taxonID	type	description	language	source
03BB87963D5D5D11A8EBF6CF11FCFC33.taxon	materials_examined	Taxonomic remarks. Kishida (1932) described a small species of Myotis from Taiwan and named it M. latirostris. Shortly after this description and also based on small specimens caught in Taiwan, Kuroda (1935) described another taxon, M. muricola orii. The very small forearm dimensions of both taxa (FA 33 – 34 mm), their tiny hind feet (6.5 mm), blackish fur and their small second upper premolars that are clearly visible laterally, are all morphological characters found in no other Myotis from Taiwan. Hence, despite the fact that the type specimens of latirostris and orii are not available for direct comparisons (Imaizumi 1962; Kaneko & Maeda 2002), there is no doubt that both taxa belong to the same species, but its systematic position is more contentious. Classical accounts variously treated latirostris (and orii) as a subspecies of M. mystacinus or of M. muricola, (Tate 1941; Findley 1972; Corbet & Hill 1992; Simmons 2005), but recent molecular reconstructions suggest that none of these species share a common ancestor (Stadelmann et al. 2007; Lack et al. 2010; Ruedi et al. 2013). The molecular data, whether issued from mitochondrial (Stadelmann et al. 2007; Fig. 3) or nuclear genes (Lack et al. 2010; Ruedi et al. 2013), rather show that the phylogenetic position of the latirostris lineage is very peculiar, being basal to all recent Myotis species tested so far. In addition to these molecular reconstructions, specimens of the latirostris lineage show that their first two lower molars have a nyctalodont configuration (i. e., where the postcristid is linking to the hypoconulid; Menu & Sigé 1971), whereas most species of Myotis have myotodont lower molars (i. e., where the postcristid is linking to the entoconid; Menu & Sigé 1971). The only exceptions are the small Myotis of the siligorensis-alticraniatus group (Tiunov et al. 2011; Ruedi et al. 2013), that can be easily recognized by their distinctly raised frontal part of the braincase and crowded premolars (Borisenko et al. 2008); these features are very different in latirostris skulls (Fig. 4 a). Multiple independent evidences therefore support that bats from the latirostris lineage belong to a distinct genus within the Myotinae radiation (Ruedi et al. 2013). In fact, while checking all relevant type material currently assigned to Asian Myotinae, we found that the very peculiar skull shape and dental characteristics of this latirostris lineage are typical of three further taxa described from continental Asia, namely Vespertilio caliginosus (Tomes 1859; type locality India), V. blanfordi (Dobson 1871; type locality northern India) and V. moupinensis (Milne-Edwards 1872; type locality Moupin, Szechuan, China). We therefore consider these four taxa as congeners, but their distinction at the species level is beyond the scope of the present study. More remarkably, we also examined the paleontological literature and found that these four species also perfectly fit the diagnosis of the Upper Miocene genus Submyotodon described by Ziegler (2003). In particular, the Miocene fossils and recent material of the latirostris lineage share not only the nyctalodont condition of the first two lower molars, but also several other dental traits (such as four-cusped third lower incisors, unreduced length of third upper molar, uncrowded premolars, etc.), a peculiar skull shape (i. e., long bony bridge between lachrymal and infraorbital foramina; ascending ramus of mandible standing nearly perpendicular to the horizontal ramus; low articular process, etc.), and very small overall dimensions (see Ziegler 2003). As the paleontological material is fragmentary, it is not possible to compare the upraised shape of posterior parts of the neurocranium, nor the elongated, relatively flattened rostrum typical of latirostris (Fig. 4 a), but as we cannot find any marked difference in other characters between the diagnosis of the Miocene Submyotodon and recent material, we propose to assign the four species of the latirostris lineage to that genus. Previous molecular dating further support this assignment, as the estimated age of the divergence time between the latirostris linage and other Myotis (credibility interval 23 – 20 million years ago, Ruedi et al. 2013 or 18 mya, Lack et al. 2010), indeed predates the paleontological age of Submyotodon (which is dated from the MN 7 / 8 stratum, i. e., 13.5 – 11 mya, Ziegler 2003). The divergence of the unrelated nyctalodont Myotis of the siligorensis – alticraniatus group is much more recent (mean 2.2 mya, Ruedi et al. 2013) and is not compatible with a Miocene divergence.	en	Ruedi, Manuel, Csorba, Gábor, Lin, Liang- Kong, Chou, Cheng-Han (2015): Molecular phylogeny and morphological revision of Myotis bats (Chiroptera: Vespertilionidae) from Taiwan and adjacent China. Zootaxa 3920 (1): 301-342, DOI: 10.11646/zootaxa.3920.2.6
03BB87963D5D5D11A8EBF6CF11FCFC33.taxon	distribution	Distribution. Submyotodon latirostris has been found so far only in Taiwan, where it is widespread and relatively common.	en	Ruedi, Manuel, Csorba, Gábor, Lin, Liang- Kong, Chou, Cheng-Han (2015): Molecular phylogeny and morphological revision of Myotis bats (Chiroptera: Vespertilionidae) from Taiwan and adjacent China. Zootaxa 3920 (1): 301-342, DOI: 10.11646/zootaxa.3920.2.6
