Search

Additional file 7: Table S6. Significant values (P < 0.001) of TreeSAAP positive and negative z-score for amino acid physiochemical properties among radical categories 6, 7 and 8.

Additional file 2: Table S2. Length and base composition of each genomic component for species used in this study.

Additional file 5: Figure S1. Phylogenetic trees reconstructed from each single gene in MrBayes and posterior probability values are shown on the nodes.

Additional file 6: Table S5. Results of positive selection based on site model and branch-site model testing in protein-coding mitochondrial genes.

Additional file 3: Table S3. PCR primers used in this study.

Additional file 4: Table S4. Polymorphic sites of 13 PCGs, 2 rRNA genes and the combined 22 tRNA genes for species in this study, and the mutation number of each single PCGs in specieslow is shown.

Additional file 1: Table S1. Voucher numbers of samples used in this study.

Cryptic species diversity in the genus Tylonycteris Previous studies have detected high levels of genetic and karyological variation among specimens identified as either T. pachypus or T. robustula collected from different geographic locations in mainland Southeast Asia (Francis et al. 2010; Huang et al. 2014). Our COI analyses further revealed the existence of three divergent geographic haplogroups for both T. pachypus and T. robustula, for each corresponding to Sumatra, northern Indochina and southern Indochina (but also northwestern India and Peninsular Malaysia for T. robustula). Our Cytb dataset confirmed the existence of these geographic haplogroups in mainland Southeast Asia. In addition, a specimen of T. pachypus collected from Borneo and originally described as T. robustula was found to be highly divergent from the two other Indochinese haplogroups (Fig. 2). In the absence of Cytb data for Sumatran specimens, it is impossible to know whether they share the same mitochondrial lineage as those from Borneo. Taken together, our mtDNA analyses show that all haplotypes sequenced for insular Tylonycteris are very divergent from those identified in mainland Southeast Asia. However, genetic inferences based on the maternally inherited mitochondrial genes are prone to be discordant with the true evolutionary history of the taxa, owing to various evolutionary processes, such as mtDNA introgression, incomplete lineage sorting or female philopatry (Avise 2000; Ballard & Whitlock 2004; Hassanin & Ropiquet 2007; Nesi et al. 2011; Rivers et al. 2005). Here, the geographic pattern of mtDNA diversity observed for the two species of Tylonycteris could be the consequence of female philopatry, i.e., the behavior of remaining in, or returning to the natal territory. Indeed, bat species with philopatric females generally display high geographic structure when relationships are examined with maternally inherited markers, such as the mitochondrial DNA. This pattern can disappear with biparentally inherited markers when adult males are able to disperse over long distances, allowing gene flow between otherwise isolated populations (Castella et al. 2001; Hassanin et al. 2015; Hulva et al. 2010; Pereira et al. 2009; Rivers et al. 2005). Behavioral and population genetic studies in southern China have shown that T. pachypus bats are philopatric to their natal area and that philopatry is especially pronounced in females (Hua et al. 2011, 2013). Although no data are available for T. robustula, female philopatry can also be predicted for this species, because it shares similar morphological, behavioural and ecological traits with T. pachypus (Medway 1972; Medway & Marshall 1970, 1972; Zhang et al. 2007). The social organization of T. pachypus and T. robustula, combined with their fragmented habitats, is therefore expected to result in limited gene flow between populations, especially among matrilines from distant geographic localities. For both species, this prediction is corroborated by the analyses of mtDNA markers, with the identification of three divergent geographically non-overlapping haplogroups. For T. robustula, this phylogeographic pattern is also supported by the nuclear sequence data, as the two Indochinese clades were recovered monophyletic with all the three introns containing enough nucleotide variation at the intra-specific level, i.e., CHPF2, HDAC1 and TUFM (Appendix 8). By contrast, our nuclear analyses (Fig. 3; Appendices 4, 8) do not support the reciprocal monophyly of the two Indochinese clades of T. pachypus, suggesting that gene flow was maintained by male dispersal or, alternatively, that their separation was too recent to be detected with our nuclear genes. Nucleotide distances estimated from mtDNA genes between northern and southern Indochinese populations of T. robustula were more than twice those of T. pachypus (6.5% vs 2.8% in COI; 9.5% vs 2.8% in Cytb), indeed supporting a more recent divergence for the latter taxon, if we assume equal evolutionary rates. Similarly, the nuclear distances between the two Indochinese clades of T. robustula were between 0.41 and 0.56%, which is more than twice those calculated between Indochinese individuals of T. pachypus (0���0.2%; Appendix 5) and in the range of interspecific distances found in other groups of Laurasiatheria, such as fruit bats of the tribes Myonycterini (Nesi et al. 2013) and Scotonycterini (Hassanin et al. 2015), or cattle and buffalo of the tribe Bovini (Hassanin et al. 2013). Although none of the nuclear markers could be sequenced for Sumatran and Bornean Tylonycteris, their high mtDNA divergence from Indochinese populations (> 5.7 % in both COI and Cytb sequences; Appendix 5) suggests they might represent distinct lineages based on nuclear markers as well. In agreement with this view, our multivariate morphological analyses revealed that Indochinese bats of the T. pachypus complex constitute a distinct group separated from those collected on Sumatra. For the T. robustula complex, morphological overlap between haplogroups is more extensive, but pairwise comparisons of their PC mean scores support the distinctness of adjacent geographical taxa, such as Tr2 and Tr3 on the Southeast Asian mainland. The close morphological similarity among taxa of Tylonycteris suggests that they have evolved under the influences of similar and specialized habitats, i.e., woody bamboo vegetation. Molecular evidence indicates, however, that T. pachypus should be split into at least two distinct species, T. pachypus on the Sunda islands (Sumatra and/or Borneo) and T. fulvida in mainland Southeast Asia, and that T. robustula should be divided into at least three species, with T. robustula on Sumatra, T. malayana in southern and western mainland Southeast Asia, and T. tonkinensis sp. nov. in northern Indochina. The evolution of Tylonyteris spp. in Southeast Asia during the Pleistocene Given that both species complexes, here named T. pachypus s. lat. and T. robustula s. lat., are usually found in sympatry across most of their geographic ranges in Southeast Asia, they are expected to share a common phylogeographical history. Our estimates of divergence times based on mtDNA sequences suggest that the genus Tylonycteris diversified during the Pliocene epoch (Cytb: 5.92 �� 0.65 Mya; COI: 4.56 �� 0.72 Mya) (Table 1). During the Miocene and until the early Pliocene, Southeast Asia was generally covered by large tracks of rain forests as a consequence of warm and humid climatic conditions (Meijaard & Groves 2006; Morley 2000). Thus, ancestors of both Tylonycteris species complexes were presumably widely distributed across Southeast Asia during the Pliocene. Our molecular dating estimates indicate that the basal geographic splits within the two species complexes, i.e., between mainland Southeast Asia and Sumatra, took place approximately at the same time during the Early Pleistocene (between 2.70 and 1.96 Mya for T. pachypus, between 3.07 and 2.22 Mya for T. robustula; Table 1). The Pleistocene epoch is characterized by the onset of repeated cycles of cold glacial and warm interglacial periods as the results of the glaciations/deglaciations of the Northern Hemisphere, which implied contraction and expansion of rain forests in Asia (An et al. 2001; Meijaard & Groves 2006; Morley 2000). As bats of the genus Tylonycteris are highly dependent on woody bamboo vegetation for roosting, foraging and mating (Kunz 1982; Medway 1972; Medway & Marshall 1970, 1972), their Pleistocene biogeographic history was firmly constrained by the distribution of such bamboo habitats. The current distribution of woody bamboo species in Asia (Bystriakova et al. 2003b; Fig. 6) indicates that eight disjunctive biogeographic regions have higher species richness (> 5 species) for some bamboo genera: southern India (Ochlandra Thwaites), northern Myanmar (Cephalostachyum Munro), southern China (Dendrocalamus Nees and Bambusa Schreb.), Hainan Island (Bambusa), northwestern Thailand (Dendrocalamus and Gigantochloa Kurz ex Munro), Peninsular Malaysia (Gigantochloa), Sumatra and Borneo (Gigantochloa and Schizostachyum Nees). All these regions may therefore have acted as distinct bamboo forest refugia during the glacial periods of the Pleistocene (Fig. 6). Evidence for a number of these postulated glacial refugia has been reported in previous studies for many organisms, including bats (Flanders et al. 2011; Khan et al. 2010; Lin et al. 2014; Mao et al. 2013). Accordingly, we propose that the contraction of woody bamboo forests into different glacial refugia had fragmented the distribution of the Pliocene ancestors of both T. pachypus s. lat. and T. robustula s. lat. In addition, we can assume that Pleistocene glacial periods resulted in higher interspecific competition between co-distributed species of Tylonycteris, because the supply of most suitable resources was more limited in glacial bamboo forest refugia (Medway & Marshall 1970). As noted in previous studies, T. pachypus s. lat. has a more manoeuvrable flight in cluttered habitats and forages on smaller insects than T. robustula s. lat. (Zhang et al. 2005, 2007). Moreover, Medway & Marshall (1970) found that the smaller T. pachypus s. lat. can roost in the internodes with small entrance holes, which the larger T. robustula s. lat. is unable to enter. These differences suggest, therefore, that the smaller T. pachypus s. lat. have greater advantages than the larger T. robustula s. lat. in interspecific competition when natural resources are limited. Hence, isolated populations of T. robustula s. lat. may have been more exposed to bottlenecks and therefore more vulnerable to local extinction than those of co-distributed T. pachypus s. lat. During interglacial periods of the Early Pleistocene, warmer and humid conditions resulted in the expansion of woody bamboo forests, which in turn may have favored the restoration of connectivity between isolated populations of both complexes. However, the isolated populations may have been connected or not, depending on their dispersal capacity and the distances between refugia. In T. robustula s. lat., these processes may have taken longer, because of its lower population abundance (Lande & Barrowclough 1987; Shaffer 1981), and may have been prevented in cases of extinction of transitional populations (Huang et al. 2014 and references therein). This scenario is supported by the fact that T. pachypus s. lat. is usually found to be more abundant than T. robustula s. lat. in bamboo forests (Zhang et al. 2004; Medway & Marshall 1972) and by the wider geographic range of