Materials

All specimens examined are listed under the corresponding taxa in Appendix I. Abbreviations referring to collections include AMNH: American Museum of Natural History, New York, United States; BMNH: The Natural History Museum (formerly British Museum [Natural History]), London, United Kingdom; CPC: Cuc Phuong National Park Reference Collection, Cuc Phuong, Vietnam; DWNP: Department of Wildlife and National Parks, Kuala Lumpur, Malaysia; FMNH: Field Museum of Natural History, Chicago, United States; HNHM: Hungarian Natural History Museum, Budapest, Hungary; IEBR: Institute of Ecology and Biological Resources, Hanoi, Vietnam; MHC: Harada Collection, Osaka City University, Osaka, Japan; MHNG: Muséum d'histoire naturelle de Genève, Geneva, Switzerland; MNHN: Museum national d'histoire naturelle, Paris, France; MZB: Museum Zoologicum Bogoriense, Bogor, Indonesia; NF: Kim Hy Nature Reserve Collection, Hanoi, Vietnam; NSMT: National Museum of Nature and Science (formerly National Science Museum), Tokyo, Japan; NMNS: National Museum of Natural Science, Taichung, Taiwan; PSUZC: Prince of Songkla University Zoological Collection, Hat Yai, Thailand; RMNH: Naturalis Biodiversity Center (formerly Rijksmuseum van Natuurlijke Historie), Leiden, Netherlands; ROM: Royal Ontario Museum, Toronto, Canada; SMC: Sumiko Matsumura Collection, Yamaguchi, Japan; SMF: Forschungsinstitut und Natur-Museum Senckenberg, Frankfurt a. M., Germany; THU: Tunghai University, Taichung, Taiwan; USNM: National Museum of Natural History (formerly United States National Museum), Washington, D.C, United States; ZMB: Museum für Naturkunde (formerly Zoological Museum), Berlin, Germany.

Measurements

Forearm length (FA) data were compiled from the literature or measured by the authors from museum specimens to the nearest 0.1 mm. Craniodental measurements were taken to the nearest 0.01 mm using digital calipers and a stereomicroscope. Measurements include only those taken from nonjuveniles, as indicated by the presence of fully ossified metacarpal–phalangeal joints. Abbreviations and definitions for craniodental measurements follow Bates and Harrison (1997) and are: GTL—greatest length of skull, from the anterior of the 1st upper incisor to the most posteriorly projecting point of the occipital region; CCL—condylo–canine length, from the exoccipital condyle to the most anterior part of the canine; CCW—greatest width across the upper canines from their buccal borders; M3M3W—greatest width across the crowns of the last upper molars from their buccal borders; IOW—least width of the interorbital constriction; ZYW—greatest width of the skull across the zygomatic arches; MAW—greatest distance across the mastoid region; BCW—greatest width of the braincase; BCH—braincase height, from the basisphenoid at the level of the hamular processes to the most dorsal part of the skull, including the sagittal crest (if present); CM3L—maxillary toothrow length, from the anterior of the upper canine to the posterior of the crown of the 3rd molar; CP4L—distance from the anterior of the upper canine to the posterior of the crown of the last premolar; ML—mandible length, from the anterior rim of the alveolus of the 1st lower incisor to the most posterior part of the condyle; cm3L—mandibular toothrow length, from the anterior of the lower canine to the posterior of the crown of the 3rd lower molar; and CPH—height of the coronoid process, from its dorsal tip to the apex of the indentation on the ventral surface of the ramus adjacent to the angular process.

All statistical analyses were carried out with R 2.13.2 (R Development Core Team 2012). Measurements were compared using Welch 2-sample t-tests. A 1-way analysis of variance (ANOVA) model with Tukey's pairwise tests was used for comparisons of selected external and craniodental measurements. All variables showed normal distributions in quantile-comparison plots and all tests were 2-tailed. To eliminate false discoveries in multiple tests, the adjustment method of Benjamini and Hochberg (1995) was applied.