03BB87963D5D5D11A8EBF6CF11FCFC33.taxon	description	Measurements. See Table 4. External morphology. The combination of very small body size (i. e., forearm about 31 – 34 mm) and small feet (5 – 7 mm), sharp notch to the outer edge of the ears and attachment of wing membranes to the outer toe, easily distinguish S. latirostris from any other species found in Taiwan and adjacent mainland China. The dorsal fur is dark slaty brown, long and shaggy, with lighter brown tips. Ventrally the fur is also dark brown with lighter, golden tips of hairs and is particularly long in the posterior parts. Ears nearly reach the nose tip when laid forward, are pointed, and have a sharp notch at the rear edge of the conch (Cheng et al. 2010). The tip of the relatively short tragus is spatulated, and bent forwards, while the basal parts have more parallel sides. The face is dark and hairy, but there is an area of bare, flesh-colored skin around the eyes. Thumbs are short (4 – 5 mm) and delicate, with small claws (1 – 1.5 mm). The feet are almost bare and very small, with small claws and are less than half the tibia length. The dark wing and tail membranes are sparsely haired below and close to the body, with short, whitish hairs extending along the uropatagium near the tibia. Wing membranes are attached to the outer toe. The calcar has a clearly keeled lobe, and extends along half of the rear edge of uropatagium. The penis is short and slightly clubshaped. Skull morphology. One immediate peculiarity of the skull profile of S. latirostris is the raised occipital part of the braincase (Fig. 4 a). The skull is delicate and smooth (with no visible crests), with a narrow and elongated rostrum, and thin zygomatic arches. Due to the elongated rostrum, the foramen lacrymale is relegated to the level of first molars, instead of premolars in Myotis. The toothrow shows a clear diastema between the second incisor and the canine, and premolars are not crowded and are all visible in lateral view. There are three upper and lower premolars and a well-pronounced paraconule to the first upper premolar. The upper canines are relatively large, clearly higher than last premolars; the lower ones weaker and are only slightly larger than third lower premolars. The nyctalodont first and second lower molars are unique to this Taiwanese species; the third lower molar is submyotodont, where the postcristid is bifurcated, one arm joining the hypoconulid, the other the entoconid (Ziegler 2003). Such dental characteristics only exist in other extant and fossil Submyotodon species and in few other Asian species of Myotis (see taxonomic remarks). Natural history. Although this forest species appears to be relatively common in Taiwan, it is mainly found in mountain areas above 1000 m, up to at least 2200 m. It occurs in lower areas as well, but apparently only during the winter period and is active throughout the year. Based on our observations, lactating females were recorded from April to June, and male with enlarged testis from August to March.	en	Ruedi, Manuel, Csorba, Gábor, Lin, Liang- Kong, Chou, Cheng-Han (2015): Molecular phylogeny and morphological revision of Myotis bats (Chiroptera: Vespertilionidae) from Taiwan and adjacent China. Zootaxa 3920 (1): 301-342, DOI: 10.11646/zootaxa.3920.2.6
03BB87963D5B5D10A8EBF47716E6F8DB.taxon	distribution	Distribution. Initially, the distribution of this species was limited to Fujian (type locality of both fimbriatus and hirsutus), SE China (Corbet & Hill 1992) and Taiwan (type locality of taiwanensis), where it is widespread and common (Allen 1938; Lin et al. 2004; Han et al. 2010). Recent morphological and molecular surveys (under the name taiwanensis), suggest that it is much more widespread along the eastern coast of China, occurring up to Anhui and central Shandong provinces (Han et al. 2010). As indicated previously, Yunnan specimens referred to this species (Zhang et al. 2009 and Ruedi et al. 2013) probably represent a distinct taxon (M. cf. fimbriatus).	en	Ruedi, Manuel, Csorba, Gábor, Lin, Liang- Kong, Chou, Cheng-Han (2015): Molecular phylogeny and morphological revision of Myotis bats (Chiroptera: Vespertilionidae) from Taiwan and adjacent China. Zootaxa 3920 (1): 301-342, DOI: 10.11646/zootaxa.3920.2.6
03BB87963D5B5D10A8EBF47716E6F8DB.taxon	description	Measurements. See Table 4. External morphology. This medium-sized Myotis (FA about 40 mm) is characterized by relatively large, hairy feet reaching more than half tibia length (Table 4). It has relatively long ears, reaching the nose tip when laid forwards. The tragus is nearly parallel, long and pointed, reaching half conch height. Although its dorsal fur is relatively short, it extends well along the tibia, which is typical for this species. The underside fur is also particularly long, including on the patagium along the humerus, and extends considerably on the uropatagium as well, especially along the tibia. These sparse hairs are long, cottony and reach beyond the margin of the uropatagium near the calcar (“ lumbar membranes ciliated ” as mentioned by Peters in the original description). The general color is greyish brown above. The ventral hairs have darker, slate-grey base but are much lighter near the tip, forming an almost pure white area near the anal region (Howell 1926). This gives an overall characteristic bicolored aspect to the pelage of M. fimbriatus, with a relatively sharp demarcation line along the sides of the body (see pictures in Lin et al. 2004; Cheng et al. 2010). Wing membranes are attached to the ankle or the proximal base of the metatarsus. The long calcar is unkeeled and extends to four-fifth of the rear edge of uropatagium. This bat bears external similarities with M. horsfieldii (large feet, long