T. pachypus s. lat. (Bates et al. 2008a, 2008b; Fig. 1). Knowing this, the body size differences between the two species complexes may be the key factor explaining why the basal divergence of northern Indochinese populations occurred earlier in T. robustula s. lat. (i.e., T. tonkinensis sp. nov) than in T. pachypus s. lat., i.e., 2.97���1.70 vs 1.35���0.79 Mya (Table 1). During Pleistocene interglacials, exchanges of Tylonycteris spp. between the continent and the islands of Sundaland were probably prevented because of the long distances between the glacial forest refugia, as well as the higher sea levels (Fig. 1). Our molecular dating estimates corroborate this scenario, as continental populations of Tylonycteris spp. from Indochina and Peninsular Malaysia diverged from insular populations (Sumatra and Borneo) in the Early Pleistocene (Table 1). Implications for conservation Previous studies considered T. pachypus and T. robustula to be common species and thus classified them as Least Concern in the IUCN Red List (Bates et al. 2008a, 2008b). Since our study reveals that both species in fact represent several species with more restricted distributions, the IUCN status of the different taxa should be reassessed urgently, including that of the new species, T. tonkinensis sp. nov. In addition, our study suggests that several biogeographic regions have acted as Pleistocene glacial refugia. This information is very important for developing more effective conservation strategies, particularly given the high current rates of deforestation affecting most natural habitats in Southeast Asia (Kingston 2010; Sodhi et al. 2010; Tordoff et al. 2012)., Published as part of Vuong Tan Tu, G��bor Csorba, Manuel Ruedi & Neil M. Furey, 2017, Comparative phylogeography of bamboo bats of the genus Tylonycteris (Chiroptera, Vespertilionidae) in Southeast Asia, pp. 1-38 in European Journal of Taxonomy 274 on pages 16-19, DOI: 10.5852/ejt.2017.274, http://zenodo.org/record/291919, {"references":["Francis C. M., Borisenko A. V., Ivanova N. V., Eger J. L., Lim B. K., Guillen-Servent A., Kruskop S. V., Mackie I. & Hebert P. D. N. 2010. The role of DNA barcodes in understanding and conservation of mammal diversity in Southeast Asia. PLoS ONE 5 (9): e 12575. http: // dx. doi. org / 10.1371 / journal. pone. 0012575","Huang C., Yu W., Xu Z., Qiu Y., Chen M., Qiu B., Motokawa M., Harada M., Li Y. & Wu Y. 2014. A cryptic species of the Tylonycteris pachypus complex (Chiroptera: Vespertilionidae) and its population genetic structure in southern China and nearby regions. International Journal of Biological Sciences 10 (2): 200 - 211. http: // dx. doi. org / 10.7150 / ijbs. 7301","Avise J. C. 2000. Phylogeography: The History and Formation of Species. Harvard University Press, Cambridge, Massachusetts.","Ballard J. W. O. & Whitlock M. C. 2004. The incomplete natural history of mitochondria. Molecular Ecology 13 (4): 729 - 744. http: // dx. doi. org / 10.1046 / j. 1365 - 294 X. 2003.02063. x","Hassanin A. & Ropiquet A. 2007. Resolving a zoological mystery: the kouprey is a real species. Proceedings of the Royal Society B 274 (1627): 2849 - 2855. http: // dx. doi. org / 10.1098 / rspb. 2007.0830","Nesi N., Nakoune E., Cruaud C. & Hassanin A. 2011. DNA barcoding of African fruit bats (Mammalia, Pteropodidae). The mitochondrial genome does not provide a reliable discrimination between Epomophorus gambianus and Micropteropus pusillus. Comptes Rendus Biologies 334 (7): 544 - 554. http: // dx. doi. org / 10.1016 / j. crvi. 2011.05.003","Rivers N. M., Butlin R. K. & Altringham J. D. 2005. Genetic population structure of Natterer's bats explained by mating at swarming sites and philopatry. Molecular Ecology 14 (14): 4299 - 4312. http: // dx. doi. org / 10.1111 / j. 1365 - 294 X. 2005.02748. x","Castella V., Ruedi M. & Excoffier L. 2001. Contrasted patterns of mitochondrial and nuclear structure among nursery colonies of the bat Myotis myotis. Journal of Evolutionary Biology 14 (5): 708 - 720. http: // dx. doi. org / 10.1046 / j. 1420 - 9101.2001.00331. x","Hassanin A., Khouider S., Gembu G. - C., Goodman S. M., Kadjo B., Nesi N., Pourrut X., Nakoune E. & Bonillo C. 2015. The comparative phylogeography of fruit bats of the tribe Scotonycterini (Chiroptera, Pteropodidae) reveals cryptic species diversity related to African Pleistocene forest refugia. Comptes Rendus Biologies 338 (3): 197 - 211. http: // dx. doi. org / 10.1016 / j. crvi. 2014.12.003","Hulva P., Fornuskova A., Chudarkova A., Evin A., Allegrini B., Benda P. & Bryja J. 2010. Mechanisms of radiation in a bat group from the genus Pipistrellus inferred by phylogeography, demography and population genetics. Molecular Ecology 19 (24): 5417 - 5431. http: // dx. doi. org / 10.1111 / j. 1365 - 294 X. 2010.04899. x","Pereira M. J. R., Salgueiro P., Rodrigues L., Coelho M. M. & Palmeirim J. M. 2009. Population structure of a cave-dwelling bat, Miniopterus schreibersii: Does it reflect history and social organization? Journal of Heredity 100 (5): 533 - 544. http: // dx. doi. org / 10.1093 / jhered / esp 032","Hua P., Zhang L., Zhu G., Jones G., Zhang S. & Rossiter S. J. 2011. Hierarchical polygyny in multiparous lesser flat-headed bats. Molecular Ecology 20 (17): 3669 - 3680. http: // dx. doi. org / 10.1111 / j. 1365 - 294 X. 2011.05192. x","Hua P., Zhang L., Guo T., Flanders J. & Zhang S. 2013. Dispersal, mating events and fine-scale genetic structure in the lesser flat-headed bats. PLoS ONE 8 (1): e 54428. http: // dx. doi. org / 10.1371 / journal. pone. 0054428","Medway L. 1972. Reproductive cycles of the flat-headed bats Tylonycteris pachypus and T. robustula (Chiroptera: Vespertilioninae) in a humid equatorial environment. Zoological Journal of the Linnean Society 51 (1): 33 - 61. http: // dx. doi. org / 10.1111 / j. 1096 - 3642.1972. tb 00772. x","Medway L. & Marshall A. G. 1970. Roost-site selection among flat-headed bats (Tylonycteris spp.). Journal of Zoology 161 (2): 237 - 245. http: // dx. doi. org / 10.1111 / j. 1469 - 7998.1970. tb 02038. x","Medway L. & Marshall A. G. 1972. Roosting associations of flat-headed bats, Tylonycteris species (Chiroptera: Vespertilionidae) in Malaysia. Journal of Zoology 168 (4): 463 - 482. http: // dx. doi. org / 10.1111 / j. 1469 - 7998.1972. tb 01362. x","Zhang L., Liang B., Parsons S., Wei L. & Zhang S. 2007. Morphology, echolocation and foraging behaviour in two sympatric sibling species of bat (Tylonycteris pachypus and Tylonycteris robustula) (Chiroptera: Vespertilionidae). Journal of Zoology 271 (3): 344 - 351. http: // dx. doi. org / 10.1111 / j. 1469 - 7998.2006.00210. x","Nesi N., Kadjo B., Pourrut X., Leroy E., Pongombo Shongo C., Cruaud C. & HassaninA. 2013. Molecular systematics and phylogeography of the tribe Myonycterini (Mammalia, Pteropodidae) inferred from mitochondrial and nuclear markers. Molecular Phylogenetics and Evolution 66 (1): 126 - 137. http: // dx. doi. org / 10.1016 / j. ympev. 2012.09.028","Hassanin A., An J., Ropiquet A., Nguyen T. T. & Couloux A. 2013. Combining multiple autosomal introns for studying shallow phylogeny and taxonomy of Laurasiatherian mammals: Application to the tribe Bovini (Cetartiodactyla, Bovidae). Molecular Phylogenetics and Evolution 66 (3): 766 - 775. http: // dx. doi. org / 10.1016 / j. ympev. 2012.11.003","Meijaard E. & Groves C. P. 2006. The geography of mammals and rivers in mainland Southeast Asia. In: Lehman S. M. & Fleagle J. G. (eds) Primate Biogeography: 305 - 329. Springer, New York.","Morley R. J. 2000. Origin and Evolution of Tropical Rain Forests. John Wiley & Sons, Chichester, U. K.","An Z., Kutzbach J. E., Prell W. L. & Porter S. C. 2001. Evolution of Asian monsoons and phased uplift of the Himalaya - Tibetan plateau since Late Miocene times. Nature 411 (6833): 62 - 66. http: // dx. doi. org / 10.1038 / 35075035","Kunz T. H. 1982. Roosting ecology of bats. In: Kunz T. H. (ed.) Ecology of Bats: 1 - 55. Springer, New York.","Bystriakova N., Kapos V., Stapleton C. & Lysenko I. 2003 b. Bamboo Biodiversity: Information for Planning Conservation and Management in the Asia-Pacific Region. UNEP-WCMC, INBAR, Cambridge, U. K.","Flanders J., Wei L., Rossiter S. J. & Zhang S. 2011. Identifying the effects of the Pleistocene on the greater horseshoe bat, Rhinolophus ferrumequinum, in East Asia using ecological niche modelling and phylogenetic analyses. Journal of Biogeography 38 (3): 439 - 452. http: // dx. doi. org / 10.1111 / j. 1365 - 2699.2010.02411. x","Khan F. A. A., Solari S., Swier V. J., Larsen P. A., Abdullah M. T. & Baker R. J. 2010. Systematics of Malaysian woolly bats (Vespertilionidae: Kerivoula) inferred from mitochondrial, nuclear, karyotypic, and morphological data. Journal of Mammalogy 91 (5): 1058 - 1072. http: // dx. doi. org / 10.1644 / 09 - MAMM-A- 361.1","Lin A. Q., Csorba G., Li L. F., Jiang T. L., Lu G. J., Thong V. D., Soisook P., Sun K. P. & Feng J. 2014. Phylogeography of Hipposideros armiger (Chiroptera: Hipposideridae) in the Oriental Region: the contribution of multiple Pleistocene glacial refugia and intrinsic factors to contemporary population genetic structure. Journal of Biogeography 41 (2): 317 - 327. http: // dx. doi. org / 10.1111 / jbi. 12163","Mao X. G., He G. M., Zhang J. P., Rossiter S. J. & Zhang S. Y. 2013. Lineage divergence and historical gene flow in the Chinese horseshoe bat (Rhinolophus sinicus). PLoS ONE 8 (2): e 56786. http: // dx. doi. org / 10.1371 / journal. pone. 0056786","Zhang L. B., Jones G., Rossiter S., Ades G., Liang B. & Zhang S. Y. 2005. Diet of flat-headed bats, Tylonycteris pachypus and T. robustula, in Guangxi, South China. Journal of Mammalogy 86 (1): 61 - 66. http: // dx. doi. org / 10.1644 / 1545 - 1542 (2005) 086 2.0. CO; 2","Lande R. & Barrowclough G. 1987. Effective population size, genetic variation, and their use in population management. In: Soule M. E. (ed.) Viable Populations for Conservation: 87 - 124. Cambridge University Press, London.","Shaffer M. L. 1981. Minimum population sizes for species conservation. BioScience 31 (2): 131 - 134. http: // dx. doi. org / 10.2307 / 1308256","Zhang L. B., Liang B., Zhou S. Y., Lu L. R. & Zhang S. Y. 2004. Group structure of lesser flat-headed bat Tylonycteris pachypus and greater flat-headed bat T. robustula. Acta Zoologica Sinica 50 (3): 326 - 333.","Bates P., Francis C., Rosell-Ambal G., Heaney L., Molur S. & Srinivasulu C. 2008 a. Tylonycteris pachypus. The IUCN Red List of Threatened Species 2008: e. T 22577 A 9377719. http: // dx. doi. org / 10.2305 / IUCN. UK. 2008. RLTS. T 22577 A 9377719. en","Bates P., Francis C., Rosell-Ambal G. & Heaney L. 2008 b. Tylonycteris robustula. The IUCN Red List of Threatened Species 2008: e. T 22578 A 9378011. http: // dx. doi. org / 10.2305 / IUCN. UK. 2008. RLTS. T 22578 A 9378011. en","Kingston T. 2010. Research priorities for bat conservation in Southeast Asia: a consensus approach. Biodiversity and Conservation 19 (2): 471 - 484. http: // dx. doi. org / 10.1007 / s 10531 - 008 - 9458 - 5","Sodhi N. S., Posa M. R. C., Lee T. M., Bickford D., Koh L. P. & Brook B. W. 2010. The state and conservation of Southeast Asian biodiversity. Biodiversity and Conservation 19 (2): 317 - 328. http: // dx. doi. org / 10.1007 / s 10531 - 009 - 9607 - 5","Tordoff A. W., Bezuijen M. R., Duckworth J. W., Fellowes J. R., Koenig K., Pollard E. H. B. & Royo A. G. 2012. Ecosystem Profile: Indo-Burma Biodiversity Hotspot Indochina Region. Final Version October 2012. Critical Ecosystem Partnership Fund, Conservation International, USA."]}