Phylogenetic reconstruction

Thirty-six Cytb sequences (1,140 base pairs) were downloaded from GenBank to build a phylogenetic tree of available species belonging to Chrysopteron and a number of other representative Myotis taxa (Table 1). In addition, homologous sequences of 3 outgroup taxa (Murina cyclotis, Mu. pluvialis, and Kerivoula papillosa) were used to root the trees. Sequences were aligned in MEGA5 (Tamura et al. 2011), and the analyses were completed in MrBayes version 3.2.1 (Ronquist and Huelsenbeck 2003) using the Bayesian interference method and in RAxML (Stamatakis 2006) to obtain a maximum-likelihood tree. Reliability of nodes in maximum-likelihood analyses was assessed by 1,000 standard bootstraps with RAxML. All analyses were done using a partitioned scheme where each codon position was allowed to have specific model parameters. The general time reversible model with “gamma” and “invariant sites” was used in each partition, as suggested by the results of jModelTest version2.1.3 (Darriba et al. 2012) applied to the alignment.

Table 1.

Origin and GenBank numbers with corresponding references of the 39 cytochrome-b sequences from Myotis analyzed. Both species of Murina and Kerivoula were used as outgroups for the phylogenetic reconstructions.

To obtain Bayesian interference trees, MrBayes was run for 10 × 10⁶ generations and sampled every 1,000 generations. The first 1 × 10⁶ generations were discarded as burn-in. Posterior probabilities were subsequently computed from the consensus of the remaining sampled trees. Two independent replicate analyses were performed on the same data set, and the results were then combined. Effective sample sizes for the estimated parameters and posterior probability were calculated with Tracer version 1.5 (Rambaut and Drummond 2007) and were all higher than 200.

The genetic distances between and within each species were calculated in MEGA5 (Tamura et al. 2011) using the Kimura 2-parameter model (Kimura 1980).

Status of Chrysopteron

The monophyletic lineage known as the “Ethiopian clade” within the phylogenetic tree of Myotis (Stadelmann et al. 2004b, 2007; Jiang et al. 2010; Ruedi et al. 2013) was recovered in our analyses with high support (> 95% posterior probability and boostrap value) and includes 6 Ethiopian, 1 Palearctic, and 2 Indomalayan species (Fig. 1). The majority of the taxa grouped in this clade had previously been regarded as closely related based on phenetic analyses (Findley 1972). The species referred to as “M. flavus” (e.g., Jiang et al. 2010) is identified by us as M. formosus, and it does not form a sister group with M. rufoniger, the other species of Indomalayan parti-colored bat (variously designated as “M. watasei” or “M. formosus” by different authors [e.g., Jiang et al. 2010]). Rather, M. formosus (as understood here) is associated with the European M. emarginatus and the African M. tricolor (low nodal support among these 3 species impedes more precise understanding of their phylogenetic relationships), whereas M. rufoniger is sister to the African M. welwitschii (with high support). The Ethiopian clade is itself part of the Old World species assemblage as detailed in a more comprehensive analysis of the genus (Ruedi et al. 2013).

Fig. 1.

Bayesian consensus tree for 19 species of Myotis and 3 outgroup species based on an alignment of 1,140 base pairs of the cytochrome-b gene. Nodal support is represented as percent posterior probabilities (PP) obtained with MrBayes and standard bootstrap value (BP) obtained with RaxML. Nodes supported by values higher than 95% in both analyses are indicated by a filled circle. Clades showing the typical dichromatic wing pattern are marked with a pictogram. For the GenBank names of the taxa pertaining to the M. formosus-complex refer to Table 1.

Although our investigations did not reveal any craniodental characters distinguishing these taxa from all other species of Myotis, 1 phenetic character was shared by all of the studied taxa. This is the characteristic reddish or yellowish dorsal fur that strongly differentiates these bats from congeners. An additional feature that may be a synapomorphic character of Chrysopteron is the peculiar texture of the hairs, variously described as “thick and woolly” (Dobson [1871:187] for V. auratus and Yoshiyuki [1989:110] for M. tsuensis), “cottony” (Tomes [1858:84] for V. rufoniger), “woolly” (Blyth [1863:34] for K. pallida, Hill and Morris [1971:43] for M. morrisi, and Dietz et al. [2009:242] for M. emarginatus), “thick, soft and cottony” (Tomes [1858:81] for V. emarginatus), or “shaggy” (Rosevear [1965:303] for M. bocagii). This feature also is reflected in the vernacular name, “hairy bats,” frequently used for Ethiopian species (Taylor 2000), and, with the exception of 2 sister species, M. goudoti and M. anjouanensis from islands in the Indian Ocean, is typical for all Chrysopteron species investigated by us.