calcar and long ears; similar to other water-loving bats), but the latter is not hairy below the patagium, its pelage is not bicolored and the wing membranes attach to the metatarsus near the outer toe, not close to the ankle as in fimbriatus. Skull morphology. The skull has an inflated braincase, especially in the taiwanensis subspecies, giving a globose appearance when viewed from above or laterally (Fig. 4 b). Upper canines are strong and higher than premolars. The latter are aligned in the toothrow and not particularly crowded, while all are visible in the lateral view (Fig. 4 b). The lower canines are weaker, but still higher than the larger premolars. Lower molars are strong, with high cusps and all are myotodont. Natural history in Taiwan. A colonial species found in underground structures, like caves (Allen 1938) or tunnels (Han et al. 2010), where it can form quite numerous breeding colonies (up to 1000 individuals in central Taiwan, Lin et al. 2004). It is apparently linked to bodies of water, where it hunts insects above the water surface, like other trawling bats. It is found in both lowland and mountain habitats. In Taiwan, newborns were recorded from November to June, suggesting an extensive breeding season. Such unusual winter reproduction was observed in artificial tunnels containing pipes transporting hot spring water, which certainly enhanced local ambient temperature to provide suitable conditions for the bats to extend their normal breeding time.	en	Ruedi, Manuel, Csorba, Gábor, Lin, Liang- Kong, Chou, Cheng-Han (2015): Molecular phylogeny and morphological revision of Myotis bats (Chiroptera: Vespertilionidae) from Taiwan and adjacent China. Zootaxa 3920 (1): 301-342, DOI: 10.11646/zootaxa.3920.2.6
03BB87963D5A5D12A8EBF71F1389FD76.taxon	distribution	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.	en	Ruedi, Manuel, Csorba, Gábor, Lin, Liang- Kong, Chou, Cheng-Han (2015): Molecular phylogeny and morphological revision of Myotis bats (Chiroptera: Vespertilionidae) from Taiwan and adjacent China. Zootaxa 3920 (1): 301-342, DOI: 10.11646/zootaxa.3920.2.6
03BB87963D5A5D12A8EBF71F1389FD76.taxon	description	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. 4 c, 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. 4 c). 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.	en	Ruedi, Manuel, Csorba, Gábor, Lin, Liang- Kong, Chou, Cheng-Han (2015): Molecular phylogeny and morphological revision of Myotis bats (Chiroptera: Vespertilionidae) from Taiwan and adjacent China. Zootaxa 3920 (1): 301-342, DOI: 10.11646/zootaxa.3920.2.6
03BB87963D585D1FA8EBF2B41078FD13.taxon	materials_examined	Holotype. Adult male collected by C. H. Chou on 8 July 2003 (field number # B 030018). The dry skin and skull are deposited in the collections of the Department of Biology, Tunghai University under accession number THUMB 30018. The complete mitochondrial Cyt b gene was sequenced from tissue extracts taken from the holotype and deposited in GenBank under accession number KP 187896 and was used in the phylogenetic analyses presented herein (Clade VIII in figure 3). Paratypes. The identity of all six paratypes listed hereafter was confirmed by both the multivariate morphological (Fig. 2) and genetic analyses (Fig. 3). These paratypes are 5 adult males and 1 adult female from near Kaohsiung City in Taoyuan District collected between 2002 and 2004 (TESRI B 0271, THUMB 92, 30017, 30020, 30051 – 2). The following GenBank accession numbers correspond to the complete mitochondrial Cyt b gene sequenced in these paratypes: KP 187889, KP 187895 – 97, KP 187899 – 00 (Table 1). Other referred material. 3 adult males from near Datong Township, Yilan County (TESRI B 0273 – 5); 1 adult female from Fuxing Township, Taoyuan County (TESRI B 0272); 3 adult males from near Wufeng Township, Hsinchu County (TESRI B 0265, B 0267, B 0268); 2 adult male and 1 adult female from near Taichung City, Heping District (NMNS t- 3474, TESRI B 0270, THUMB 30013); 1 adult male and 1 adult female from near Yuli Township, Hualian County (THUMB 30023, 30031); 2 adult females and 1 adult male from Taoyuan District, Kaohsiung City (THUMB 30053, 30016, 30019); 1 adult male from Yuli Township, Hualian County (THUMB 30055); 1 adult male from Donghe Township, Taitung County (TESRI B 0269). Type locality. The designated type series originate from the forest areas near Kaohsiung City (Taoyuan District) in south-central Taiwan ROC (approximate coordinates: 23 ° 08 ’ N, 120 ° 48 ’ E).	en	Ruedi, Manuel, Csorba, Gábor, Lin, Liang- Kong, Chou, Cheng-Han (2015): Molecular phylogeny and morphological revision of Myotis bats (Chiroptera: Vespertilionidae) from Taiwan and adjacent China. Zootaxa 3920 (1): 301-342, DOI: 10.11646/zootaxa.3920.2.6
03BB87963D585D1FA8EBF2B41078FD13.taxon	distribution	Distribution. M. secundus sp. n. has so far only been recorded in Taiwan, where it is relatively common and widespread in forested habitats across most of the central regions. It was found near locations 1 - 4, 7, 15, 17, 19, 21 and 21 (Fig. 1).	en	Ruedi, Manuel, Csorba, Gábor, Lin, Liang- Kong, Chou, Cheng-Han (2015): Molecular phylogeny and morphological revision of Myotis bats (Chiroptera: Vespertilionidae) from Taiwan and adjacent China. Zootaxa 3920 (1): 301-342, DOI: 10.11646/zootaxa.3920.2.6