Submyotodon latirostris (Kishida, 1932) Synonymy. Myotis latirostris Kishida, 1932. Type locality C Taiwan. Myotis muricola orii Kuroda, 1935. Type locality Taihezon, Taihokusiu, N Taiwan. Myotis mystacinus latirostris: Tate 1941. Name combination. Myotis mystacinus orii: Tate 1941. Name combination. Myotis mystacinus latirostris: Ellerman & Morrison-Scott 1966. Name combination. Myotis muricola latirostris: Findley 1972. Name combination. Myotis muricola orii: Corbet and Hill 1992. Name combination. Myotis latirostris: Lin et al. 2004. Name combination. Myotis muricola latirostris: Simmons 2005. Name combination. Myotis latirostris: Stadelmann et al. 2007. Name combination. Myotis latirostris: Cheng et al. 2010. Name combination. Myotis latirostris: Lack et al. 2010. Name combination. Myotis latirostris: Ruedi et al. 2013. Name combination. 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 & Sig�� 1971), whereas most species of Myotis have myotodont lower molars (i.e., where the postcristid is linking to the entoconid; Menu & Sig�� 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. Distribution. Submyotodon latirostris has been found so far only in Taiwan, where it is widespread and relatively common. 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 1000m, 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., Published as part of Ruedi, Manuel, Csorba, G��bor, Lin, Liang- Kong & Chou, Cheng-Han, 2015, Molecular phylogeny and morphological revision of Myotis bats (Chiroptera: Vespertilionidae) from Taiwan and adjacent China, pp. 301-342 in Zootaxa 3920 (1) on pages 319-321, DOI: 10.11646/zootaxa.3920.2.6, http://zenodo.org/record/287922, {"references":["Kishida, K. (1932) Notes on a Formosan whiskered bat. Lansania, 4, 153 - 160.","Kuroda, N. (1935) Formosan Mammals preserved in the collection of Marquis Yamishina. Journal of Mammalogy, 16, 277 - 291. http: // dx. doi. org / 10.2307 / 1374636","Tate, G. H. (1941) A review of the genus Myotis (Chiroptera) of Eurasia, with special reference to species occurring in the East Indies. Bulletin of the American Museum of Natural History, 78, 537 - 565.","Ellerman, J. R. & Morrison-Scott, T. C. S. (1966) Checklist of Palaearctic and Indian mammals, 1758 to 1946. 2 nd Edition. Alden Press, Oxford, 810 pp. http: // dx. doi. org / 10.1126 / science. 115.2990.431","Findley, J. S. (1972) Phenetic relationships among bats of the genus Myotis. Systematic Zoology, 21, 31 - 52. http: // dx. doi. org / 10.2307 / 2412256","Corbet, G. B. & Hill, J. E. (1992) The mammals of the Indomalayan region: a systematic review. Oxford University Press, Oxford, 488 pp. http: // dx. doi. org / 10.1017 / s 0030605300020718","Lin, L. K., Lee, L. L. & Cheng, H. C. (2004) Bats of Taiwan (2 nd ed.). National Museum of Natural Sciences, Taipei, 177 pp. [in Chinese]","Simmons, N. B. (2005) Order Chiroptera. In: Wilson, D. E. & Reeder, D. M. (Eds.), Mammal species of the world. A taxonomic and geographic reference. Johns Hopkins University Press, Washington, pp. 312 - 529.","Stadelmann, B., Kunz, T. H., Lin, L. K. & Ruedi, M. (2007) Molecular phylogeny of New World Myotis (Chiroptera, Vespertilionidae) inferred from mitochondrial and nuclear DNA genes. Molecular Phylogenetics and Evolution, 43, 32 - 48. http: // dx. doi. org / 10.1016 / j. ympev. 2006.06.019","Cheng, H. C., Fang, Y. P. & Chou, C. H. (2010) A photographic guide to the bats of Taiwan. The Agriculture Committee Taiwan Endemic Species Research Institute, Taipei, 144 pp. [in Chinese]","Lack, J. B., Roehrs, Z. P., Stanley, C. E. Jr., Ruedi, M. & Van den Bussche, R. A. (2010) Molecular phylogenetics of Myotis indicate familial-level divergence for the genus Cistugo (Chiroptera). Journal of Mammalogy, 91, 976 - 992. http: // dx. doi. org / 10.1644 / 09 - mamm-a- 192.1","Ruedi, M., Stadelmann, B., Gager, Y., Douzery, E. J. P., Francis, C. M., Lin, L. - K., Guillen-Servent, A. & Cibois, A. (2013) Molecular phylogenetic reconstructions identify East Asia as the cradle for the evolution of the cosmopolitan genus Myotis (Mammalia, Chiroptera). Molecular Phylogenetics and Evolution, 69, 437 - 449. http: // dx. doi. org / 10.1016 / j. ympev. 2013.08.011","Imaizumi, Y. (1962) List of type specimens of mammals which are present or had been present in the past in Japan. Proceedings of the Japanese Society of Systematic Zoology, 27, 6 - 10.","Kaneko, Y. & Maeda, K. (2002) A list of scientific names and the types of mammals published by Japanese researchers. Honyurui Kagaku (Mammalian Science), 42, 1 - 21.","Menu, H. & Sige, B. (1971) Nyctalodontie et myotodontie, importants caracteres de grades evolutifs chez les chiropteres entomophages. Comptes Rendus de l'Academie des Sciences de Paris, 272, 1735 - 1738.","Tiunov, M. P., Kruskop, S. V. & Feng, J. (2011) A new mouse-eared bat (Mammalia: Chiroptera, Vespertilionidae) from South China. Acta Chiropterologica, 13, 271 - 278. http: // dx. doi. org / 10.3161 / 150811011 x 624758","Borisenko, A. V., Kruskop, S. V. & Ivanova, N. V. (2008) A new mouse-eared bat (Mammalia: Chiroptera: Vespertilionidae) from Vietnam. Russian Journal of Theriology, 7, 57 - 69. http: // dx. doi. org / 10.1515 / mamm. 2001.65.1.63","Tomes, R. F. (1859) Descriptions of six hitherto undescribed species of bats. Proceedings of the Zoological Society of London, 1859, 68 - 79.","Dobson, G. E. (1871) Notes on nine new species of Indian and Indo-Chinese Vespertilionidae, with remarks on the synonymy and classification of some other species of the same family. Proceedings of the Asiatic Society of Bengal, 1871, 210 - 215. http: // dx. doi. org / 10.1126 / science. 1124187","Milne-Edwards, A. (1872) Memoire de la faune mammalogique du Tibet Oriental et principalement de la principaute de Moupin. In: Milne-Edwards, H. (Ed), Recherches pour servir a l'histoire naturelle des mammiferes comprenant des considerations sur la classification de ces animaux. G. Masson, Paris, pp. 231 - 304. http: // dx. doi. org / 10.5962 / bhl. title. 59889","Ziegler, R. (2003) Bats (Chiroptera, Mammalia) from Middle Miocene karstic fissure fillings of Petersbuch near Eichstatt, Southern Franconian Alb (Bavaria). Geobios, 36, 447 - 490."]}

Myotis fimbriatus (Peters, 1870) Synonymy. Vespertilio fimbriatus Peters, 1870. Type locality Amoy, Fujian, China. Myotis taiwanensis ��rnb��ck-Christie-Linde, 1908. Type locality Takao, Anping, Tainan, Taiwan. Myotis hirsutus Howell, 1926. Type locality Yenpingfu, Fujian, China. Myotis fimbriatus: Tate 1941. First use of current name combination. Myotis taiwanensis: Lin et al. 2004. Name combination. Myotis adversus taiwanensis: Simmons 2005. Name combination. Myotis taiwanensis: Cheng et al. 2010. Name combination. Myotis taiwanensis: Han et al. 2010. Name combination. Myotis taiwanensis: Ruedi et al. 2013. Name combination. Taxonomic remarks. Peters (1870) described two new Chinese species of Vespertilio (now Myotis) living in sympatry in Amoy (now Xiamen, Fujian Province). He named the smaller, rarer species with woolly pelage V. laniger (see below for its description), whereas the larger, more common species was named V. fimbriatus, and was characterized by the ���margins of interfemoral and lumbar membranes ciliated���; he also mentioned that the wing membrane extended to the middle of the metatarsus, but closer examination of the type specimen (and all recent material) suggest that it rather connects to the ankle, close to the proximal end of the metatarsus. In addition, Peters (1870) also mentioned that the second upper premolar was situated at the inner side of the third, suggesting that it might be displaced inwards from the toothrow, which is not the case on the type and all other referred material. Based on a new series of specimens collected in the same area in Fujian, Howell (1926) described another two species, a smaller form with wing membranes attaching to the base of toes (= M. sowerbyi), and a larger one, named M. hirsutus. The latter species is characterized by very hairy feet and membranes, and resembles the European M. capaccinii, as Howell suggested. However, he overlooked Peters��� (1870) description of M. fimbriatus, which was based on bats likely caught in the same cave (Allen 1938). Direct comparisons of the type material indicate that fimbriatus and hirsutus indeed represent the same species, as suggested by Allen (1938) and both are very similar to the type of taiwanensis (��rnb��ck-Christie-Linde 1908). External dimensions of these three taxa are also very similar, but the latter taxon has a slightly larger skull and a more globose braincase than the type of fimbriatus or hirsutus, and thus should be retained as a valid subspecies (i.e., M. fimbriatus taiwanensis) endemic to Taiwan. Specimens referred to M. fimbriatus s.l. from Taiwan and mainland China are genetically very similar or identical (Fig. 3), and confirm that they belong to the same species. Contrary to previous suggestions based on phenetic comparisons (Findley 1972; Corbet & Hill 1992), molecular reconstructions (Han et al. 2010; Ruedi et al. 2013) indicate that M. fimbriatus is not particularly related to M. capaccinii, nor to M. adversus, but is comprised in Clade X (Fig. 3). This East Asian clade includes other large-footed, trawling bats such as M. pilosus (a senior synonym of M. ricketti), M. cf. fimbriatus, M. petax and M. macrodactylus (Fig. 3). Records of M. fimbriatus from Yunnan (Zhang et al. 2009; Ruedi et al. 2013) refer to specimens that also fit the general morphological characteristics of fimbriatus, but they are larger (FA 42.9 mm; GLS 16.1 mm; CM 3 5.9 mm) and are genetically divergent (Ruedi et al. 2013). They might represent a distinct species in this group and should best be referred as M. cf. fimbriatus (as in Clade X of figure 3) until more detailed taxonomic comparisons are made. 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). 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., Published as part of Ruedi, Manuel, Csorba, G��bor, Lin, Liang- Kong & Chou, Cheng-Han, 2015, Molecular phylogeny and morphological revision of Myotis bats (Chiroptera: Vespertilionidae) from Taiwan and adjacent China, pp. 301-342 in Zootaxa 3920 (1) on pages 321-322, DOI: 10.11646/zootaxa.3920.2.6, http://zenodo.org/record/287922, {"references":["Peters, W. (1870) Catalogue of Mammals of China. In: Swinhoe, R. (Ed.), Proceedings of the Zoological Society of London, 3, 615 - 653.","Arnback-Christie-Linde, A. (1908) A collection of bats from Formosa. Annals and Magazine of Natural History, 2, 235 - 238. http: // dx. doi. org / 10.1080 / 00222930808692477","Howell, A. B. (1926) Three new mammals from China. Proceedings of the Biological Society of Washington, 39, 137 - 140.","Tate, G. H. (1941) A review of the genus Myotis (Chiroptera) of Eurasia, with special reference to species occurring in the East Indies. Bulletin of the American Museum of Natural History, 78, 537 - 565.","Lin, L. K., Lee, L. L. & Cheng, H. C. (2004) Bats of Taiwan (2 nd ed.). National Museum of Natural Sciences, Taipei, 177 pp. [in Chinese]","Simmons, N. B. (2005) Order Chiroptera. In: Wilson, D. E. & Reeder, D. M. (Eds.), Mammal species of the world. A taxonomic and geographic reference. Johns Hopkins University Press, Washington, pp. 312 - 529.","Cheng, H. C., Fang, Y. P. & Chou, C. H. (2010) A photographic guide to the bats of Taiwan. The Agriculture Committee Taiwan Endemic Species Research Institute, Taipei, 144 pp. [in Chinese]","Han, N. J., Zhang, J. S., Reardon, T., Lin, L. K., Zhang, J. P. & Zhang, S. Y. (2010) Revalidation of Myotis taiwanensis Arnback- Christie-Linde 1908 and its molecular relationship with M. adversus (Horsfield 1824) (Vespertilionidae, Chiroptera). Acta Chiropterologica, 12, 449 - 456. http: // dx. doi. org / 10.3161 / 150811010 x 538016","Ruedi, M., Stadelmann, B., Gager, Y., Douzery, E. J. P., Francis, C. M., Lin, L. - K., Guillen-Servent, A. & Cibois, A. (2013) Molecular phylogenetic reconstructions identify East Asia as the cradle for the evolution of the cosmopolitan genus Myotis (Mammalia, Chiroptera). Molecular Phylogenetics and Evolution, 69, 437 - 449. http: // dx. doi. org / 10.1016 / j. ympev. 2013.08.011","Allen, G. M. (1938) The mammals of China and Mongolia. Vol. 11. Part. 1. The American Museum of Natural History, New York, 620 pp.","Findley, J. S. (1972) Phenetic relationships among bats of the genus Myotis. Systematic Zoology, 21, 31 - 52. http: // dx. doi. org / 10.2307 / 2412256","Corbet, G. B. & Hill, J. E. (1992) The mammals of the Indomalayan region: a systematic review. Oxford University Press, Oxford, 488 pp. http: // dx. doi. org / 10.1017 / s 0030605300020718","Zhang, Z. Z., Tan, X. Y., Sun, K. P., Liu, S., Xu, L. J. & Feng, J. (2009) Molecular systematics of the Chinese Myotis (Chiroptera, Vespertilionidae) inferred from cytochrome-b sequences. Mammalia, 73, 323 - 330. http: // dx. doi. org / 10.1515 / mamm. 2009.058"]}

