taxonID	type	description	language	source
03D787D05637FF9BFF460B01D4E2174F.taxon	description	* Corresponding author. Email: yangqs @ ioz. ac. cn [Ve r s i o n o f R e c o r d, p u b l i s h e d o n l i n e 9 Ju l y 2 0 1 9; http: // zoobank. org / urn: lsid: zoobank. org: pub: 0122 DEEF- 3 F 68 - 4 D 2 F-A 119 - 378 D 8 C 4 CA 5 CF]	en	Ge, Deyan, Feijó, Anderson, Cheng, Jilong, Lu, Liang, Liu, Rongrong, Abramov, Alexei V., Xia, Lin, Wen, Zhixin, Zhang, Weiyong, Shi, Lei, Yang, Qisen (2019): Evolutionary history of field mice (Murinae: Apodemus), with emphasis on morphological variation among species in China and description of a new species. Zoological Journal of the Linnean Society 187: 518-534
03D787D05637FF9AFCDE08FED1AF176C.taxon	description	on the Danish islands Lolland and Faster (Andersen et al., 2017), which is not typical for mice of the genus Apodemus. More than 100 species or subspecies of this genus have been established in the historical literature (Thomas, 1922; Allen, 1938; Ellerman & Morrison-Scott, 1951; Xia, 1984; Musser et al., 1996; Musser & Carleton, 2005). Species of Apodemus are most abundant in the broadleaf forests of Palaearctic and Oriental areas. Several of these species host multiple human pathogens (Klein et al., 2015; Ma et al., 2015), notably hantavirus (Guzzetta et al., 2017; Tian et al., 2017) and Ljungan virus (Hauffe et al., 2010). A study on the evolutionary history and dispersal of this genus is, therefore, important for public health and biodiversity conservation in Eurasia and North Africa. Rapid radiation of Apodemus during the Late Miocene is thought to have been associated with global forest changes, when the flora changed from tropical to temperate during the Late Miocene (Serizawa et al., 2000). The earliest fossil records of this genus and its sister genera were dated to the early Late Miocene, approximately 11 Mya (Freudenthal, 1976; Martin Suarez & Mein, 1998; Kimura et al., 2017), but based on molecular data, the earliest divergence time of this genus was estimated to be approximately 6 – 8 Mya (Michaux et al., 2002; Liu et al., 2004; Suzuki et al., 2008; Darvish et al., 2015). Three crucial evolutionary events of this genus were recognized in previous studies: (1) initial broad dispersal and radiation approximately 6 Mya, (2) regional radiation in Europe and China approximately 2 Mya and (3) westward dispersal of A. agrarius to Europe in the Quaternary (Suzuki et al., 2008). These conclusions were proposed without evidence from fossil information or robust statistical analyses. A comprehensive study including extant and fossil species is important for understanding the evolutionary history of this genus. According to the most comprehensive mammalian checklists, the composition of Apodemus is still disputable. Mammal species of the world and the Handbook of the mammals of the world listed 20 species of Apodemus (Musser & Carleton, 2005; Wilson et al., 2016). Traditional taxonomy divided Apodemus into three groups: the Apodemus group, the Sylvaemus group and the Alsomys group (Zimmermann, 1962) or the Argenteus group (Musser et al., 1996). Studies based on complete mitochondrial cytochrome b (Cytb) sorted Apodemus species into four groups or subgenera: the Sylvaemus group [A. uralensis (Pallas, 1811), A. flavicolllis (Melchior, 1834), A. alpicola Heinrich, 1952, A. sylvaticus, A. mystacinus (Danford & Alston, 1877), A. hermonensis (Filippucci et al., 1989)], the Apodemus group [(A. agrarius, A. chevrieri (Milne-Edwards, 1868), A. speciosus (Temminck, 1844), A. draco (Barrett-Hamilton, 1900), A. ilex Thomas, 1922, A. semotus Thomas, 1908, A. latronum Thomas, 1911 and A. peninsulae (Thomas, 1907)] and A. argenteus (Temminck, 1844) and A. gurkha Thomas, 1924, constitute two other distinct groups (Serizawa et al., 2000; Liu et al., 2004; Suzuki et al., 2008). Generally, different aspects of species in Europe and eastern Asia have been well studied. For example, genetic variation among species using protein electrophoresis suggested a recent separation of members of the subgenus Sylvaemus from a common ancestor, followed by rapid radiation (Filippucci et al., 2002), and higher genetic variability of field mice was found in the Mediterranean peninsulas than in northern Europe (Michaux et al., 2003). Moreover, a large number of fossil species from Europe were described that were absent from glacial assemblages, but are always present in