03BB87963D585D1FA8EBF2B41078FD13.taxon	etymology	Etymology. We name it secundus (meaning second in Latin), as it was the second unnamed taxon found in Taiwan and initially mentioned under the name Myotis sp. 2 (Chou 2004; Lin et al. 2004; Cheng et al. 2010; Ruedi et al. 2013). In the photographic guide to the bats of Taiwan (Cheng et al. 2010), it was illustrated under the vernacular name of Long-toed Myotis, although this attribute is not particularly developed in this species.	en	Ruedi, Manuel, Csorba, Gábor, Lin, Liang- Kong, Chou, Cheng-Han (2015): Molecular phylogeny and morphological revision of Myotis bats (Chiroptera: Vespertilionidae) from Taiwan and adjacent China. Zootaxa 3920 (1): 301-342, DOI: 10.11646/zootaxa.3920.2.6
03BB87963D585D1FA8EBF2B41078FD13.taxon	description	Measurements of the holotype. Measurements are in mm. Head and body length, 37; tail length, 41; forearm length, 35.7; hind foot length (including claw), 7.1; tibia length, 17.5; thumb length, 6.6; ear length, 12.8; tragus length, 6.2; greatest skull length, including incisors, 14.1; greatest zygomatic breadth, 8.4; postorbital breadth, 3.3; mastoid breadth, 6.7; greatest braincase width, 6.3; upper canine-molar toothrow, 5.4; width across upper canines, 3.4; width across 3 rd upper molars, 5.6.	en	Ruedi, Manuel, Csorba, Gábor, Lin, Liang- Kong, Chou, Cheng-Han (2015): Molecular phylogeny and morphological revision of Myotis bats (Chiroptera: Vespertilionidae) from Taiwan and adjacent China. Zootaxa 3920 (1): 301-342, DOI: 10.11646/zootaxa.3920.2.6
03BB87963D585D1FA8EBF2B41078FD13.taxon	diagnosis	Diagnosis. Small-sized Myotis (weight 3 – 5 g) with dark brown pelage color, slightly lighter ventrally, without demarcation between dorsal and ventral sides. Face and ears dark brown becoming more flesh-colored near the base of ears and above the eyes. Wing membranes attached to base of outer toe. Feet about half tibia length. Tail approximately same length as head and body length. Ears relatively long and narrow with a slight notch on the proximal third of the rear edge of conch. Tragus elongated and narrow, reaching the ear notch. Skull profile slender and relatively angular, with nearly flat braincase apex and abruptly falling posterior parts (Fig. 6 a); frontal part of braincase gently rising from rostrum to the top. Anterior part of the rostrum narrow and convergent. Teeth relatively robust with upper premolars aligned in toothrow and all visible in side view (Figs. 4 d, 6 a). The second upper and lower premolars are distinctly smaller than corresponding first premolars. Canines relatively strong, slightly larger than premolars. All lower molars are myotodont.	en	Ruedi, Manuel, Csorba, Gábor, Lin, Liang- Kong, Chou, Cheng-Han (2015): Molecular phylogeny and morphological revision of Myotis bats (Chiroptera: Vespertilionidae) from Taiwan and adjacent China. Zootaxa 3920 (1): 301-342, DOI: 10.11646/zootaxa.3920.2.6
03BB87963D585D1FA8EBF2B41078FD13.taxon	description	Description. This small Myotis has a long, shaggy and very dark brown pelage, with a tinge of lighter brown color at hair tips, giving a frosted appearance to its fur (unlike e. g., M. yanbarensis). The underparts are lighter, buffier, also with pale brown hair tips. The face is densely haired, with a small bare part located above the eyes (Cheng et al. 2010). The patagium is essentially naked and dark brown, with wing membrane attaching to the base of outer toe. The ears are relatively long and narrow, with a small but distinct notch. The tragus is narrow, straight, slightly bent forward close to the tip; it extends to about a third of the distance to the tip of ear. Feet (about 7 – 9 mm, including claw) and thumbs are slender, both with sparse guard hairs and slender claws. The tibia is long (16 – 17.7 mm). Tail (about 40 mm) is as long as head and body length. When viewed in profile (Figs. 4 d, 6 a), the braincase of the skull has gently rising frontal parts, and is nearly flat on its summit; the occipital region is rounded, smooth with no visible crests. The frontal part of the rostrum is narrow and, in front of the canines, is convergent. Mean and range of external and skull measurements of M. secundus sp. n. from Taiwan are given in Table 4. Dental formula I 2 / 3 C 1 / 1 PM 3 / 3 M 3 / 3, comprising the adult dentition of 38 teeth. The teeth are not particularly weak, although the canines (both upper and lower) are only slightly higher than the corresponding last premolars (Figs. 4 d, 6 a). The second incisor is clearly smaller than the first (in lateral view), although in occlusal view both are of similar basal dimensions. All premolars are in the toothrow and visible in lateral view, but the second one (both upper and lower) is much smaller than the first one (Fig. 4 d). The third upper premolar has a distinct paraconule. Lower molars are all myotodont. Comparisons. In recent taxonomic accounts (e. g., Corbet & Hill 1992, Simmons 2005), a single small species of Myotinae was found on Taiwan and was called Myotis muricola latirostris (see above for its current taxonomic position). However, since 2004, bat surveys showed that a second small (weight 3 – 5 g), relatively common Myotis species, that was not fitting the characteristics of latirostris, was living in various habitats of the island. It was referred subsequently as Myotis sp. 2, and hence named here M. secundus sp. n. Although both species are indeed small and have a dark, almost black dorsal pelage, the new species is easily distinguished from S. latirostris by its larger external dimensions such as thumb or tibia length, or by mostly larger skull dimensions as well (Table 4). M. secundus sp. n. has myotodont lower molars, whereas they are nyctalodont in S. latirostris. This character also distinguishes the new species from other