Key to the Myotinae from Taiwan and adjacent mainland China 1 Size large, forearm 45 mm or more, upper toothrow (CM 3) 7 mm or more..................................... 2 - Smaller size, forearm less than 45 mm, upper toothrow less than 7 mm........................................ 8 2 Wing membranes distinctly particolored, rufous and blackish............................................... 3 - Wing membranes unicolored, brownish................................................................ 4 3 Ears conspicuously tipped and marginated with black; feet and thumbs entirely black; ventral fur tricolored (dark grey, pale rufous and bright reddish tips); second upper premolar in toothrow; breadth across molars ��� 7.6 mm............................................................................................ M. rufoniger (incl. watasei) - Ears only faintly or not marginated with black; feet and thumbs essentially brown; ventral fur yellow, without rusty tips; second upper premolar crowded, displaced inwards from toothrow; breadth across molars> 7.5 mm.............................................................................................. M. formosus (incl. flavus) 4 Wing membrane attached to tibia; feet greatly enlarged, about size of tibia, with strongly curved claws............................................................................................... M. pilosus (= ricketti) - Wing membrane attached to ankle or metatarsus; feet not enlarged, less than half tibia length, claws not strongly curved. 5 5 Very large, forearm ��� 63 mm; belly dark greyish; massive teeth..................................... M. chinensis - Smaller, forearm M. altarium - Ears shorter, barely reaching nose tip.................................................................. 7 7 Medium size, forearm about 45 mm; long, straight tragus reaching beyond ear notch; distinctive fringe of stiff hairs along margin of uropatagium.................................................................... M. pequinius - Larger size, forearm about 60 mm; shorter tragus reaching half ear size; no stiff hairs at uropatagium; colour pale brown above............................................................................................................................................................................... M. blythii ancilla 8 Wing membrane attached to ankle or base of metatarsus; feet larger than half of tibia length; feet and uropatagium hairy along tibia....................................................................................... 9 - Wing membrane attached near base of toe; smaller feet about half or less tibia length........................... 10 9 Size small, forearm less than 38 mm; ears relatively long, reaching beyond (about 5 mm) tip of nose; dorsal pelage greyishbrown, long and soft; upper canines weak, exceeding slightly the larger premolar in height.................. M. laniger - Larger size, forearm 38 mm or more; ears relatively short, not reaching tip of nose; dorsal pelage short, richer brown; upper canines strong, much larger than premolars.................................... M. fimbriatus (incl. taiwanensis) 10 Size very small, forearm less than 35; tibia 15 mm or less................................................. 11 - Larger size, forearm more than 34; longer tibia, 15 mm or more............................................ 13 11 Second upper premolars crowded, displaced lingually from toothrow; canines strong; braincase globose; feet (incl. claw) 8 mm or more; lower molars myotodont.......................................................... M. davidii - Second upper premolars not crowded, visible laterally; small feet (7 mm or less); first and second lower molars nyctalodont.............................................................................................. 12 12 Ears deeply notched; paraconule of third premolar reaching height of second premolar; canines longer than premolars; braincase flattened, distinctively raised behind.................................................. S. latirostris - Ears without significant notch; small or no paraconule at third premolar; canines weak comparable in height to third premolars; braincase flattened, but not raised behind.............................................. M. alticraniatus 1 [1 the slightly larger Myotis badius, recently described from Yunnan (Tiunov et al. 2011) has also nyctalodont lower molars, but would be distinguished by a more domed, globose braincase] 13 Second upper premolar visible in lateral view; ears long, reaching nearly nose tip, without notable notch; rostrum not shortened; globose profile of braincase; forearm 38 mm or smaller.............................................. 14 - Second upper premolar displaced inwards, not visible laterally; ears short, with a distinct notch on rear edge; short rostrum with elevated forehead and angular braincase; forearm more than 38 mm..................................... 16 14 Upper canines similar in height to the larger premolar; calcar short, smaller than half length of rear edge of uropatagium; dorsal fur with frosted appearance................................................................... 15 - Strong upper canines, much higher than premolars; calcar long, about �� of rear edge of uropatagium; dorsal fur dark brown without frosted appearance................................................................. M. horsfieldii 15 Upper and lower canines weak; inflated braincase with frontal part rising abruptly; distal part of rostrum rounded; endemic to SE China............................................................................. M. sowerbyi - Stronger canines; braincase rising gently from the rostrum to the apex; distal part of the rostrum more pointed; endemic to Taiwan........................................................................... M. secundus sp. n. 16 Pelage dark brown, without frosted appearance; tail as long as head and body length; feet much less than half tibia length; tragus nearly reaching ear notch; forearm less than 42 mm........................................... M. frater - Pelage rich cinnamon red with golden tips of dorsal hairs giving a frosted appearance; tail clearly shorter than head and body length; feet about half tibia length; tragus clearly shorter than ear notch; forearm about 42 mm...... M. soror sp. n., Published as part of Ruedi, Manuel, Csorba, G��bor, Lin, Liang- Kong & Chou, Cheng-Han, 2015, Molecular phylogeny and morphological revision of Myotis bats (Chiroptera: Vespertilionidae) from Taiwan and adjacent China, pp. 301-342 in Zootaxa 3920 (1) on page 337, DOI: 10.11646/zootaxa.3920.2.6, http://zenodo.org/record/287922, {"references":["Tiunov, M. P., Kruskop, S. V. & Feng, J. (2011) A new mouse-eared bat (Mammalia: Chiroptera, Vespertilionidae) from South China. Acta Chiropterologica, 13, 271 - 278. http: // dx. doi. org / 10.3161 / 150811011 x 624758"]}

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: Top��l 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 (Top��l 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 4 c, 5 b). 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 (Top��l 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. 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., Published as part of Ruedi, Manuel, Csorba, G��bor, Lin, Liang- Kong & Chou, Cheng-Han, 2015, Molecular phylogeny and morphological revision of Myotis bats (Chiroptera: Vespertilionidae) from Taiwan and adjacent China, pp. 301-342 in Zootaxa 3920 (1) on pages 322-324, DOI: 10.11646/zootaxa.3920.2.6, http://zenodo.org/record/287922, {"references":["Peters, W. (1870) Catalogue of Mammals of China. In: Swinhoe, R. (Ed.), Proceedings of the Zoological Society of London, 3, 615 - 653.","Dobson, G. E. (1878) Catalogue of the Chiroptera in the collection of the British Museum. Taylor & Francis, London, 567 pp. http: // dx. doi. org / 10.5962 / bhl. title. 55341","Allen, G. M. (1938) The mammals of China and Mongolia. Vol. 11. Part. 1. The American Museum of Natural History, New York, 620 pp.","Tate, G. H. (1941) A review of the genus Myotis (Chiroptera) of Eurasia, with special reference to species occurring in the East Indies. Bulletin of the American Museum of Natural History, 78, 537 - 565.","Bates, P. J. J. & Harrison, D. L. (1997) Bats of the Indian Subcontinent (Vol. 43). Harrison Zoological Museum, Sevenoaks, 258 pp.","Topal, G. (1997) A new mouse-eared bat species, from Nepal, with statistical analyses of some other species of subgenus Leuconoe (Chiroptera, Vespertilionidae). Acta Zoologica Academiae Scientiarum Hungaricae, 43, 375 - 402.","Lin, L. K., Lee, L. L. & Cheng, H. C. (1997) Bats of Taiwan (1 st ed.). National Museum of Natural Sciences, Taipei, 165 pp. [in Chinese]","Bates, P. J. J., Hendrichsen, D. K., Walston, J. L. & Hayes, B. (1999) A review of the mouse-eared bats (Chiroptera: Vespertilionidae: Myotis) from Vietnam with significant new records. Acta Chiropterologica, 1, 47 - 74.","Lin, L. K., Lee, L. L. & Cheng, H. C. (2004) Bats of Taiwan (2 nd ed.). National Museum of Natural Sciences, Taipei, 177 pp. [in Chinese]","Borisenko, A. V., Kruskop, S. V. & Ivanova, N. V. (2008) A new mouse-eared bat (Mammalia: Chiroptera: Vespertilionidae) from Vietnam. Russian Journal of Theriology, 7, 57 - 69. http: // dx. doi. org / 10.1515 / mamm. 2001.65.1.63","Cheng, H. C., Fang, Y. P. & Chou, C. H. (2010) A photographic guide to the bats of Taiwan. The Agriculture Committee Taiwan Endemic Species Research Institute, Taipei, 144 pp. [in Chinese]","Ruedi, M., Stadelmann, B., Gager, Y., Douzery, E. J. P., Francis, C. M., Lin, L. - K., Guillen-Servent, A. & Cibois, A. (2013) Molecular phylogenetic reconstructions identify East Asia as the cradle for the evolution of the cosmopolitan genus Myotis (Mammalia, Chiroptera). Molecular Phylogenetics and Evolution, 69, 437 - 449. http: // dx. doi. org / 10.1016 / j. ympev. 2013.08.011","Shamel, H. H. (1942) A collection of bats from Thailand (Siam). Journal of Mammalogy, 23, 317 - 328. http: // dx. doi. org / 10.2307 / 1375002","Corbet, G. B. & Hill, J. E. (1992) The mammals of the Indomalayan region: a systematic review. Oxford University Press, Oxford, 488 pp. http: // dx. doi. org / 10.1017 / s 0030605300020718","Simmons, N. B. (2005) Order Chiroptera. In: Wilson, D. E. & Reeder, D. M. (Eds.), Mammal species of the world. A taxonomic and geographic reference. Johns Hopkins University Press, Washington, pp. 312 - 529.","Kruskop, S. V. & Tsytsulina, K. A. (2001) A new big-footed mouse-eared bat Myotis annamiticus sp. nov. (Vespertilionidae, Chiroptera) from Vietnam. Mammalia, 65, 63 - 72. http: // dx. doi. org / 10.1515 / mamm. 2001.65.1.63","Zhang, Z. Z., Tan, X. Y., Sun, K. P., Liu, S., Xu, L. J. & Feng, J. (2009) Molecular systematics of the Chinese Myotis (Chiroptera, Vespertilionidae) inferred from cytochrome-b sequences. Mammalia, 73, 323 - 330. http: // dx. doi. org / 10.1515 / mamm. 2009.058","Chou, C. - H. (2004) Taxonomic status of the genus Myotis (Chiroptera: Vespertilionidae) in Taiwan. Department of Wildlife Ecology, Tunghai University, Tunghai, 115 pp. [in Chinese]","Allen, G. M. (1923) New Chinese bats. American Museum Novitates, 85, 1 - 8."]}