Chrysopteron is the earliest available name for the “Ethiopian clade.” Its type species, bartelsi Jentink, 1910 from Java, is currently considered a subspecies of M. formosus (Simmons 2005), which, in turn, is part of the “Ethiopian clade.” This renders Chrysopteron the only traditional subgenus of Myotis currently validated by molecular systematics.

With the separation of Cistugo from Myotis (Stadelmann et al. [2004b]; subsequently confirmed by Horácek et al. [2006] and Lack et al. [2010]), all sub-Saharan species of Myotis were regarded (or anticipated in the case of M. morrisi) as part of the “Ethiopian clade” by Stadelmann et al. (2004b). Our morphological examination of the holotype of M. morrisi indicated that the species has the fur characters regarded here as diagnostic of Chrysopteron, thereby justifying its inclusion in the “Ethiopian clade.” The 2 Indian Ocean insular species, M. goudoti and M. anjouanensis, which are clearly part of the Ethiopian clade in molecular reconstructions (Ruedi et al. 2013; Fig. 1), also have very strong overall reddish to orange dorsal fur. In contrast, the recently recognized species M. dieteri Happold, 2005 may be distinct from the “Ethiopian clade.” Although M. dieteri resembles M. bocagii in size, Happold (2005) noted its dorsal fur coloration as dark brown with a pale auburn tip (among other distinguishing characters), instead of reddish brown with a rufous tip, as in M. bocagii. Based on this external characteristic M. dieteri would be the only sub-Saharan Myotis that does not belong to Chrysopteron, but molecular data are needed to substantiate this interpretation.

Although Stadelmann et al. (2004b) supposed the dichromatic wing coloration evolved only once within the “Ethiopian clade,” the presence of the conspicuous black-and-orange pattern in 2 independent lineages of Chrysopteron and in several genetically unsampled species suggests that the developmental pathway may be conserved in many (if not all) species of the subgenus. It is, however, worth noting that within the subgenus this wing coloration is always accompanied by large body size and is observed only in M. welwitschii, the largest Ethiopian species (Taylor 2000; Ratcliffe 2002), and much less conspicuously so in some specimens M. tricolor from Malawi (M. Happold, pers. comm.) and in similar-sized Asian taxa. Interestingly, the distantly related K. picta (Vespertilionidae: Kerivoulinae) also shows the very same dichromatic pattern, suggesting that this peculiar coloration has appeared more than once in the evolution of bats.

Biogeographic inferences based on a likelihood model of ancestral area reconstruction (DEC model—Ree and Smith 2008) and a nearly complete, worldwide taxonomic sampling of Myotis species (Ruedi et al. 2013) indicate that species of Chrysopteron evolved after an initial range expansion of an Asian ancestor. This widespread ancestor radiated in sub-Saharan Africa to give rise to several taxa. The 2 lineages that led to the current, nonsister species M. rufoniger and M. formosus apparently resulted from 2 independent recolonizations of the Asian continent by these African forms.

Revision of Asian species of Chrysopteron

The 1st publication that grouped all of the Asian dichromatic taxa into a single species was that of Findley (1972), who provided limited justification for this action. However, his opinion was adopted by Honacki et al. (1982), Koopman (1989, 1994), and, with the exception of recognizing M. hermani as a distinct species, by Corbet and Hill (1992) and Simmons (2005). The presence of more than 1 species within M. formosus sensu lato is suggested by more recent publications employing external traits (Cheng et al. 2010), craniodental features (Heaney et al. 1998; Chou 2004), and molecular genetic studies (Jiang et al. 2010; Ruedi et al. 2013).

Our phylogenetic reconstruction (Fig. 1) confirms that specimens formerly included in the M. formosus complex are not monophyletic, but form 2 strongly supported, nonsister clades. The nomenclature used in the tree is based on our identification of voucher specimens or is inferred from distributional information and genetic distances.