small, blackish Myotis from continental Asia pertaining to the siligorensis group, such as M. siligorensis, M. badius, M. alticraniatus, etc. (see Tiunov et al. 2011). Another small Myotis living in south-east China and described originally as M. sowerbyi (Howell 1926), is also very similar to M. secundus sp. n. (similar external dimensions, wing insertion to the base of the toes, myotodont lower molars), but is clove-brown, not blackish, has much more slender upper (Fig. 8 b) and lower canines, has a first lower premolar nearly as large as the canine (it is only half its size in M. secundus sp. n., Fig. 4 d), and in skull profile it has a more inflated and elevated braincase and rostrum (Fig. 6 b). M. davidii is also a small myotodont species, but has the upper premolars crowded, the second being completely displaced lingually from toothrow and is now considered as belonging to the mystacinus morpho-group (see illustrations in Benda et al. 2012). Compared to the closest relatives of M. secundus sp. n. in phylogenetic analyses (Clade VIII in figure 3), M. pruinosus has a less elevated frontal part of skull and relatively flat braincase (Fig. 6 c), whereas the skull of M. yanbarensis (known from Okinawa, Japan) is characterized by a longer rostrum, more globose braincase (Fig. 6 d) and is larger (e. g., CBL over 14 mm,) than M. secundus sp. n. (CBL about 13 mm). Furthermore, the tibia is particularly long in M. secundus sp. n. (about 17 mm), unlike in related M. pruinosus from Japan (13 mm). The tail of M. secundus sp. n. is nearly as long as head and body length, whereas it is distinctly longer in M. yanbarensis. Phylogenetic relationships. The complete (1140 bp) Cyt b gene of the holotype and paratypes of M. secundus sp. n. are all similar (with less than 1 % K 2 P distance) and diverge by at least 10.5 % K 2 P distance from their closest relative, M. yanbarensis (Fig. 3) and by at least 20 % for any other sympatric Myotis from Taiwan (Table 5). In a more general phylogenetic context, M. secundus sp. n. (named Myotis sp. 2 in Ruedi et al. 2013) is part of the strongly supported Clade VIII within the Myotinae radiation, together with M. yanbarensis, M. pruinosus, and representatives of the M. montivagus species complex. It also differs notably from species of the mystacinus morpho-group that appear in Clade VI (Fig. 3 and Ruedi et al. 2013). Natural history. M. secundus sp. n. is essentially a forest-dwelling species, but we observed breeding females also occupying small holes or chinks in caves. It is common both in the lowlands and higher mountains, suggesting that this species may have a relatively broad ecological niche. It was found in sympatry with S. latirostris in the higher elevations. Lactating females were observed from May to July, while males with enlarged testis were recorded from August to March.	en	Ruedi, Manuel, Csorba, Gábor, Lin, Liang- Kong, Chou, Cheng-Han (2015): Molecular phylogeny and morphological revision of Myotis bats (Chiroptera: Vespertilionidae) from Taiwan and adjacent China. Zootaxa 3920 (1): 301-342, DOI: 10.11646/zootaxa.3920.2.6
03BB87963D555D19A8EBF54513E1F867.taxon	materials_examined	Holotype. Adult female collected by G. Csorba, C. - C. Huang and H. - C. Kuo on 5 September 2003 (field number # CSOTA 141). The dry skin and skull are deposited in the collections of the Hungarian Natural History Museum (HNHM) under accession number 2003.36.20. Part of the mitochondrial Cyt b (413 bp) was sequenced from tissue extracts taken from the holotype; this sequence is deposited in GenBank under accession number KP 187901. The Cyt b gene was used in the phylogenetic analyses presented herein (Fig. 3). Type locality. The holotype and only known specimen was caught along the “ water pipe road ”, near the Highland Experimental Farm of National Taiwan University, Nantou County, Taiwan ROC (approximate coordinates: 24 ° 05 ’ N, 121 ° 09 ’ E), at an altitude of 2100 m above sea level (location 9 in figure 1).	en	Ruedi, Manuel, Csorba, Gábor, Lin, Liang- Kong, Chou, Cheng-Han (2015): Molecular phylogeny and morphological revision of Myotis bats (Chiroptera: Vespertilionidae) from Taiwan and adjacent China. Zootaxa 3920 (1): 301-342, DOI: 10.11646/zootaxa.3920.2.6
03BB87963D555D19A8EBF54513E1F867.taxon	distribution	Distribution. The only known specimen of M. soror sp. n. is an adult female caught in a mist-net placed across a small road at Meifong, Nantou County, in the slopes of Mt Hehuan, in central Taiwan (location 9 in figure 1). The evergreen, temperate forests surrounding this area is the typical vegetation found at this elevation (2100 m). Other species of bat caught in the same area during our survey include an unknown species of Pipistrellus, Harpiola isodon, M. frater, M. laniger, M. secundus sp. n., S. latirostris, Murina gracilis, Plecotus taivanus, Miniopterus fuliginosus, Barbastella leucomelas, Rhinolophus monoceros and R. formosae. As none of the previous, intensive surveys of the bat fauna of Taiwan evidenced this species elsewhere (Chou 2004; Lin et al. 2004), M. soror sp. n. might indeed be a very rare forest species. Nothing is known about its natural history besides that the female was in a post-lactating stage (with teats still enlarged and devoid of hairs) when it was collected, and thus must have been breeding recently in the area.	en	Ruedi, Manuel, Csorba, Gábor, Lin, Liang- Kong, Chou, Cheng-Han (2015): Molecular phylogeny and morphological revision of Myotis bats (Chiroptera: Vespertilionidae) from Taiwan and adjacent China. Zootaxa 3920 (1): 301-342, DOI: 10.11646/zootaxa.3920.2.6