Myotis formosus (Hodgson, 1835) Synonymy. Vespertilio formosa Hodgson, 1835. Type locality Kathmandu Valley, Nepal. Kerivoula pallida Blyth, 1863. Type locality Chaibassa, Orissa, India. Vespertilio auratus Dobson, 1871. Type locality Darjeeling, West Bengal, India. Vespertilio dobsoni Anderson, 1881. Type locality Purnia, Bihar, India. Not V. dobsoni Trouessart, 1878. Vespertilio Andersoni Trouessart, 1897. Replacement name for Vespertilio dobsoni Anderson, 1881, preoccupied by V. dobsoni Trouessart, 1878. Myotis formosus: Tate 1941. First use of current name combination. Myotis flavus Shamel, 1944. Type locality Yuanli, Miaoli, Taiwan. Myotis formosus formosus: Koopman 1994. Name combination. Myotis formosus watasei: Lin et al. 1997. Name combination. Myotis flavus: Lin et al. 2004. Name combination. Myotis formosus flavus: Cheng et al. 2010. Name combination. Myotis flavus: Jiang et al. 2010. Name combination. Myotis formosus flavus: Ruedi et al. 2013. Name combination. Taxonomic remarks. A full taxonomic treatment of species from the subgenus Chrysopteron (to which M. formosus belongs) has been published recently (Csorba et al. 2014). To avoid repetitions, we outline hereafter only the main distinguishing characters of the two species living in Taiwan and China (M. formosus and M. rufoniger). All species classified in this subgenus are phylogenetically part of the Ethiopian Clade (Csorba et al. 2014) and constitute a robust, monophyletic clade (Fig. 3). Following Csorba et al. (2014), the population of Taiwan (being significantly larger than their continental counterparts) is considered here as a distinct subspecies, M. formosus flavus. 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. 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., Published as part of Ruedi, Manuel, Csorba, G��bor, Lin, Liang- Kong & Chou, Cheng-Han, 2015, Molecular phylogeny and morphological revision of Myotis bats (Chiroptera: Vespertilionidae) from Taiwan and adjacent China, pp. 301-342 in Zootaxa 3920 (1) on pages 334-335, DOI: 10.11646/zootaxa.3920.2.6, http://zenodo.org/record/287922, {"references":["Hodgson, B. H. (1835) Synopsis of the Vespertilionidae of Nepal. Journal of the Asiatic Society of Bengal, 4, 699 - 701.","Blyth, E. (1863) Catalogue of the Mammalia in the Museum Asiatic Society. Savielle and Cronenburgh, London, 187 pp.","Dobson, G. E. (1871) Notes on nine new species of Indian and Indo-Chinese Vespertilionidae, with remarks on the synonymy and classification of some other species of the same family. Proceedings of the Asiatic Society of Bengal, 1871, 210 - 215. http: // dx. doi. org / 10.1126 / science. 1124187","Anderson, J. (1881) Catalogue of Mammalia in the Indian Museum, Calcutta. Indian Museum, Calcutta, 223 pp.","Trouessart, E. - L. (1878) Catalogue des mammiferes vivants et fossiles. Chiroptera. Revue et magazine de zoologie pure et appliquee, 3, 200 - 254.","Trouessart, E. - L. (1897) Catalogus mammalium tam viventium quam fossilium. Fasciculus I. - II. Friedlander and Sohn, Berlin, 1469 pp.","Tate, G. H. (1941) A review of the genus Myotis (Chiroptera) of Eurasia, with special reference to species occurring in the East Indies. Bulletin of the American Museum of Natural History, 78, 537 - 565.","Shamel, H. H. (1944) A new Myotis from Formosa. Journal of Mammalogy, 25, 191 - 192.","Koopman, K. F. (1994) Chiroptera: Systematics. In: Niethammer, J., Schliemann, H. & Starck, D. (Eds.), Handbuch der Zoologie. de Gruyter, Berlin, pp. 100 - 109.","Lin, L. K., Lee, L. L. & Cheng, H. C. (1997) Bats of Taiwan (1 st ed.). National Museum of Natural Sciences, Taipei, 165 pp. [in Chinese]","Lin, L. K., Lee, L. L. & Cheng, H. C. (2004) Bats of Taiwan (2 nd ed.). National Museum of Natural Sciences, Taipei, 177 pp. [in Chinese]","Cheng, H. C., Fang, Y. P. & Chou, C. H. (2010) A photographic guide to the bats of Taiwan. The Agriculture Committee Taiwan Endemic Species Research Institute, Taipei, 144 pp. [in Chinese]","Jiang, T. L., Sun, K. P., Chou, C. H., Zhang, Z. Z. & Feng, J. (2010) First record of Myotis flavus (Chiroptera: Vespertilionidae) from mainland China and a reassessment of its taxonomic status. Zootaxa, 2414, 41 - 51.","Ruedi, M., Stadelmann, B., Gager, Y., Douzery, E. J. P., Francis, C. M., Lin, L. - K., Guillen-Servent, A. & Cibois, A. (2013) Molecular phylogenetic reconstructions identify East Asia as the cradle for the evolution of the cosmopolitan genus Myotis (Mammalia, Chiroptera). Molecular Phylogenetics and Evolution, 69, 437 - 449. http: // dx. doi. org / 10.1016 / j. ympev. 2013.08.011","Csorba, G., Chou, C. - H., Ruedi, M., Gorfol, T., Motokawa, M., Wiantoro, S., Thong, V. D., Son, N. T., Lin, L. - K. & Furey, N. (2014) The reds and the yellows: a review of Asian Chrysopteron Jentink, 1910 (Chiroptera: Vespertilionidae: Myotis). Journal of Mammalogy, 95, 663 - 678. http: // dx. doi. org / 10.1644 / 13 - mamm-a- 200","Swinhoe, R. (1870) Catalogue of the mammals of China (south of the River Yangtsze) and of the island of Formosa. Proceedings of the Zoological Society of London, 42, 615 - 652."]}

Glischropus aquilus n. sp. Figures 1���4; Tables 1. Synonymy. Glischropus sp.: Huang et al. (2014). Holotype. MZB 35030 (field number # JH 20110815.2), adult male, in alcohol, skull removed. Collected by Joe Chun-Chia Huang on 15 August 2011. Type locality. Sukabanjar village, Lampung, Sumatra, Indonesia, 4 �� 56 ���S, 103 �� 52 ���E, 768 m a.s.l. External measurements of the holotype (in mm): FA= 32.1, EAR = 11.0, HB= 35.4, TAIL = 38.8, TIB= 15.1, TH= 5.6, HF= 6.2, body mass (g)= 4.8. Etymology. The specific epithet /a.kvi.lus/ (meaning dark-coloured in English) refers to the blackish ears and generally darker pelage of the new species relative to its congeners. The proposed English name is Dark Thickthumbed bat. Diagnosis. A rather large representative of the genus (FA over 32 mm), with dark brown fur, elevated frontal part, globose braincase and gradually narrowing interorbital region. Description. Forearm length= 32.1 mm (Table 1); ears are blackish, wide, and broadly rounded; tragus Pipistrellus -like, relatively narrow and bluntly pointed (Fig. 1). The dorsal pelage is dark brown with no reddish hints; ventrally lighter, medium brown. The individual hairs on the back are mostly unicoloured dark brown with only the tips a lighter brown; the hairs on the belly have two bands, the basal 2 / 3 being dark brown, the distal portion medium brown. The plagiopatagium is attached to the base of the toe. The calcar is short, extends to less than half of the free edge of the uropatagium; the lobe on the calcar is well-developed, elongated, and with a supporting median cartilage. The thumb has a large pinkish pad, characteristic of the genus, oval in outline and 3 mm in length. The sole of the foot is pink and fleshy. The penis is dorsoventrally flattened, 5.2 mm in length; the proximal half is practically naked, whereas the distal half is strongly pilose, with stiff, whitish hairs on the dorsal surface and around the glans (Fig. 2). The skull has an elevated frontal region and a relatively globose braincase (Fig. 3). The narial emargination is wide and U-shaped; the sagittal crest obsolete, the lambdoid crests are moderately developed. The zygoma is thin and nearly straight with no dorsal eminence. The basioccipital pits are almost imperceptible. The tips of the four upper incisors are situated in a nearly straight line; the concavity of the second upper incisor (I 3) is turned outwards. The first incisor (I 2) is bifid, I 3 reaches half the height of I 2. The main cusp of I 3 is much longer than the faint secondary cusplet of the same tooth and its tip is directed downwards. Basal dimension of I 3 equals that of I 2. The first upper premolar (P 2) is basally as large as I 2, partly intruded from the toothrow and visible in the lateral view; its cusp reaches well beyond the cingulum of the posterior premolar. The upper and lower molars show no specific modifications and the lower molars are nyctalodont (Fig. 4). Comparisons. Glischropus aquilus n. sp. is readily distinguishable externally from all other Glischropus species by the general impression of the dorsal fur being dark brown instead of reddish-yellow and the nearly black ear and tragus (vs. brown in all other species of the genus) (Fig. 1). Beside the external features, G. aquilus n. sp. is also clearly different from both G. tylopus and G. javanus in skull proportions, having a deeper rostrum, more elevated frontal part and higher occipital region (Fig. 3). G. aquilus n. sp. is larger in FA, M 3 M 3 W, RW and IOW than any G. tylopus specimen investigated. G. aquilus n. sp. was also different from all investigated Sumatran G. tylopus (see Comparative material) having a wide ���U��� shaped narial emargination (vs. elongated and posteriorly narrowing). Although in skull shape G. aquilus n. sp. is more similar to G. bucephalus and except the PDW value (which is smaller in all investigated G. bucephalus specimens) its craniodental measurements fall within the range of the latter species, its braincase in general is less swollen, interorbital region gradually narrowing (vs. abruptly narrowing in G. bucephalus); and I 3 nearly equals I 2 in basal dimensions (vs. much smaller). Multivariate analyses. In addition to the morphological comparisons a Principal Component Analysis was also performed. The 49 specimens clustered into two main groups: G. tylopus on the left and G. bucephalus on the right of the PC 1 axis (Fig. 5). The single G. j a v a n u s type fell near the group of G. t y l o p u s, whereas G. aquilus n. sp. is far from all other Glischropus specimens. The PC 1 axis is a size axis and shows the wideness of the skull; it accounted for a relatively large (71.49 %) proportion of the total variation, whereas PC 2 is responsible for 19.40 %of variation and indicates a shape factor dominated by PDW (Table 2). Phylogenetic reconstruction. Because G. javanus is represented only by the holotype, the phylogenetic analysis did not include this species. The analyzed Glischropus cytb sequences grouped into a monophyletic clade and G. aquilus n. sp. is clearly separated from other congeners (Fig. 6). The genetic distance between G. aquilus n. sp. and other Glischropus species���including G. bucephalus paratypes and a G. tylopus specimen collected close to the type locality���is between 12.1���14.6 %, which clearly supports that G. aquilus n. sp. is a separate species. The two G. bucephalus paratypes differed only in a few nucleotides, but the two G. tylopus differed considerably, by 5.2 % from each other (Table 3). Ecological notes. The only known specimen was caught along a trail in a secondary forest with a four-bank harp-trap which was set near a bamboo stand and a small stream. Another 14 species were recorded in the forest (including Tylonycteris robustula another bamboo specialist) and an additional eight species from a plantation nearby; hence, 23 bat species were confirmed from the site (Huang et al. 2014)., Published as part of Csorba, G��bor, G��rf��l, Tam��s, Wiantoro, Sigit, Kingston, Tigga, Bates, Paul J. J. & Huang, Joe Chun-Chia, 2015, Thumb-pads up ��� a new species of thick-thumbed bat from Sumatra (Chiroptera: Vespertilionidae: Glischropus), pp. 267-278 in Zootaxa 3980 (2) on pages 269-273, DOI: 10.11646/zootaxa.3980.2.7, http://zenodo.org/record/234638, {"references":["Huang, J. C. - C., Jazdzyk, E. L., Nusalawo, M., Maryanto, I., Maharadatunkamsi, Wiantoro, S. & Kingston, T. (2014) A recent bat survey reveals Bukit Barisan Selatan Landscape as a chiropteran diversity hotspot in Sumatra. Acta Chiropterologica, 16, 413 - 449. http: // dx. doi. org / 10.3161 / 150811014 X 687369"]}