One of these clades includes all representatives of M. formosus (= M. flavus sensu Jiang et al. 2010), whereas the remaining sequences, including those representing M. rufoniger (= M. watasei sensu Jiang et al. 2010), group in a distinct clade. The latter clade is clearly sister to M. welwitschii from Africa in all reconstructions (Fig. 1), but the phylogenetic position of the M. formosus clade is uncertain within a group that also contains M. emarginatus and M. tricolor with no bootstrap or posterior probability supporting either sister-group relationship. The mean Kimura 2-parameter genetic distance is about 2 orders of magnitude larger between (18.9%) than within (0.2%) these 2 clades and corresponds well to a pattern of interspecific and intraspecific comparisons, respectively (Bradley and Baker 2001).

Myotis formosus sensu lato, in fact, contains several forms that are morphologically distinct and can be found either in sympatry (e.g., in Taiwan) or in widely separated geographic localities. Having investigated specimens from the entire distribution, we recognize 6 species (M. bartelsi, M. formosus, M. hermani, M. rufoniger, M. rufopictus, and M. weberi), revise their distributions, and summarize their distinguishing characters. In view of the history of taxonomic confusion, a brief review of previous taxonomic opinions also is included.

Description

This is a large species of Chrysopteron (Table 2). Individual dorsal hairs are black basally, pale yellow distally, then darken to deep red before terminating in a black tip. The general impression of the dorsal fur is thus red tipped with black. Individual ventral hairs possess a black base, followed by either a pale yellow section that progressively darkens distally to deep red, or are otherwise entirely deep red. The ear is conspicuously edged with black and the thumb and underside of hind foot are entirely black. This combination of fur and flesh coloration is henceforth referred to as the “rufoniger-type,” this being the 1st taxon described to have these characters (Figs. 2 and 3a).

Table 2.

Selected external and craniodental measurements (in mm) of Asian Chrysopteron (Myotis) species. Values are given as mean ± SD (n ≥ 5), minimum–maximum (n). Acronyms and definitions for measurements are given in the text.

Fig. 2.

Live Myotis rufoniger from China, illustrating “M. rufoniger-type” pelage (photo: Wen-Hua Yu and Yi Wu).

Fig. 3.

Detailed view of dorsal pelage of a) Myotis rufoniger from South Korea (HNHM 2003.37.8.) and b) M. formosus from Nepal (HNHM 98.8.22.).

The skull has a definite, albeit shallow, rostral depression, and an elongated supraorbital region. The sagittal crest is prominent, and the lambdoid crests are strong. The dentition is robust. The upper canine has a wide base, and attains a height twice that of P4. P3 is fully out of line of the rest of the toothrow, has less than half the basal area of P2, and is not visible in lateral view of the skull. The mandible of the holotype is missing (Fig. 4).

Fig. 4.

Lateral view of skull and occlusal view of left upper dentition of Myotis bartelsi (holotype, MZB 10573). Scale bar in millimeters.

Taxonomic remarks

In describing C. bartelsi (separated from C. weberi from Sulawesi based on its larger size and small differences in color patterns), Jentink (1910) differentiated the cranial (flatter head) and dental (smaller middle upper premolar and number of cusps on lower incisors) features of these 2 species from another, unrelated genus containing parti-colored bats, Kerivoula, and placed both in his newly established genus Chrysopteron. He also noted that the species of Chrysopteron “have some characters in common with … Myotis” (Jentink 1910:73) and tentatively included M. formosus in the new genus.

This taxon is quite similar to M. hermani in craniodental traits. Although M. bartelsi is smaller in several mensural characters (e.g., FA, M3M3W, IOW, and CM3L; and is therefore tentatively regarded here as a separate species), in other measurements (e.g., GTL, CCW, and ZYW) it falls within the known variation of M. hermani. When more specimens and genetic information for both taxa become available, they may prove to be conspecific.

Distribution

Indonesia (Java and Bali [Fig. 5]). Kitchener and Foley (1985) recorded the species (as M. formosus) from Bali.

Fig. 5.

Distribution map of Asian Chrysopteron species: Myotis formosus (empty circles), M. rufoniger (full circles), M. rufopictus (full squares), M. hermani (full triangles), M. bartelsi (empty squares), and M. weberi (empty triangles).

