier for these small mammals by the early Pliocene. http://zoobank.org/urn:lsid:zoobank.org:pub:9EE286AA-0C00-4041-8F0C-877B35283181. SUPPLEMENTAL DATA—Supplemental materials are available for this article for free at www.tandfonline.com/UJVP Citation for this article: Li, Q., T. A. Stidham, X. Ni, and L. Li. 2018. Two new Pliocene hamsters (Cricetidae, Rodentia) from southwestern Tibet (China), and their implications for rodent dispersal ‘into Tibet’. Journal of Vertebrate Paleontology. DOI: 10.1080/02724634.2017.1403443. INTRODUCTION Cricetine fossils are distributed widely in the low-elevation regions of northern and southern China during the late Neogene, Hamsters are a diverse group of small rodents and comprise but they are extremely rare in the high-altitude Tibetan (Qing- seven extant genera and 18 species placed in the subfamily Crice- hai-Xizang) Plateau. Currently, the only known record of fossil tinae within the family Cricetidae (superfamily Muriodea). They cricetines in the hinterland of the Tibetan Plateau are specimens are allied with Arvicolinae, Lophiomyinae, Neotominae, Sigmo- referred to the late Miocene ‘Plesiodipus’ thibetensis (including dontinae, and Tylomyinae (Musser and Carleton, 2005). Living ‘Himalayactaga liui’) from the Gyirong ( D Guizhong or Chi- hamsters form a monophyletic group, subfamily Cricetinae, orig- long) Basin (Li and Chi, 1981). Even to this day, Cricetulus is the inating in the Neogene, whose monophyly is supported by both sole extant genus of hamsters living on the Tibetan Plateau (Luo morphological traits and phylogenetic analysis of nuclear gene et al., 2000). Fossils of two new extinct cricetine species from Pli- sequences (Carleton and Musser, 1984; Michaux and Catzeflis, ocene sediments in the Zanda ( D Zhada) Basin, southwestern 2000; Michaux et al., 2001). If the fossil record and extinct taxa Tibetan Plateau (China), expand the known past diversity of the are included, Cricetinae comprises at least 22 genera, and all of group and demonstrate the longevity of the group in the region. its members were once considered as part of family Muridae The Zanda Basin is in Ngari District in southwestern Xizang (McKenna and Bell, 1997). Hamsters are particularly diverse in (Tibet) Autonomous Region, China. It is a late Neogene pull- China, with a fossil record extending into the late Miocene and apart sag basin located just north of the high Himalaya ridge including 10 genera (Sinocricetus, Nannocricetus, Kowalskia crest, and has a mean altitude of about 4000 m (Fig. S1, Supple- [including Neocricetodon and Chuanocricetus], Amblycricetus, mentary Data 1). Tectonically, it is mainly controlled by the Cricetinus, Allocricetus, Bahomys, Cricetus, Cricetulus [including Oligo–Miocene Great Counter Thrust (GCT) to the north and Tscherskia and Allocricetulus], and Phodopus) (Zdansky, 1928; the South Tibetan Detachment System (STDS) to the south Schaub, 1930, 1934; Chow and Li, 1965; Zheng, 1984, 1993; Wu, (Murphy and Yin, 2003; Wang et al., 2008b). The Zanda Basin 1991; Zheng and Cai, 1991; Qiu, 1995; Luo et al., 2000; Qiu and has a vast current outcrop extent of basin fill (>9000 km2), with Storch, 2000; Qiu and Li, 2003; Zhang et al., 2008; Li, 2010). about 800 m of maximum thickness (Wang et al., 2008a; Saylor et al., 2010). These sediments have yielded surprisingly abundant and diverse mammalian fossils (Deng et al., 2011, 2012; Wang *Corresponding author. et al., 2013a, 2013b, 2014, 2015a, 2015b, 2016; Tseng et al., Color versions of one or more of the figures in the article can be found 2013a, 2013b, 2016; Li and Wang, 2015). The discoveries of the online at www.tandfonline.com/ujvp. earliest woolly rhinoceros (Coelodonta thibetana) initiated the Published online 16 Feb 2018 Li et al.—Pliocene hamsters from Tibet (e1403443-2) ‘out of Tibet’ hypothesis that proposes that earlier and more morphology on a Zeiss MA EVO25 scanning electron micro- primitive large mammals from the Pliocene of Tibet gave rise to scope without gold-plating and with 3.0 kV voltage. The teeth part of the Ice Age megafauna that extended across much of were measured with an Olympus SZX7 microscope to the near- what has been called the Mammoth Steppe (Deng et al., 2011; est 0.01 mm. Wang et al., 2015b). In contrast, the small mammals seem to have followed an ‘into Tibet’ pattern. This biogeographic pattern SYSTEMATIC PALEONTOLOGY is the case for zokors (family Spalacidae) and hamsters. Zokor dispersal into Tibet is recorded in Kunlun Pass and Zanda Basins Class MAMMALIA Linnaeus, 1758 in the form of Prosiphneus (Li and Wang, 2015). The hamster Order RODENTIA Bowdich, 1821 fossils discussed below were collected from Institute of Verte- Family CRICETIDAE Fischer von Waldheim, 1817 brate Paleontology and Paleoanthropology (IVPP) field locality Subfamily CRICETINAE Fischer von Waldheim, 1817 0 0 ZD1001 (31 40 N, 79 45 E, »4114 m above sea level; Fig. S1), NANNOCRICETUS Schaub, 1934 where the hamster material is associated with specimens of ancient snow leopard (Panthera blytheae), arctic fox (Vulpes Type Species—Nannocricetus mongolicus Schaub, 1934. zhudingi), and zokor (Prosiphneus eriksoni) (Tseng et al., 2013b; Referred Species—N. primitivus Zhang, Zheng, and Liu, 2008; Wang et al., 2014; Li and Wang, 2015). Locality ZD1001 bearing N. qiui, sp. nov. (this paper). fossil hamsters is correlated to the geomagnetic polarity time Distribution—Early late Miocene to late Pliocene in northern, scale (GPTS) magnetochron C3n.1 r, with an age of »4.40 Ma northwestern, and southwestern China; Late Miocene, (Wang et al., 2013b). Mongolia. Institutional Abbreviations—CAS, Chinese Academy of Sci- ences, Beijing, China; IVPP, Institute of Vertebrate Paleontol- NANNOCRICETUS QIUI, sp. nov. ogy and Paleoanthropology of CAS, Beijing, China; IZ, Institute of Zoology of CAS, Beijing, China; NGS, National Geographic Nannocricetus sp.: Deng et al., 2011:table S1. Society, Washington, D.C., U.S.A.; NSFC, National Nature and Nannocricetus sp.: Wang et al., 2013a:282. Science Foundation of China, Beijing, China; V, prefix to verte- Nannocricetus sp.: Wang et al., 2013b:93, table 3. brate fossils of IVPP; ZD, Zanda Basin, prefix to IVPP field numbers Holotype—IVPP V 23220, a left M1 (Fig. 1A). Paratypes—IVPP V 23221.1–16, 16 isolated molars including four left M1s (one with only the posterior portion preserved), MATERIALS AND METHODS two M2s (one left and one right), one left M3, four m1s (one left, The new fossil material includes 43 isolated molars
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