Myotis frater Allen, 1923 Synonymy. Myotis frater Allen, 1923. Type locality Yenping, Fukien Province, China. Myotis sp. 3: Lin et al. 2004. Vernacular, unavailable name. Myotis sp. 3: Cheng et al. 2010. Vernacular, unavailable name. Myotis sp. 3: Ruedi et al. 2013. Vernacular, unavailable name. Taxonomic remarks. The angular skull, short toothrow and displaced upper middle premolar differentiate the taxa of the frater complex from other Myotis species (Tsytsulina & Strelkov 2001), but this group is morphologically a polytypic species complex comprising various forms, with variation in skull profiles and in extension of displacement of the second upper premolar. Tsytsulina and Strelkov (2001) separated the larger, pale desert form isolated in western Asia as a distinct species, M. bucharensis Kuzyakin, 1950. They commented on the taxonomy of the other, more boreal forms (longicaudatus, kaguyae and eniseensis), but did not compare them with material from the type locality in SE China, where M. frater s. s. was originally described by Allen (1923). Morphological comparisons of skulls (Figs. 9 b, 9 c) indicate that M. frater from Taiwan are very similar to the type of frater from adjacent China and thus represent the same species. Another specimen, described here as a new species in Taiwan, M. soror sp. n., also belongs to the frater species complex, but differs in external and cranial morphology (Figs. 8 a, 8 b, 9) and is genetically divergent from its congeners (Fig. 3; Table 5). Previous molecular analyses (Kruskop et al. 2012) further showed that the subspecies eniseensis and longicaudatus, although geographically isolated and morphologically separable, differ only slightly at the mitochondrial COI gene (about 2 %), suggesting that both belong to a single boreal species. Our phylogenetic reconstructions (Fig. 3) further indicate that specimens from Manchuria (i.e., representing longicaudatus) and Hokkaido (i.e., representing kaguyae) also differ minimally from each other at another mitochondrial gene (less than 1 % at Cyt b), again suggesting that all three taxa living in boreal and temperate forests are subspecies of a single species. These molecular data further suggest that bats living in more tropical regions (e.g., those sampled in Taiwan and nearby China) differ markedly in their Cyt b gene sequences (> 13 %) and in the size of the baculum (Tsytsulina & Strelkov 2001), and thus are clearly distinct from all other boreal taxa (Fig. 3). The genetic differences are comparable to those separating the two sympatric species in Taiwan (M. soror sp. n. and M. frater, 11.4 %; Table 5). Besides M. soror sp. n., we thus recommend treating M. frater s. s. from Taiwan and adjacent China as a species distinct from the other, more temperate taxa in this group. According to this new taxonomic arrangement and to designate the boreal-temperate species, M. longicaudatus Ognev, 1927 is the oldest available name, which has priority over kaguyae Imaizumi, 1956 and eniseensis Tsytsulina and Strelkov, 2001. Owing to relatively important morphological variability of this taxon in the Japanese archipelago, more comprehensive genetic sampling across Hokkaido and Honshu is still needed to confirm that a single taxon is living throughout this archipelago. Distribution. The known distribution of M. frater s. s. as understood here is restricted to Fujian Province (formerly Fukien) in eastern China, and Taiwan. 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)., Published as part of Ruedi, Manuel, Csorba, G��bor, Lin, Liang- Kong & Chou, Cheng-Han, 2015, Molecular phylogeny and morphological revision of Myotis bats (Chiroptera: Vespertilionidae) from Taiwan and adjacent China, pp. 301-342 in Zootaxa 3920 (1) on pages 331-334, DOI: 10.11646/zootaxa.3920.2.6, http://zenodo.org/record/287922, {"references":["Allen, G. M. (1923) New Chinese bats. American Museum Novitates, 85, 1 - 8.","Lin, L. K., Lee, L. L. & Cheng, H. C. (2004) Bats of Taiwan (2 nd ed.). National Museum of Natural Sciences, Taipei, 177 pp. [in Chinese]","Cheng, H. C., Fang, Y. P. & Chou, C. H. (2010) A photographic guide to the bats of Taiwan. The Agriculture Committee Taiwan Endemic Species Research Institute, Taipei, 144 pp. [in Chinese]","Ruedi, M., Stadelmann, B., Gager, Y., Douzery, E. J. P., Francis, C. M., Lin, L. - K., Guillen-Servent, A. & Cibois, A. (2013) Molecular phylogenetic reconstructions identify East Asia as the cradle for the evolution of the cosmopolitan genus Myotis (Mammalia, Chiroptera). Molecular Phylogenetics and Evolution, 69, 437 - 449. http: // dx. doi. org / 10.1016 / j. ympev. 2013.08.011","Tsytsulina, K. & Strelkov, P. P. (2001) Taxonomy of the Myotis frater species group (Vespertilionidae, Chiroptera). Bonner zoologischer Beitrage, 50, 15 - 26.","Kruskop, S. V., Borisenko, A. V., Ivanova, N. V., Lim, B. K. & Eger, J. L. (2012) Genetic diversity of northeastern Palaearctic bats as revealed by DNA barcodes. Acta Chiropterologica, 14, 1 - 14. http: // dx. doi. org / 10.3161 / 150811012 x 654222"]}

Myotis soror sp. n. Figures 2, 3, 5, 8 a, and 9 a 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). 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. 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. 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). 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. 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)., Published as part of Ruedi, Manuel, Csorba, G��bor, Lin, Liang- Kong & Chou, Cheng-Han, 2015, Molecular phylogeny and morphological revision of Myotis bats (Chiroptera: Vespertilionidae) from Taiwan and adjacent China, pp. 301-342 in Zootaxa 3920 (1) on pages 327-329, DOI: 10.11646/zootaxa.3920.2.6, http://zenodo.org/record/287922, {"references":["Chou, C. - H. (2004) Taxonomic status of the genus Myotis (Chiroptera: Vespertilionidae) in Taiwan. Department of Wildlife Ecology, Tunghai University, Tunghai, 115 pp. [in Chinese]","Lin, L. K., Lee, L. L. & Cheng, H. C. (2004) Bats of Taiwan (2 nd ed.). National Museum of Natural Sciences, Taipei, 177 pp. [in Chinese]","Tsytsulina, K. & Strelkov, P. P. (2001) Taxonomy of the Myotis frater species group (Vespertilionidae, Chiroptera). Bonner zoologischer Beitrage, 50, 15 - 26.","Yoshiyuki, M. (1989) A systematic study of the Japanese chiroptera. National Science Museum monographs, 7, 1 - 242.","Csorba, G., Chou, C. - H., Ruedi, M., Gorfol, T., Motokawa, M., Wiantoro, S., Thong, V. D., Son, N. T., Lin, L. - K. & Furey, N. (2014) The reds and the yellows: a review of Asian Chrysopteron Jentink, 1910 (Chiroptera: Vespertilionidae: Myotis). Journal of Mammalogy, 95, 663 - 678. http: // dx. doi. org / 10.1644 / 13 - mamm-a- 200"]}

G��rf��l, Tam��s, Est��k, P��ter, G��bor, Csorba (2013): The Subspecies Of Myotis Montivagus - Taxonomic Revision And Species Limits (Mammalia: Chiroptera: Vespertilionidae). Acta Zoologica Academiae Scientiarum Hungaricae 59 (1): 41-59, DOI: http://doi.org/10.5281/zenodo.5736085, {"references":["GhOSh, M. K. (1989) Newlocality-recordsforMyotismontivaguspeytoniWroughton & Ryley, 1913, andMurinacyclotiscyclotisDobson, 1872 (Chiroptera: Vespertilionidae) intheeasternghatsofAndhraPradesh, India. JournalofBombayNaturalHistorySociety 86: 93-94.","HeLLeR, K.-G. & VOLLeTh, M. (1988) FledermauseausMalaysia. 1. Beobachtungenzur biologie, morphologieundtaxonomie (Mammalia: Chiroptera). SenckenbergianaBiologica 69: 243-276.","HeNDRichSeN, D. K., BATeS, P. J. J. & HAyeS, B. D. (2001a) Recentrecordsofbats (Chiroptera) fromCambodia. ActaChiropterologica 3(1): 21-32.","HeNDRichSeN, D. K., BATeS, P. J. J., HAyeS, B. D. & WALSTON, J. L. (2001b) Recent records ofbats (Mammalia: Chiroptera) fromVietnamwithsixspeciesnewtothecountry. Myotis 39: 35-122.","HiLL, J. E. (1962) NotesonsomeinsectivoresandbatsfromUpperBurma. Proceedingsofthe Zoological Society of London 139(1): 119-137.","HiLL, J. E. & FRANciS, C. M. (1984) New bats (Mammalia: Chiroptera) and new records of bats from Borneo and Malaya. Bulletin of the British Museum (Natural History), Zoology Series 47(5): 303-329.","HiLL, J. E. & ThONgLONgyA, K. (1972) Bats from Thailand and Cambodia. Bulletin of the British Museum (Natural History), Zoology Series 22(6): 173-196.","HOThORN, T., HORNiK, K. & ZeiLeiS, A. (2006) Unbiasedrecursivepartitioning: Acondition- al inference framework. Journal of Computational and Graphical Statistics 15(3): 651-674.","KOOpmAN, K. F. (1994) Chiroptera: Systematics. Pp. 217. In: NieThAmmeR, J., SchLiemANN, H. & STARcK, D. (eds): HandbookofZoology. WalterdeGruyter, Berlin.","KRuSKOp, S. V. & TSyTSuLiNA, K. A. (2001) Anewbig-footedmouse-earedbatMyotisanna- miticussp. nov. (Vespertilionidae, Chiroptera) fromVietnam. Mammalia 65(1): 63-72.","KuzNeTSOv, G. V., BORiSeNKO, A. V. & ROzhNOv, V. V. (2001) Asynopsisofthemammalfauna oftheVuQuangNatureReserve. Pp. 35-46. In: KORzuN, L. P. & KALyAKiN, M. V. (eds): Materials of zoological and botanical studies in Vu Quang Nature Reserve (Ha Tinh Province, Vietnam). JointRussian-VietnameseScienceandTechnologicalTropicalCentre, MoscowandHanoi.","LuNDe, D. P., MuSSeR, G. G. & PhAmDucTieN (2003) Recordsofsomelittleknownbats (Chiroptera: Vespertilionidae) fromVietnam. Mammalia 67(3): 459-461.","MANDAL, A. K., PODDAR, A. K. & BhATTAchARyyA, T. P. (2000) Further new records of bats from Mizoram, India. Records of the Zoological Survey of India 98(2): 147-154.","PAyNe, J., FRANciS, C. M. & PhiLLippS, K. (1985) A field guide to the mammals of Borneo. The SabahSocietyandWorldWildlifeFundMalaysia, KotaKinabaluandKualaLumpur, 332 pp.","RCOReTeAm (2012) R: Alanguageandenvironmentforstatisticalcomputing. RFounda- tionforStatisticalComputing, Vienna. http://www.R-project.org","SmiTh, A. T. & Xie, Y. (2008) AguidetothemammalsofChina. PrincetonUniversityPress, Princeton, 544 pp.","SuyANTO, A. & STRueBig, M. J. (2007) BatsoftheSangkuliranglimestonekarstformations, EastKalimantan - apriorityregionforBorneanbatconservation. ActaChiropterologica 9(1): 67-95.","TATe, G. H. H. (1941) ResultsoftheArchboldExpeditions 39. ReviewofMyotisofEurasia. Bulletin of the American Museum of Natural History 78(8): 537-565.","TATe, G. H. H. (1942) ResultsoftheArchboldExpeditions 47. ReviewoftheVespertilionine bats, withspecialattentiontogeneraandspeciesoftheArchboldCollections. Bulletin of the American Museum of Natural History 80(7): 221-297.","ThOmAS, O. (1916) ListofMicrochiroptera, otherthanleaf- nosebats, inthecollectionof theFederatedMalayStatesMuseums. JournaloftheFederatedMalayStatesMuseums 7(1): 1-6.","ThOmAS, O. (1920) ScientificresultsfromtheMammalSurveyXXIII. Anewbatofthegenus Myotis from Sikkim. Journal of Bombay Natural History Society 27: 248-249.","TOpAL, Gy. (1970) OnthesystematicstatusofPipistrellusannectansDobson, 1871 andMyo- tisprimulaThomas, 1920 (Mammalia). Annaleshistorico-naturalesMuseinationalis hungarici 62: 373-379.","VANiThARANi, J. (2006) NoteworthyrepresentativesofbatspeciesinAgasthyamalaiBio- sphereReserve, TamilNadu. JournalofTheoreticalandExperimentalBiology 2(2): 47-59.","VANiThARANi, J., MALAThi, U. S. U. & SuNDARi, A. K. (2005) New records of bats from Ka- lakadMundanthuraiTigerReserve, India. BatNetCCINSANewsletter 6(1): 13-14.","VeLAzcO, P. M., GARDNeR, A. L. & PATTeRSON, B. D. (2010) SystematicsofthePlatyrrhinus hellerispeciescomplex (Chiroptera: Phyllostomidae), withdescriptionsoftwonew species. Zoological Journal of the Linnean Society 159(3): 785-812.","WANg, Y. X. (2003) A complete checklist of mammal species and subspecies in China: a taxonomic andgeographicreference. ChinaForestryPublishingHouse, Beijing, 394 pp.","WiLSON, D. E., HeLgeN, K. M., YuN, C. S. & GimAN, B. (2006) Smallmammalsurveyattwo sitesinplantedforestzone, Bintulu, Sarawak. MalayanNatureJournal 59(2): 165-187.","WROughTON, R. C. & RyLey, K. V. (1913) ScientificresultsfromtheMammalSurveyIII. A new species of Myotis from Kanara. Journal of Bombay Natural History Society 22: 13-21."]}