Description

A medium-sized species of Chrysopteron (Table 2). Individual dorsal hairs are mid-brown at base (approximately 10% of hair length), pale yellow distally for 80–100% of hair length, or instead terminate in a mid-brown tip slightly darker than the base. Banding is sometimes not evident as the color changes gradually. The general aspect of the dorsal fur is light yellow-brown. Ventral hairs either possess a mid-brown base followed distally by pale yellow, or are entirely light yellow. The darker hair bases are not apparent from a superficial view. The ear is only faintly edged with black, and the thumb and hind foot are brown, not black. This combination of fur and flesh coloration is henceforth referred to as the “formosus-type,” this being the 1st taxon having these characters to be described (Figs. 3b and 6).

Fig. 6.

Live Myotis formosus from Taiwan, illustrating “M. formosus-type” pelage (photo: Cheng-Han Chou).

The skull has a distinctly elevated frontal region, with a globose braincase. The sagittal crest is missing or very weak and the lambdoid crests are weakly developed. The upper canine is moderately robust. P3 occupies at most half the basal area of P2, is fully or mostly out of the toothrow (sometimes missing), and is not visible in lateral view of the skull. The lower middle premolar (p3) occupies approximately half the basal area of p2 and is usually situated within the toothrow (Fig. 7).

Fig. 7.

Lateral view of skull and occlusal view of left upper and right lower dentition of Myotis formosus (Nepal, HNHM 98.8.22.). Scale bar in millimeters.

Taxonomic remarks

The very vague description of K. pallida given by Blyth (1863:34) contains a short comparison with Kerivoula picta and is limited to some external measurements.

Dobson's (1871:187) description of V. auratus “… hairs tipped with light golden brown; beneath light fawn color” defines it as having the same general color as M. formosus, with which he later (together with K. pallida) synonymized the taxon (Dobson 1878). Without referring to M. formosus, Shamel (1944:191) described M. flavus from Taiwan and characterized it as a “pale yellow bat.” He pointed out that this taxon is larger than M. rufoniger and M. watasei and that its P3 is much more reduced. He also described the characteristically differing coloration of M. watasei from a Taiwanese specimen (Shamel 1944:192). Lacking exact information on the characters of M. formosus sensu stricto, Jiang et al. (2010) mistakenly thought that M. flavus and M. formosus differ in coloration and craniodental traits (they likely regarded M. formosus as the same taxon as M. rufoniger). Although they correctly recognized that 2 separate species occur in Taiwan and in mainland Asia, these populations were subsequently associated with the wrong names. As a consequence, they named the larger, yellow species “flavus” and the smaller, reddish one “formosus,” and M. watasei was regarded as a junior synonym of M. formosus. The specific identity of the Jianxi (China) specimen (Jiang et al. 2010) is confirmed by the accompanying photographs (Jiang et al. 2010:46–47) and its Cytb sequence (Fig. 1).

Although the limited geographic coverage does not suggest major genetic subdivision within M. formosus (n = 19), based on Welch 2-sample t-tests, the Taiwanese population (M. flavus, n = 12) is significantly larger than the mainland population in the following measurements: M3M3W (t_10.34 = 3.42, P = 0.006), CM3L (t_9.26 = 3.89, P = 0.003), CP4L (t_10.89 = 3.36, P = 0.006), cm3L (t_7.62 = 3.75, P = 0.006), ML (t_7.39 = 2.92, P= 0.021) and CCW (t_8.54 = 2.41, P = 0.041). After controlling for false discoveries in multiple tests, differences in M3M3W, CM3L, CP4L, and cm3L remain significant (P = 0.023 in each case). We regard M. formosus flavus herein as a valid subspecies.

Distribution

Afghanistan, India (Jammu and Kashmir, Himachal Pradesh, Punjab, Maharashtra, Uttar Pradesh, Bihar, West Bengal, Sikkim, Assam, and Meghalaya—for details see Bates and Harrison [1997] and Mandal et al. [2000]), Nepal, China (Tibet and Jianxi), Taiwan, and Vietnam (Fig. 5). The IEBR XL-15B specimen from Thanh Hoa Province, Vietnam (an adult female, collected on 18 April 2012), represents the 1st and only record of M. formosus sensu stricto from the country.

Description

The largest species of the subgenus (Table 2). The coloration is of the “rufoniger-type” (Figs. 2 and 3a).