03BB87963D555D19A8EBF54513E1F867.taxon	etymology	Etymology. We name it soror (meaning sister in Latin) as it is clearly related to Myotis frater, its sister species in phylogenetic reconstructions (see Clade III in figure 3). Owing to its peculiar coloration, we suggest Reddish Myotis as an appropriate vernacular name.	en	Ruedi, Manuel, Csorba, Gábor, Lin, Liang- Kong, Chou, Cheng-Han (2015): Molecular phylogeny and morphological revision of Myotis bats (Chiroptera: Vespertilionidae) from Taiwan and adjacent China. Zootaxa 3920 (1): 301-342, DOI: 10.11646/zootaxa.3920.2.6
03BB87963D555D19A8EBF54513E1F867.taxon	description	Measurements of the holotype. Measurements are in mm and, unlike in other species, were taken on the prepared specimen. Head and body length, 48; tail length, 41; forearm length, 42.1; hind foot length (including claw), 7.6; tibia length, 17.2; thumb length, 5.1 and claw 2.5; ear length, 11; tragus length, 6; greatest skull length, including incisors, 13.8; greatest zygomatic breadth, 8.9; postorbital breadth, 4.1; mastoid breadth, 7.8; greatest braincase width, 7.2; upper canine-molar toothrow, 5.2; width across upper canines, 4.2; width across 3 rd upper molars, 5.7 (Table 4).	en	Ruedi, Manuel, Csorba, Gábor, Lin, Liang- Kong, Chou, Cheng-Han (2015): Molecular phylogeny and morphological revision of Myotis bats (Chiroptera: Vespertilionidae) from Taiwan and adjacent China. Zootaxa 3920 (1): 301-342, DOI: 10.11646/zootaxa.3920.2.6
03BB87963D555D19A8EBF54513E1F867.taxon	diagnosis	Diagnosis. Medium-sized Myotis with striking pelage color, rich cinnamon-brown with lighter, golden hair tips dorsally (Fig. 7 a) and only slightly lighter ventrally. Color of face brownish and other bare parts dark brown. Wing membranes attached close to base of outer toe, on the distal part of the metatarsus (Fig. 7 b). Feet nearly half tibia length. Tail notably shorter than head and body length. Ears relatively short and broad with a distinct notch on the middle of the rear edge of conch (Fig. 7 c). The inner and outer sides of the conch are covered with sparse, cinnamon-rufous hairs. Skull angular, with short rostrum and abruptly raised frontal part of braincase (as seen in profile, Fig. 8 a). Teeth relatively robust with second upper premolars small and moderately displaced inside from toothrow, but not visible in side view. Upper canines strong, with a marked groove along the labial edge.	en	Ruedi, Manuel, Csorba, Gábor, Lin, Liang- Kong, Chou, Cheng-Han (2015): Molecular phylogeny and morphological revision of Myotis bats (Chiroptera: Vespertilionidae) from Taiwan and adjacent China. Zootaxa 3920 (1): 301-342, DOI: 10.11646/zootaxa.3920.2.6
03BB87963D555D19A8EBF54513E1F867.taxon	description	Description. The pelage is dense, soft and relatively long on the dorsum (about 6 mm); dorsal hairs cinnamon brown on the base and becoming progressively lighter towards the tips, the last 2 mm being golden yellow, giving a frosted appearance to the dorsal fur. Ventral parts appear slightly paler, with hairs brown to the middle, becoming lighter, almost creamy towards the tip. Bare parts, including ears and patagium, reddish-brown, with face lighter, flesh colored. Ears short and broad, not reaching the nose tip when laid forward; conspicuous notch at two-thirds of its height on the rear edge (Fig. 7 c). Long (up to 1.6 mm), sparse cinnamon hairs are present on both inner and outer sides of the conch. Tragus relatively short, but reaching the ear notch; the anterior edge is slightly convex but the exterior margin has a small lobe at the base (hidden into the conch), and a larger one along half of its length; the tragus is convergent and tapers medially forming a slightly rounded head at the tip (Fig. 7 c). Wing membranes are essentially naked (except the underparts, close to the body), and attached to the distal parts of the metacarpus, near the base of the outer toe (Fig. 7 b). Feet are robust, with strong curved claws, and about half the size of tibia length. Uropatagium broad with tail almost fully included in the membrane. The calcar is short (about one-third uropatagium edge length), and bears an indistinct, unkeeled lobe. The tail is notably shorter than head and body length. When viewed in profile, the skull has a relatively short rostrum, and an abruptly raising frontal part of the braincase (Fig. 9 a). The occipital part looks like a square in profile or when viewed from above (Fig. 8 a). Lambdoid crests are visible laterally, but no notable sagittal crest is present. Dental formula I 2 / 3 C 1 / 1 PM 3 / 3 M 3 / 3, comprising the adult dentition of 38 teeth. Teeth are robust, with upper canines longer than third premolars, but the lower canines are much weaker, barely reaching the height of lower premolars (Fig. 8 a). The two upper incisive are nearly as high as wide, short, of comparable size and both are visible in side view. The upper canines have a distinct groove along the labial edge. The first upper premolar is small and aligned in toothrow, but the second is minute and displaced inwards and barely visible in side view. The third premolar is large, but bears an inconspicuous paraconule in its front margin. Molars are robust, but low, with ill-defined paraconules. The three lower premolars are in a row, not particularly crowded. All lower molars are myotodont. Comparisons. Owing to its short rostrum, raised frontal parts of braincase, angular skull shape (Figs. 8 a, 9 a) and inwards displaced second upper premolars, Myotis soror sp. n. is clearly