Background: There have been reported cases of host- switching in avian and lizard species of Plasmodium (Apicomplexa, Haemosporidia), as well as in those infecting different primate species. However, no evidence has previously been found for host-swapping between wild birds and mammals. Methods: This paper presents the results of the sampling of blood parasites of wild-captured bats from Madagascar and Cambodia. The presence of Haemosporidia infection in these animals is confirmed and cytochrome b gene sequences were used to construct a phylogenetic analysis. Results: Results reveal at least three different and independent Haemosporidia evolutionary histories in three different bat lineages from Madagascar and Cambodia. Conclusion: Phylogenetic analysis strongly suggests multiple host-switching of Haemosporidia parasites in bats with those from avian and primate hosts.

Murina harrisoni sp. nov. Holotype HZM 1.36316 (field number JLW 04.07), adult &female;, body in alcohol, skull ex- tracted. Collected by J. L. Walston on 5 February, 2000. Type locality OTuk Chehn, KiriromNationalPark,Kompong Speu Province, Cambodia, 11��29.611���N, 104��12.746���E. Diagnosis This isamediumsizedtube-nosedbat (FA35.9 mm), withalargeskull(totallengthofskull18.39mm; Table1) andwiththeplagiopatagiumattached tothe baseof thefirsttoe(Fig. 1). Theanteriorpartoftherostrumisbulbous toaccommodatethelargecanines(Fig. 2A). Thefirstupperin- cisor(I 2) isratherslender, bifidand withits secondary cuspveryshort; theprincipalcuspoftherobustsecondincisor(I 3 ) isrel- ativelyshort, onlyslightlyexceeding halftheheightof I 2. Description Ear lengthis 14 mmandtheearconchis without an emargination on the posterior border. Thetragusistypicalof thegenus, slightly bent backwardsand7.4 mmin length.Thedorsalfur(describedfromthewetspecimen) isauniform reddish-brown; theindividualhairsare withouta wellde- finedbanding, althoughtheterminalpartis aslightlydarker reddish-brown. On theventralaspect(includingthethroat), the hairsarewhitethroughouttheirlength. Thetailmembraneis evenlyfurredaboveandthelastvertebraisfreefromtheuro- patagium. Theplagiopatagiumisattachedtothebase ofthefirsttoe. Theskullisheavilybuiltwiththeros- trumdeepandmassive, anteriorlyratherbulbous to accommodatethelarge canines(Fig. 2A). Thesagittalandlambdoidcrestsaredistinct. Thenarialemarginationisnar- row, withthe lengthconsiderablyexceedingthewidth; the lengthofthepalatal emar- ginationalsoslightlyexceedsthe width. Thereisnobasioccipitalfissure. Thefirstupperincisor (I2) isratherslender, its sec- ondarycuspbeing onlyacingularcusp; the principal cusp of the second incisor (I 3) is relatively short, only slightly exceeding half the height of I 2. I 3 is situated alongside, rather than posterior to, I 2. I 3 is less than half the height of upper canine (C 1), which greatly exceeds the height of the second up- per premolar (P 4). In lateral view, the first upper premolar (P 2) is only slightly shorter than the second (P 4) and the crown area is subequal (Fig. 3A). The mesostyle of the first (M 1) and second (M 2) upper molars are not reduced and the metacones are distinct- ly higher than the paracones. The angular process of the mandible is elongated, with a tall coronoid process (Fig. 3B). The lower canine (C 1) greatly exceeds the first (P 2) and second (P 4) lower premolars in height when viewed laterally; P 2 and P 4 are sube- qual in height; P 2 is about half the crown area of P 4. The hypoconid and entoconid of the first (M 1) and second (M 2) lower mo- lars are well separated from the anterior cusps and clearly form a posterior trigonid. Craniodental measurements (in mm) are as follows: STOTL 18.39; CBL 16.73; CCW 5.2; M 3 M 3 W 6.37; ZYW 11.33; MAW 9.61; IOW 4.51; CM 3 L 6.49; CP 4 L 3.37; ML 13.03; CM 3 L 7.15; CP 4 L 3.19; CPH 5.21. Measurements M. harrisoni n. sp. (holotype) M. hilgendorfi M. huttoni M. leucogaster M. puta M. rozendaali x n min���max n min���max n min���max n min���max n min���max FA 35.90? 40��� 45? 33��� 37? 40 ��� 44? 30��� 37? 32���34 STOTL 18.39618.39��� 19.64 1316.39��� 18.15 118.802016.63���18.09 2 15.51���15.80 CM 3 L6.4996.16��� 6.59 165.73��� 6.08 16.08205.79���6.26 2 5.21���5.34 CP4 L3.3793.04��� 3.37 162.65��� 2.99 12.93202.69���3.01 2 2.44 ���2.54 ML13.03912.88��� 13.87 1411.17��� 12.54 112.821711.43��� 12.45 CM 3 L7.1596.75��� 7.28 146.26��� 6.70 16.60176.34��� 6.78 CPH 5.21 9 4.61��� 5.21 133.87 ��� 4.54 14.29 173.86��� 4.51 Etymology Named in honour of Dr. David Lakin Harrison, who, as Chairman of Trustees, has supported, encouraged and actively par- ticipated in the extensive researches of the Harrison Institute into the bats of Southern and South-East Asia. COMPARISONS The insertion point of the plagiopa- tagium (which is attached to the base of the first toe in M. harrisoni) differentiates this taxon from all other Murina species (the attachment point of which is the base of the first claw) except for M. hilgendorfi (including intermedia, ognevi and sibirica), which has a similar point of insertion. However, unlike M. hilgendorfi and all other Murina taxa, the second upper incisor (I 3) of M. harrisoni is much shorter than the first (I 2), with the principal cusp of I 3 only slightly exceeding half the height of I 2. In addition, the rostral profile of M. harrisoni differs markedly from that of M. hilgendorfi . In M. harrisoni, the rostrum is deep and massive and anteriorly bulbous to accommodate the large canines; in consequence, the rostral profile is concave (Fig. 2A). In M. hilgendorfi, the rostrum is not enlarged and the rostral profile is straighter (Fig. 2B). M. harrisoni is further distinguished from M. hilgendorfi by the first upper premolar (P 2) which is subequal in height with the second (P 4) in M. harrisoni whilst in M. hilgendorfi it is about two-thirds the height and with a relatively smaller crown area (Fig. 3C). The pelage of M. harrisoni is reddish-brown rather than grey and the fore- arm is considerably shorter (FA = 35.9 mm) as compared to> 40 mm in M. hilgendorfi. The cranial dimensions of M. harrisoni are superficially similar to those of M. le- ucogaster rubex. However, in addition to the difference in the insertion point of the plagiopatagium and the relative size of the upper incisor (as outlined above), the canine of M. harrisoni exceeds that of rubex in length, the narial and palatal emarginations are narrower, the rostral profile is more con- cave, the rostrum distinctly more bulbous and the crown area of P 2 is larger relative to that of P 4. In M. harrisoni, P 2 and P 4 are subequal in size; in M. l. rubex, the crown area of P 2 is less than half that of P 4. Within the ��� cyclotis -group���, M. harri- soni has a similar development of the meso- style of the first (M 1) and second (M 2) up- per molars and the posterior trigonid of the first (M 1) and second (M 2) lower molars to those of M. rozendaali, M. huttoni and M. puta. However, in addition to the diagnostic characters of M. harrisoni listed above, these latter taxa are also smaller in all cra- nial and dental measurements (Table 1). Habitat The single specimen was collected at 22:50 hours in a mist net which was set across a river in disturbed semi-evergreen gallery forest with many immature trees. The area had been selectively logged in the past. The river was about 5 m wide with medium-fast flowing water (J. L. Walston, pers. comm.). Distribution Currently this species is only known from a single specimen collected in Kiri- rom National Park, Cambodia (11��30���N, 104��13���E). However, future studies may show it to be more widespread in the forests of South-East Asia. ACKNOWLEDGEMENTS We are indebted to Joe Walston of the Wildlife and Conservation Society, Cambodia for making the specimens available for study. We thank Charles Francis and Hao-Chi Kuo for their expert advice on Murina characters. Paula Jenkins (The Natural History Museum, London), Ling-Ling Lee (National Taiwan University, Taipei), Hsi-Chi Cheng (Endemic Species Research Institute, Chichi) and Yen-Jean Chen (National Museum of Natural Science, Tai- chung) kindly provided access to the specimens under their care. We thank David Harrison, Malcolm Pearch and Karen Bates at the Harrison Institute for their help with the preparation of specimens and their advice concerning the manuscript and Peter Ujhelyi for the final elaboration of the drawings. Finally, we would like to thank the Darwin Initiative of the UK Government for their support of taxonomic bat studies in South-East Asia., Published as part of Csorba, G��bor & Bates, Paul J. J., 2005, Description of a new species of Murina from Cambodia (Chiroptera: Vespertilionidae: Murininae), pp. 1-7 in Acta Chiropterologica 7 (1) on pages 2-6, DOI: 10.3161/1733-5329(2005)7[1:doanso]2.0.co;2, http://zenodo.org/record/3944989

Rhinolophus affinis HORSFIELD, 1823 In the original description of the species HORSFIELD (1823) indicated no type specimen. Beside a specimen (labelled as holotype) stored in the BM(NH), JENTINK (1887) listed two more specimens in the RMNH, Leiden marked as types. However, the two RMNH individuals (RMNH 25236, cat. ost. b and RMNH 25237, cat. ost. c) represented by skulls only, proved to be Hipposideros larvatus. The possible reason of the confusion should be the fact, that HORSFIELD worked with the two species in question at the same time (the descriptions appeared in the same book). Since there was no holotype designation in HORSFIELD's work, these three specimens are regarded as syntypes; consequently, the BM(NH) specimen (No. 79.11.21.70) as the only R. affinis is designated herein as lectotype., Published as part of Csorba, G., 2002, Remarks on some types of the genus Rhinolophus (Mammalia, Chiroptera), pp. 217-226 in Annales Historico-Naturales Musei Nationalis Hungarici 94 on page 221, DOI: 10.5281/zenodo.3839717