interglacial assemblages (Martin Suarez & Mein, 1998; Knitlova & Horacek, 2017 a, b). However, it is unclear how the evolutionary history of Apodemus formed its current distributional pattern. Faunas in many regions of China remain relatively poorly explored, impeding a comprehensive understanding of interspecific differentiation. Allen (1938) recognized five species of Apodemus in China and Xia (1984) listed six: A. sylvaticus, A. draco, A. peninsulae, A. latronum, A. chevrieri and A. agrarius. Zheng (1993) reported fossil occurrences of A. chevrieri, A. agrarius, A. sylvaticus and A. cf. peninsulae that were dated to the Late Pleistocene in Chongqing and Guizhou Provinces. Based on a large number of Apodemus specimens from China, Musser et al. (1996) recognized seven species, A. agrarius, A. chevrieri, A. peninsulae, A. latronum, A. draco, A. semotus and A. uralensis, but excluded A. sylvaticus. Smith & Xie (2008) included one more species, A. pallipes (Barrett-Hamilton, 1900), than the study of Musser et al. (1996), but this number was not followed by Wilson et al. (2016), detailed information are given in Table 1. In recent studies, A. ilex is also considered a distinct species (Liu et al., 2004, 2012, 2017). A previous study based on a wider range of sampling, and using cytochrome oxidase subunit I (Cox 1) recognized A. agrarius, A. chevrieri, A. peninsulae, A. latronum, A. ilex, A. draco and A. uralensis, and identified a distinct genetic lineage from Guizhou Province (Liu et al., 2017). However, it is unclear how to identify these species based on morphology and a detailed description of the distinct genetic lineage is lacking. An integrative study on the phylogeny of Apodemus and taxonomy of species in this genus in China is, therefore, warranted. In recent years, we collected a large number of samples of the genus Apodemus in China and examined collections in several museums that preserve Chinese specimens. Here, our aim is to (1) investigate the evolutionary history of this taxon by integrating fossil occurrences and extant species, (2) use the broadest geographic coverage of Apodemus to date, to assess the phylogenetic relationships and morphological variation of all species of this genus recorded in China and (3) test whether the newly discovered genetic lineage in our previous study represents a distinct species. By integrating molecular and morphological data, we established a new species of the genus Apodemus from high mountains in south-eastern China (27.83 ºN, 108.76 ºE).	en	Ge, Deyan, Feijó, Anderson, Cheng, Jilong, Lu, Liang, Liu, Rongrong, Abramov, Alexei V., Xia, Lin, Wen, Zhixin, Zhang, Weiyong, Shi, Lei, Yang, Qisen (2019): Evolutionary history of field mice (Murinae: Apodemus), with emphasis on morphological variation among species in China and description of a new species. Zoological Journal of the Linnean Society 187: 518-534
03D787D05630FF97FCD60B69D61C1406.taxon	description	The landscape of China is a complex mosaic, characterized by high-plateau mountain steppes in the west, temperate dry steppes and deserts in the north, a complex of mountain ranges harbouring dense rainforest in the centre and subtropical, low-elevation forests on the east coast, the so-called three levels of terrain in China. Such habitat heterogeneity appears to favour the diversification of Apodemus species. The large genetic distances and significant morphological variations among these species probably resulted from a long history of differentiation. For example, the split of A. draco and A. ilex, the youngest sister species of this genus distributed in China, was dated to approximately 3.26 Mya. Based on the locations of molecular voucher specimens, A. agrarius is widely distributed in the north to the south-east of China, A. peninsulae occurs along the ‘ Hu Huanyong Line’ (Hu, 1935) or ‘ the 400 mm annual precipitation line’ from north-east to south-west China, A. uralensis inhabits the north-west Qinghai – Tibet plateau and Xinjiang, and A. semotus is endemic to Taiwan (Liu et al., 2017). The remaining species, A. draco, A. chevrieri, A. latronum, A. ilex and A. nigrus are endemic to the mountainous region of southern China. This region harbours a high diversity of species and reflects a marked heterogeneous habitat zonation along mountain slopes, in some cases mirroring tropical – temperate forest transitions. Several new mammal species were described from this zone in