related to the M. frater species complex (sensu Tsytsulina & Strelkov 2001). However, it differs genetically from sympatric specimens of M. frater by at least 11 % (K 2 P divergence at the Cyt b gene; Table 5). Its rich cinnamon pelage (Fig. 7 a) with golden frosted appearance of the dorsal fur is also unique in this group, all other taxa living in temperate regions being darker brown (sometimes lighter brown in juvenile specimens, Yoshiyuki 1989). The desert form M. bucharensis is decidedly much paler and slightly larger. With a forearm length of 42.1 mm, M. soror sp. n. is also larger than the Far Eastern taxa (longicaudatus, kaguyae and eniseensis) and slightly larger than sympatric M. frater s. s. (FA barely reaching 41 mm; Table 4). The tail of M. soror sp. n. is shorter than head and body length, and thus is considerably shorter than in other taxa in this group (Table 4). Compared to M. frater s. s. the shape and pilosity of ears differ notably, the notch on the rear edge being less visible in the later species, with pilosity confined to inner side (hairy on both sides in M. soror sp. n., Fig. 7 c). Due to an absolute shorter tibia length (about 17 mm) compared to other taxa in the frater s. l. group (about 20 mm or more), the feet (about 8 mm) appear relatively large, almost half the tibia length, whereas they are much shorter than 50 % tibia length in the other related taxa. Other, unrelated Asian species of Myotis with inwards displaced second upper premolars, such as M. davidii, are either much smaller (FA about 31 mm) and have a darker, blackish pelage color, or are much larger (FA about 50 mm) and have particolored wings (see M. formosus s. l. group, Csorba et al. 2014). Phylogenetic relationships. The partial (413 bp) Cyt b gene of the holotype of M. soror sp. n. diverges by at least 11 % K 2 P distance from any other homologous sequences, including from its sympatric, sister taxon M. f. frater (Fig. 2; Table 5).	en	Ruedi, Manuel, Csorba, Gábor, Lin, Liang- Kong, Chou, Cheng-Han (2015): Molecular phylogeny and morphological revision of Myotis bats (Chiroptera: Vespertilionidae) from Taiwan and adjacent China. Zootaxa 3920 (1): 301-342, DOI: 10.11646/zootaxa.3920.2.6
03BB87963D515D24A8EBF2BF1748FC32.taxon	distribution	Distribution. The known distribution of M. frater s. s. as understood here is restricted to Fujian Province (formerly Fukien) in eastern China, and Taiwan.	en	Ruedi, Manuel, Csorba, Gábor, Lin, Liang- Kong, Chou, Cheng-Han (2015): Molecular phylogeny and morphological revision of Myotis bats (Chiroptera: Vespertilionidae) from Taiwan and adjacent China. Zootaxa 3920 (1): 301-342, DOI: 10.11646/zootaxa.3920.2.6
03BB87963D515D24A8EBF2BF1748FC32.taxon	description	Measurements. See Table 4. External morphology. Medium sized bat with a dense, soft pelage that extends well over the underparts of the tail and wing membranes, up to the elbows and around the anal region. The dorsal color is dark brown, without gloss, whereas the underparts are lighter, creamy brown, most of the hair base being darker brown. The wing membranes are broad and attached near the base of the outer toe, at the distal end of the metacarpus (see illustration in Cheng et al. 2010). Feet are relatively small, much shorter than half tibia length. Ears are short, relatively angular in shape, the front edge being vertical to three-quarters of it length, then bending sharply backwards at an angle of 45 °; the rear edge has a distinct notch at mid-height; the inner side of the conch along the front edge is hairy, especially at its base, where long hairs form a comb-like structure; the outer surface of the ear is naked (see illustration in Cheng et al. 2010), unlike in M. soror sp. n. The calcar is long (half of the free edge of the uropatagium), with a narrow but unkeeled lobe. The uropatagium is particularly large, and is sparsely haired on the underside, near the anal region. The tragus is relatively long (longer than notch height), nearly straight but bending forwards and lacking a distal lobe. The external morphology resembles that of M. soror sp. n., except for color (richer cinnamon and frosted aspect of fur in the later), but differs from this new species by having longer tibia (20 mm instead of 17 mm) and slightly longer tail (about the same as head and body length, instead of being shorter in M. soror sp. n.). Skull morphology. The outlines of the skull viewed from above are angular, fitting almost in a cube. In profile, the short rostrum, abruptly raised frontal part of the braincase and angular occipital region are characteristic of the M. frater complex (Fig. 9). The upper canines are strong and bear a typical, deep groove along their labial edge. The incisors are short but robust. The second upper premolar is displaced inwards and invisible in profile (see the type of frater, Fig. 9 b), but the extent of this intrusion is variable within the same population. Natural history. In Taiwan, this forest dwelling bat is widely distributed, from low to higher elevations. It was found roosting in tree holes, but never in caves. Lactating females were found from June to August, depending on altitude. Males started showing enlarged testis in August. In Yenping (Fujian) three specimens (including the type) were taken in holes of live bamboo stems in a mountain area at an altitude of about 1000 m (Allen 1923).	en	Ruedi, Manuel, Csorba, Gábor, Lin, Liang- Kong, Chou, Cheng-Han (2015): Molecular phylogeny and morphological revision of Myotis bats (Chiroptera: Vespertilionidae) from Taiwan and adjacent China. Zootaxa 3920 (1): 301-342, DOI: 10.11646/zootaxa.3920.2.6