The skull is very robust with a shallow but distinct frontal depression, posteriorly elongated supraoccipitale, and exceptionally developed sagittal and lambdoid crests. The basal dimensions of C1 exceed those of P4, whereas P3 is minute, fully displaced lingually, and obscured in lateral view of the skull. The lower middle premolar (p3) is half the size of p2 and partly out of the toothrow (Fig. 8).

Fig. 8.

Lateral view of skull and occlusal view of left upper and right lower dentition of Myotis hermani (holotype, BMNH 23.1.2.13). Scale bar in millimeters.

Taxonomic remarks

Thomas (1923) allied M. hermani with M. weberi and M. bartelsi and noted that M. hermani is much larger size and has a well-marked sagittal crest. He also briefly discussed the earlier generic placements of M. weberi and M. bartelsi (in Kerivoula and Chrysopteron, respectively) and was the first to recognize that all 3 species belong to Myotis.

Although larger, M. hermani is evidently closely related to, and might be conspecific with M. bartelsi, in which case M. bartelsi would have priority (see also remarks under that species).

Distribution

Indonesia (Sumatra), Thailand (Songkhla [in Bumrungsri et al. 2006]), and Malaysia (Perak [in Francis 1995; Fig. 5]).

Description

On average one of the smallest of Chrysopteron species in Asia (Table 2). Coloration is of the “rufoniger-type” (Figs. 2 and 3a).

The skull has a slightly, but distinctly, elevated frontal part, and moderately strong sagittal and lambdoid crests. The dentition (including the canines) is moderately robust. The basal area of P3 is approximately two-thirds that of P2, and is usually in line and visible from outside, but rarely displaced inward. The lower middle premolar (p3) is well developed and at least two-thirds of the size of p2 basally, but often closely approaches its basal dimensions (Fig. 9).

Fig. 9.

Lateral view of skull (Taiwan, MHC 7223) and occlusal view of left upper and right lower dentition (holotype, BMNH 57.4.16.1) of Myotis rufoniger. Scale bar in millimeters.

Taxonomic remarks

Tomes (1858) provided a diagnosis for his V. rufoniger based on the color differences (ears edged with black, and dorsal and ventral hairs tipped with bright rufous) but hesitated to recognize it as a distinct species or a “variety” of V. formosus.

When Kuroda (1922) described M. tsuensis as having reddish brown fur dorsally and ventrally and compared it only with M. macrodactylus and M. nattereri bombinus, he unsurprisingly found it specifically distinct. Details provided by Kishida (1924:40) for M. watasei unambiguously define the species from Taiwan as belonging to the “rufoniger-type”: “ear red brown at base, edged with black … upper [dorsal] and under [ventral] body fur basal two-thirds brown-yellow, terminal one-third brown … feet black” (translated from Japanese).

Mori (1928:390) compared M. chofukusei, characterized by its “capucine orange” dorsal fur and ears emarginated with dark margins, with M. bechsteinii. Under the heading of formosus, Howell (1929:15) stated “the mainland bat of this rufous and black type has been described under the name rufo-niger Tomes, but I believe that the validity of the latter has not been satisfactorily established.” The geographical basis of this remark likely stemmed from his erroneous idea that M. formosus was described from Formosa (= Taiwan).

Imaizumi (1970:223), however, regarded M. tsuensis as “indistinguishable from M. chofukusei described from Korea” and gave its range as Tsushima and the Korean Peninsula. Yoshiyuki (1989) and Yoon (2010) also regarded the Korean population as belonging to the subspecies M. formosus tsuensis.

Kim et al. (2011) published the complete mitochondrial genome of a specimen of “M. formosus” from South Korea (GenBank accession number is HQ184048, not HQ184084 as published in Kim et al. 2011); genetically, this corresponds fully (100% match at Cytb, see Fig. 1) to “M. formosus” in Kawai et al. (2003) and “M. watasei” in Jiang et al. (2010) and thus represents M. rufoniger, not M. formosus sensu stricto.