Rhinolophus thomasi septentrionalis SANBORN, 1939 The taxon septentrionalis once was described and later widely accepted as a subspecies of thomasi, differing from the nominotypical race by its larger size and slightly extruded anterior upper premolars (SANBORN 1939). The holotype of septentrionalis (FMNH 33291) and other specimens from Yunnan stored in the FMNH and USNM agree in every respect with each other. However, it is much bigger in external measurements than thomasi and latifolius (FA 51.1-55.5 against 40.5-48.0; SL 19.79-20.98 against 17.87-19.98; andC M 3 7.65-8.40 against 6.82-7.67 mm), and has strong, widely based, long canines. These differences support the view, that septentrionalis differs from R. thomasi at specific level. The taxon sinicus was described as a subspecies of R. rouxi by ANDERSEN (1905) who separated it on the basis of its smaller skull and toothrow measurements. As ANDERSEN remarked, the general size of sinicus as is the smallest example of the typical form of R. rouxi. This taxonomical position of sinicus was generally accepted, but T H O M A S (1997) in her detailed work, based on phenetic analysis and DNA techniques, verified that sinicus represents a distinct species occuring in the Himalayas, Myanmar, northern Vietnam and southern China. Nevertheless, the relation and the specific boundary between R. sinicus and R. thomasi is unclear. The extremely hastate, excessively shortened lancet thought to be diagnostic for R. thomasi (ANDERSEN 1905, CORBET & HILL 1992, K O O P M A N 1994) is not clearly expressed in all specimens of that species, while a similar shortening of lancet can be found in several R. sinicus. The types of both species are unusually small specimens and almost all subsequently collected individuals are larger. It means that although the type of R. sinicus is much larger than the type of R. thomasi (therefore justifies the distinctness on species level), it overlaps in size with the majority of the known R. thomasi specimens (determined hereby the slender upper and lower canine only). On average, R. sinicus is much bigger than R. thomasi. The form septentrionalis is therefore better referable to R. sinicus; the large external measurements (the forearm length is over 50 mm) validate the subspecific separation within the species., Published as part of Csorba, G., 2002, Remarks on some types of the genus Rhinolophus (Mammalia, Chiroptera), pp. 217-226 in Annales Historico-Naturales Musei Nationalis Hungarici 94 on pages 221-223, DOI: 10.5281/zenodo.3839717, {"references":["SANBORN, C. C. (1939): Eight new bats of the genus Rhinolophus. - Publications of the Field Museum of Natural History Zoological Series 24: 37 - 43.","K O O P M A N, K. F. (1994): Chiroptera: Systematics. Handbook of Zoology. Mammalia, part 60. - Walter de Gruyter, Berlin, 217 pp."]}

The description of this taxon as a subspecies of R. lepidus from Upper Burma (Myanmar) was published by O L D F I E L D THOMAS on behalf of ANDERSEN (1918), based on the short notes of the latter. The diagnostic characters of shortridgei ("skull and teeth averaging larger") appeared only in the key given for the species and subspecies of the pusillus-group but even without comparison of the measurements with the other named forms. According to SlNHA (1973) shortridgei differs from R. lepidus lepidus in having a longer hind foot (55-63% of the tibia, against 45.8-47.5%) and longer mandible. However, investigation of the type skull (BM(NH) 18.8.3.1) and other specimens (housed in the collection of USNM, FMNH, HNHM) revealed well-defined differences as compared with the other subspecies of R. lepidus; upper canines are strong, wide-based; sagittal crest extending posteriorly to the lambda and skull length is over 17 mm. Consequently, the taxon shortridgei is considered as a full species., Published as part of Csorba, G., 2002, Remarks on some types of the genus Rhinolophus (Mammalia, Chiroptera), pp. 217-226 in Annales Historico-Naturales Musei Nationalis Hungarici 94 on pages 219-220, DOI: 10.5281/zenodo.3839717, {"references":["SlNHA, Y. P. (1973): Taxonomic studies on the Indian horseshoe bats of the genus Rhinolophus Lacepede. - Mammalia 37 (4): 603 - 630."]}

Rhinolophus philippinensis montanus GOODWIN, 1979 G O O D W I N (1979) discussed the differences between his new montanus and the other subspecies of R. philippinensis, and noted its much smaller size, differently shaped sella and connecting process, more pronounced nasal swellings and more crowded situation of the small premolars. Investigation of the known speci�� mens (holotype, paratype and two more individuals collected together the types, A M N H 237811-237814) has shown that these differences are definitely beyond intraspecific variation of R. philippinensis and leave no doubt that montanus is a distinct species. The external appearence of the noseleaf of montanus is intermediate between R. philippinensis and R. macrotis. As already ANDERSEN (1907) noted, R. macrotis is an example of "a type of low level of evolution which has no closer relative than the primitive forms of the Rh. philippinensis group" and "the sella of macrotis might properly be described as that of a philippinensis deprived of its lateral expansions; the shape of the connecting process and lancet also point towards relationship with philippinensis ". The noseleaf features of the much later described R. montanus are filling this gap. *, Published as part of Csorba, G., 2002, Remarks on some types of the genus Rhinolophus (Mammalia, Chiroptera), pp. 217-226 in Annales Historico-Naturales Musei Nationalis Hungarici 94 on pages 223-225, DOI: 10.5281/zenodo.3839717, {"references":["G O O D W I N, R. E. (1979): The bats of Timor: systematics and ecology. - Bulletin of the American Museum of Natural History 163: 73 - 122."]}

The confusing history of the name borneensis was reviewed in detail by A N �� DERSEN (1905) who described it as "accumulation of errors and wrong identifications" which resulted in the fact that " Rh. borneensis has for many years been completely confused not only with several more or less closely related species, but also with the widely different Rh. minor " (= R. pusillus). One of the possible reason of the confusion should be the mis-matching of labels and/or skulls in the Museum f��r Naturkunde, Berlin (MNB). There are two skulls (in very bad condition) in the type collection of MNB (2533.1 and 2533.2) which certainly represent specimens of R. borneensis, although both labelled as " Rhinolophus minor, type, Labuan, Java ". On the other hand, the type of R. minor is in the BM (NH). Since the type lo- cality of R. borneensis is also Labuan (the Malayan island off Borneo, not in Java), and according to PETERS (1871) its type is deposited in the Berlin Museum, the MNB 2533.1 and 2533.2 specimens are undoubtedly the mis-labelled types of R. borneensis., Published as part of Csorba, G., 2002, Remarks on some types of the genus Rhinolophus (Mammalia, Chiroptera), pp. 217-226 in Annales Historico-Naturales Musei Nationalis Hungarici 94 on pages 220-221, DOI: 10.5281/zenodo.3839717, {"references":["PETERS, W. (1871): Uber die Gattungen und Arten der Hufeisennasen, Rhinolophi. - Monatsberichte der Koniglichen Preuss Akademie der Wissenschaften zu Berlin 301 - 332."]}

Csorba, G . (2002): Remarks on some types of the genus Rhinolophus (Mammalia, Chiroptera). Annales Historico-Naturales Musei Nationalis Hungarici 94: 217-226, DOI: http://doi.org/10.5281/zenodo.3839717, {"references":["ANDERSEN , K . (1905): On some bats of the genus Rhinolophus, with remarks on their mutual affinities, and descriptions of twenty-six new forms. - Proceedings of the Zoological Society of London 2: 75-145.","ANDERSEN , K. (1907): Chiropteran notes. - Annali del Museo Civico di Storia Naturale Giacomo Doria, Genova (Ser. 3) 3: 5-45.","ANDERSEN , K. (1918): Diagnoses of new bats of the families Rhinolophidae and Megadermatidae.- The Annals and Magazine of Natural History (Ser. 9) 2: 374-384.","CORBET , G. B. & H I L L , J . E. (1992): The mammals of the Indomalayan region. - Oxford University Press, Oxford, 488 pp.","DOBSON , G. E. (1878): Catalogue of the Chiroptera in the collection of the British Museum. - Order of the Trustees, London, 567 pp.","GOODWIN , R. E. (1979): The bats of Timor: systematics and ecology. - Bulletin of the American Museum of Natural History 163: 73-122.","HORSFIELD , T. (1823): Zoological researches in Java. - Kingbury, Parbury and Allen, London.","JENTINK, F. A. ( 1887): Catalogue osteologique des mammiferes. - Museum d'Histoire Naturelle des Pays-Bas 9: 1-360.","JENTINK , F. A. (1888): Catalogue systematique des mammiferes (rongeurs, insectivores, cheiropteres, edentes et marsupiaux). - Museum d'Histoire Naturelle des Pays-Bas 12: 1-280.","KOOPMAN , K. F. (1994): Chiroptera: Systematics. Handbook of Zoology. Mammalia, part 60. - Walter de Gruyter, Berlin, 217 pp.","PETERS, W. (1871): Uber die Gattungen und Arten der Hufeisennasen, Rhinolophi. - Monatsberichte der Koniglichen Preuss Akademie der Wissenschaften zu Berlin 301-332.","SANBORN, C. C. (1939): Eight new bats of the genus Rhinolophus. - Publications of the Field Museum of Natural History Zoological Series 24: 37-43.","SlNHA, Y. P. (1973): Taxonomic studies on the Indian horseshoe bats of the genus Rhinolophus Lacepede. - Mammalia 37 (4): 603-630.","THOMAS, N. M. (1997): A systematic review of selected Afro-Asiatic Rhinolophidae (Mammalia: Chiroptera): an evaluation of taxonomic methodologies. - Unpublished PhD. Thesis. Harrison Zoological Museum, Sevenoaks, 211 pp."]}

Rhinolophus pusillus T E M M I N C K, 1834 The types of R. pusillus in the RMNH, Leiden caused a lot of mental labour. DOBSON (1878) investigated the types and concluded that they were "undoubtedly specimens of R. hipposide rus " which led to the statement that R. hipposideros should occur in Java. ANDERSEN (1905) later speculated that "an interchange of la�� bels has taken place in that Museum". The small specimens in the Leiden Museum were always kept in glass vials from the very beginning, with a little round label glued on the cork cover (C. SMEENK pers. comm.). Needless to say, these labels may have come off, so there was always the danger of specimens becoming mislabelled or interchanged. In fact, the type series in Leiden consists of five syntypes (RMNH 35177-35181), of which three represent R. hipposideros (RMNH 35178 [= Rh. hipposideros cat. syst, b] and 35179 [= Rh. hipposideros cat. syst, c] mounted specimens, skulls intact; RMNH 35181 [= Rh. hipposideros cat. ost. a] separate skull). One of the remaining two specimens (RMNH 35177 [= Rh. hipposideros cat. syst a, cat. ost. b]) consists of a skull of R. pusillus and a mounted skin of R. hipposideros', the another one (RMNH 35180 [= Rh. hipposideros cat. syst, d] mounted with skull intact) is with no doubt R. pusillus. These facts explain why DOBSON (1878) and JENTINK (1887) referred R. pusillus as a synonym of R. hipposideros, and make it clear that ANDERSEN (1905) was right when accepted T E M M I N C K ' S statement that the types of R. pusillus were brought back from Java. Since the cranial characters are widely used features in the group, from the syntypes representing genuine R. pusillus the RMNH 35177 specimen (a cleaned skull) is designated herein as lectotype; the skin of R. hipposideros bearing the same number is regarded as mis-labelled. The RMNH 35180 mounted specimen is the paralectotype of R. pusillus. The shape of the rostral profile of R. pusillus was desribed by CORBET and HILL (1992) as being nearly straight, almost horizontal (contrary to the up- ward-curving rostral profile of R. lepidus ). Taking into consideration of the lectotype specimen of the former and the variability of both species, this character is not typical or uniform, and cannot be used for distinction of the two species. The development of the posterior median swellings (which affects the shape of the rostral profil) is either a variable feature of both species or it has a taxonomical significance not fully understood as yet., Published as part of Csorba, G., 2002, Remarks on some types of the genus Rhinolophus (Mammalia, Chiroptera), pp. 217-226 in Annales Historico-Naturales Musei Nationalis Hungarici 94 on pages 218-219, DOI: 10.5281/zenodo.3839717