recent years (e. g. Fan et al., 2017; Ge et al., 2018 b), suggesting a still overlooked biodiversity in China. The discovery of the new species A. nigrus from the Wuling Mountain chain in Guizhou and Congqing Provinces reinforces this trend. The Wuling Mountains are located in central China, which is a transitional zone connecting the lowest and highest of the three levels of terrain in China, with a mean elevation of approximately 1000 m. This region constitutes one of the major components of the Jiangnan orogenic belt in southern China. The climate of the Pliocene here was characterized by a warm period from the Early to the Middle Pliocene (2 – 3 ° C higher than today) and a transition from relatively warm climates to the prevailing cooler climates of the Pleistocene. The greatest warming appears to have occurred at higher latitudes, where temperatures were often sufficiently elevated to allow species of animals and plants to exist at higher latitudes than their closest modern relatives. Historical exploration rarely paid attention to the Wuling Mountains, which is evident today by sparse material for different taxa from this region in museums. The new species A. nigrus is found on the Fanjing and Jinfo Mountains, both belonging to the Wuling Mountain chain. Fanjing Mountain (or Fanjingshan) is the highest mountain in the Wuling Mountains. The peak of this mountain is 2570 m above sea level. The mountain acts as an ‘ isolated ecological island’ with a high degree of endemism of plants and animals. Fanjing Mountain is unique in its geological history, landforms, geographical location and climatic conditions, and all of these factors have created a terrestrial island with a specific ecological environment. The main peak of Fanjing Mountain is separated from the surrounding karst areas by its high elevation, and it was selected as a World Heritage Site in 2018. It is an ecological island that hosts the endangered Guizhou golden monkey (Rhinopithecus brelichi Thomas, 1903), the Chinese dove tree (Davidia involucrata Baill.), various alpine rhododendrons (Rhododendron spp.) and many other rare animals and plants. With an elevation of 2238 m, Jinfo Mountain is the highest mountain in Chongqing Province, and it is famous for its high biodiversity. The split of A. nigrus from its sister taxa likely occurred during the Middle Pliocene, approximately 4 Mya. Climate warming drove its preferred habitat upwards, which caused this species to inhabit higher elevations. However, this region was likely less affected by climatic oscillations in the Late Quaternary than other regions of the world, for example, the stable demographic history of Eothenomys melanogaster (Milne-Edwards, 1871) and the survival of five Chinese giant salamander species that diverged over four million years ago in the same region (Lv et al., 2018; Yan et al., 2018). Subsequent global cooling allowed this species to migrate slightly to lower elevations, but the dependence of the species on local vegetation and climate warming at lower latitudes likely impeded widespread dispersal. Abbreviations for species are the same as Table 2. ‘ N ’ gives the total number of intact adult specimens included in analyses. BM = body mass, EM = external measurements, HBL = head and body length, TL = tail length, HFL = hind foot length, EL = ear length, CM = craniodental measurements, TLC = total length of the cranium, NL = nasal length, GWS = greatest width of the ‘ snout’, SDO = shortest distance between orbits, ZB = zygomatic breadth, GMB = greatest mastoid breadth, BL = basal length, BSL = basilar length, PL = palatal length, IFL = incisive foramen length, WIF = width of the incisive foramen, GPB = greatest palatal breadth, LTB = length of the tympanic bulla, ULMM = length of the maxillary molars, ULMD = length of the maxillary diastema, ML = mandibular length, LLMM = length of the mandibular molars, LLMD = length of the mandibular diastema. TAXONOMIC ACCOUNT Order: Rodentia Family: Muridae Subfamily: Murinae Apodemus Kaup, 1829. APODEMUS NIGRUS DEYAN GE, ANDERSON FEIJÓ &	en	Ge, Deyan, Feijó, Anderson, Cheng, Jilong, Lu, Liang, Liu, Rongrong, Abramov, Alexei V., Xia, Lin, Wen, Zhixin, Zhang, Weiyong, Shi, Lei, Yang, Qisen (2019): Evolutionary history of field mice (Murinae: Apodemus), with emphasis on morphological variation among species in China and description of a new species. Zoological Journal of the Linnean Society 187: 518-534