Although earlier studies (Imaizumi and Yoshiyuki 1969) found no differences between specimens from Tsushima Island (M. tsuensis) and the geographically closer Korean Peninsula (M. chofukusei), in 1-way ANOVA models the Tsushima Island population (n = 4) proved to be significantly smaller than either M. chofukusei (n = 10) or the Taiwan population (M. watasei, n = 8), respectively, in the following measurements GTL (P = 0.020 and 0.006, F = 4.82), MAW (P = 0.005 and 0.010, F = 5.36), CM3L (P = 0.037 and 0.001, F = 5.39), CP4L (P = 0.005 and 0.001, F = 11.11), and CPH (P = 0.021 and 0.001, F = 9.36). After controlling for false discoveries in multiple tests, differences between M. tsuensis and M. chofukusei in MAW and CP4L remained significant (P = 0.038 in both cases). For M. tsuensis versus M. watasei, all of the above differences remained significant (P = 0.023, 0.03, 0.005, 0.005, and 0.005, respectively). Abe et al. (2008) asserted that the Tsushima population was vagrant from the Korean Peninsula and did not breed on the island although a female with young had been reported on Tsushima Island in August by Imaizumi (1970). Recent observations indicate these bats are present year-round on the island (S. Matsumura, pers. comm.). Specimens from Tsushima have not yet been included in phylogenetic analysis, but all other samples studied show almost identical Cytb sequences, indicating no major genetic subdivision within the species.

The only known Lao record of M. rufoniger is included in several phylogenetic reconstructions (Stadelmann et al. 2004b; Jiang et al. 2010; Ruedi et al. 2013; present paper). Its specific identity is supported by the color plate provided by Francis et al. (1999) and by its genetic data (e.g., approximately 0.8% sequence divergence from South Korean or Taiwanese M. rufoniger).

Distribution

North Korea, South Korea, Japan (Tsushima), China (Fujian, Jiangxi, Jilin, Shanghai, and Sichuan), Taiwan, Laos, and Vietnam (Fig. 5). The Vietnamese specimens from the provinces of Hai Duong, Bac Kan, and Ninh Binh represent the 1st published records of the species from the country.

Description

A medium-sized species of the Asian members of the subgenus (Table 2). The fur color is the “formosus-type” (Figs. 3b and 6).

The skull profile ascends almost evenly with no frontal depression. The sagittal and lambdoid crests are only moderately developed, whereas the skull is globose posteriorly. The canines are moderately strong, and the upper molars are relatively robust with developed talons. The P3s are missing from the holotype. The p3s are very small and intruded lingually half-way out of the line of the toothrow (Fig. 10).

Fig. 10.

Lateral view of skull and occlusal view of left upper and right lower dentition of Myotis rufopictus (holotype, BMNH 7.1.1.533). Scale bar in millimeters.

Taxonomic remarks

Waterhouse (1845) gave a detailed description of the fur of his new species, which was compared by him only with K. picta and separated from that species by its much larger size and differently shaped ear.

Tomes (1858) directly compared the type specimens of M. rufopictus and M. formosus and noted the larger size (despite immaturity) and missing middle premolars of the former.

Large differences in skull measurements within the Philippines (CCL 16.1–17.9, and CM3L 7.1–8.5, n = 4) are reported by Ingle and Heaney (1992) and records of a reddish form in the islands (L. Heaney, Field Museum of Natural History, pers. comm., 2013) imply the presence of a 2nd species in addition to M. rufopictus.

Distribution

Philippines (Fig. 5). An overview of the distribution and habitat of the species is given by Ong et al. (2008).

Description

A relatively large species of the subgenus (Table 2). The fur coloration is the “rufoniger-type” (Figs. 2 and 3a).

The skull has a distinct frontal depression, moderately developed sagittal and lambdoid crests, and posteriorly elongated supraoccipital region. The upper canine has a wide base but is not especially high, whereas P2 is much reduced in size, fully or partly displaced lingually, and is accordingly obscured or visible in the lateral view. The p3 occupies half the basal area of p2 at most and is situated within the toothrow (Fig. 11).

Fig. 11.

Lateral view of skull and occlusal view of left upper and right lower dentition of Myotis weberi (holotype, RMNH 35827). Scale bar in millimeters.

Taxonomic remarks

In placing M. weberi in the genus Kerivoula, Jentink (1890) was misled by the apparent similarity of their wing patterns, and within that genus he distinguished his new species by its larger size; comparisons were inappropriately confined to K. picta.

Myotis weberi is erroneously given by Yoshiyuki (1989) as the type species of Chrysopteron.

Distribution

Indonesia (Sulawesi [Fig. 5]).