03BB87963D6E5D27A8EBF477129AFD31.taxon	distribution	Distribution. M. formosus is a rare, but relatively widespread species, found from Afghanistan, along the foothills of the Himalaya (Csorba et al. 2014) east to Jiangxi province in China (Jiang et al. 2010), and Taiwan.	en	Ruedi, Manuel, Csorba, Gábor, Lin, Liang- Kong, Chou, Cheng-Han (2015): Molecular phylogeny and morphological revision of Myotis bats (Chiroptera: Vespertilionidae) from Taiwan and adjacent China. Zootaxa 3920 (1): 301-342, DOI: 10.11646/zootaxa.3920.2.6
03BB87963D6E5D27A8EBF477129AFD31.taxon	description	Measurements. See Table 4 for measurements of the Taiwanese subspecies, M. f. flavus. Measurements for the nominal subspecies are given in Csorba et al. (2014). External morphology. This spectacular, relatively large Myotis has a unique cottony and yellowish fur, both above and below, the dorsal parts being only slightly darker at the hair tips (i. e., no ” smoked ” aspect of fur as in M. rufoniger). The wing membranes are parti-colored, with a characteristic orange (along bones) and black patterning (see pictures in Lin et al. 2004). The uropatagium is essentially orange, as are the ears (unlike in M. rufoniger, which has conspicuous black margins to the ears) and face. Thumbs and hairy feet are also largely orange, except close to the claws, which are strong and black. Wings are attached to the base of the outer toe. Skull morphology. The skull is large, with massive canines and strong molars. The second upper premolar is much smaller, less than half the size of the first, and is usually completely displaced lingually from toothrow and thus invisible in side view (Fig. 8 c). The posterior parts of the braincase are high and globose, with very weak or no occipital or lambdoid crests (unlike in M. rufoniger, which has a more angular skull and marked crests). Natural History. This beautiful, but rare bat is known from few specimens, and owing to its large distribution across Asia is likely to have different ecologies throughout its range. In Taiwan, it is found exclusively in the lowlands, where it roosts either among tree foliage (Swinhoe 1870; Chen et al. 2010), or in buildings (Lin et al. 2004). In both situations, the bats are fully exposed to the light and seem to rely on their particular coloration to avoid predation. They occupy the breeding colonies (which may number up to several hundred individuals) between March and July. In October, they disappear from the breeding colonies to hibernate in unknown winter roosts.	en	Ruedi, Manuel, Csorba, Gábor, Lin, Liang- Kong, Chou, Cheng-Han (2015): Molecular phylogeny and morphological revision of Myotis bats (Chiroptera: Vespertilionidae) from Taiwan and adjacent China. Zootaxa 3920 (1): 301-342, DOI: 10.11646/zootaxa.3920.2.6
03BB87963D6D5D26A8EBF30C10B7FE2A.taxon	distribution	Distribution. M. rufoniger is apparently also a rare species that has only been collected in a handful of localities. It has been found in northern Vietnam and Laos, Taiwan, throughout China, North and South Korea and on Tsushima Island in the Japanese archipelago (see details in Csorba et al. 2014).	en	Ruedi, Manuel, Csorba, Gábor, Lin, Liang- Kong, Chou, Cheng-Han (2015): Molecular phylogeny and morphological revision of Myotis bats (Chiroptera: Vespertilionidae) from Taiwan and adjacent China. Zootaxa 3920 (1): 301-342, DOI: 10.11646/zootaxa.3920.2.6
03BB87963D6D5D26A8EBF30C10B7FE2A.taxon	description	Measurements. See Table 4 for measurements of the specimens from Taiwan. External morphology. M. rufoniger is a second, large species of Oriental Myotis with a striking orange and black patterning of the wing membranes similar to that of M. formosus. However, unlike the latter, its wooly fur is darker, more rusty orange (both above and below), and hairs on the dorsal parts are tricolored (slate grey at the base, yellowish in the middle and dark brown near the tips), which gives a smoked appearance to its dorsal pelage (see figure at pages 100 and 112 in Lin et al. 2004). Other typical external characters include the ears (which are orange bordered extensively with black along the edge of the conch), and the nose tip, thumbs and feet (which are also entirely black, not orange). The face and uropatagium are essentially orange, but darker than in formosus. Wings are attached to the base of the outer toe, and are furred along the body. Feet are strong and hairy, about one third of tibia length. The tragus is sickle-shaped, bending forwards, and is relatively pointed. Skull morphology. The dimensions of the skull and teeth of M. rufoniger are slightly smaller than those of M. formosus (see Table 4). Differences include a more angular posterior part of the braincase, and the presence of marked occipital and lambdoid crests (Fig. 8 d). The second upper premolar is only slightly smaller than the first, and usually only barely displaced from toothrow (Fig. 8 d), and hence visible when viewed from profile (unlike in M. formosus, Fig. 8 c). Natural history. Myotis rufoniger is also an uncommon bat whose biology and habitat requirements are largely unknown. In Korea and Taiwan (Lin et al. 2004), it is found in mountain forests, where is roosts in caves, and a female with young was reported in August.	en	Ruedi, Manuel, Csorba, Gábor, Lin, Liang- Kong, Chou, Cheng-Han (2015): Molecular phylogeny and morphological revision of Myotis bats (Chiroptera: Vespertilionidae) from Taiwan and adjacent China. Zootaxa 3920 (1): 301-342, DOI: 10.11646/zootaxa.3920.2.6
