Molecular phylogeny and morphological revision of Myotis bats (Chiroptera: Vespertilionidae) from Taiwan and adjacent China Author Ruedi, Manuel Author Csorba, Gábor Author Lin, Liang- Kong Author Chou, Cheng-Han text Zootaxa 2015 3920 1 301 342 journal article 10.11646/zootaxa.3920.2.6 b05da2f8-5e97-4918-97e2-ae85ac43eecf 1175-5326 287922 8B991675-0C48-40D4-87D2-DACA524D17C2 Myotis laniger ( Peters, 1870 ) Synonymy. Vespertilio laniger Peters, 1870 . Type locality Amoy, Fujian, China . Vespertilio fimbriatus : Dobson 1878 (part). Name combination. Myotis sowerbyi : Allen 1938 (part) Name combination. Myotis daubentonii laniger : Tate, 1941 . First use of current name combination. Myotis daubentonii : Bates and Harrison 1997 (part). Name combination. Myotis laniger : Topál 1997 . Name combination. Myotis sp.: Lin et al. 1997 . Myotis daubentonii laniger : Bates et al . 1999 . Name combination. Myotis sp. 1: Lin et al. 2004 . Vernacular, unavailable name. Myotis cf. daubentonii : Borisenko et al 2008 . Name combination. Myotis sp.: Cheng et al. 2010 . Vernacular, unavailable name. Myotis sp. 1: Ruedi et al. 2013 . Vernacular, unavailable name. Taxonomic remarks. Peters’ description of M. laniger in Swinhoe’s book (1870) is relatively brief, but some major distinguishing characters for this species are mentioned (i.e., relatively long feet reaching more than half tibia length, wing membranes attached to mid-metatarsus, weak canines). Allen (1938) , in his influential book, unfortunately largely based his morphological description of M. laniger on the type series of another species, M. sowerbyi , which triggered much subsequent confusion about the exact characteristics of Peters’ laniger . Shamel (1942) reexamined M. sowerbyi and assigned it to a different group (i.e., as a subspecies of M. siligorensis ), which is its current taxonomic position today ( Corbet & Hill 1992 ; Simmons 2005 ). Unfortunately, the confusion about the exact content of M. laniger persisted. For instance following Tate (1941) , Bates and Harrison (1997) relied on Allen’s book (1938) to detail its morphology, and concluded that “ laniger ” should be regarded as a synonym of M. daubentonii (now confined to the western Palaearctic region and unrelated to M. laniger , see Clade III in figure 3). These authors overlooked that the attachment point of the wing membrane in true laniger is close to the ankle, not the outer metatarsal, and that it has long ears (versus short ones in M. daubentonii ). Topal’s (1997) concept of M. laniger was more accurate as he relied on the true laniger series from Amoy, and indeed stressed their morphological similarities with M. longipes and M. csorbai , both of them faunal elements of the Himalayan foothills. The apparent high morphological similarities between laniger , annamiticus and longipes ( Topál 1997 ; Kruskop & Tsytsulina 2001 ) also adds to this confusion, notably in the recent phylogenetic literature (e.g., Zhang et al. 2009 ), and more specific studies including material from the type localities are needed to fully understand the species limits in this complex group. It is, however, obvious from phylogenetic reconstructions that neither M. daubentonii (part of Clade III), nor M. capaccinii (outside Clade IX) pertain to this group of long-footed, Oriental species ( Fig. 3 ). After comparing the lectotype of M . laniger (and Peters’ 1870 original description) with the unknown Myotis discovered in the mountains of Taiwan and designated as Myotis sp. 1 ( Lin et al. 1997 ; Chou 2004 ; Lin et al. 2004 ), they proved to be undistinguishable. The characteristically weak dentition of the partial skull of the lectotype ( Fig. 5 a) also conforms to the dentition of specimens from Taiwan (Figs 4c, 5b). Genetically, all specimens from Taiwan are closely related to M. laniger sequenced in adjacent mainland China ( Fig. 3 ), which confirms their conspecificity. Distribution. The distribution of M. laniger as understood here is limited to the Fujian ( type locality) and Henan provinces in China , and Taiwan , but if further morphologic and genetic studies confirm that specimens from Southeast Asia also pertain to this species ( Topál 1997 ), then the distribution of M. laniger might be much more extensive. Measurements. See Table 4 . External morphology. This medium-sized bat has a dense, woolly fur, extending to the face, which is also distinctly hairy ( Cheng et al. 2010 ). The strong pilosity of this bat is also evident on the under parts of the wing and tail membranes, where relatively dense creamy-white hairs run along the sides of the body and extend on the membranes up to the elbows, knees, and base of tail; in this respect it resembles a small version of M. fimbriatus . The general color is greyish-brown above, with darker underfur. It is distinctly whitish on the ventral parts (pure white near the anal region). As mentioned by Peters (1870) , individuals can rarely be completely rufescent both above and below; this peculiar color morph has also been observed in Taiwan and is illustrated in Cheng et al. (2010) . The ears are long and narrow, relatively pointed, with an inconspicuous notch along the rear edge; they extend much beyond the nose tip if laid forward. The tragus is relatively straight and narrow, and reaches nearly half the conch height. The hairy face is dark brown, but the bare parts around the eyes are lighter, fleshy colored. The wing membrane is attached close to the ankle, but is prolonged by a narrow strip of membrane to the base of the metatarsus (see illustration in Cheng et al. 2010 ) and hence may give the impression of a more distal attachment. The calcar is long, about three-quarters the length of the free edge of uropatagium, without lobe or keel. The last vertebra is not inserted within the tail membrane. The feet are large ( 10.5 mm including claw) and more than half the tibia length, bear long, curved claws and are hairy. These characteristics of the feet are distinct from the smaller-footed species found in continental China (e.g., M. sowerbyi ) or in Taiwan (e.g., the unnamed Myotis sp. 2, see below). The penis is club-shaped. Skull morphology. The profile of the skull of M. laniger typically rises sharply after the postorbital constriction, but is nearly horizontal above, and rounded in the occipital region (Figs. 4c, 5). The surface of the braincase is smooth, with no visible crests. The teeth are typically weak, with the upper canines barely reaching the size of the third premolars; the lower ones are even smaller (Fig. 4c). The incisive and first upper premolars are nearly the same size, the second premolars being smaller. These premolars are uncrowded and aligned in the toothrow. The inferior teeth are weak, all molars being myotodont. Natural history. Allen (1923) commented that this bat was relatively uncommon in Fujian (compared to M. fimbriatus ), where he procured only three specimens. In Taiwan , it looks also to be a rare, cave dwelling bat, confined to the east of Taiwan . As in mainland China , it was also found in the same cave roosts as the larger M. fimbriatus . The large, hairy feet and uropatagium of M. laniger suggest that it is also a trawling species hunting close to or above water bodies.