Abstract
Murina is a wide-ranging genus of insectivorous bats that occurs throughout Asia whose known diversity has recently been increasing with the description of 26 new species in the past 20 years. The Philippine archipelago—mostly comprised of islands with oceanic origin and thus a history of isolation—has produced a fauna rich in endemic species, including bats. Here, we describe the morphological and genetic diversity among Murina specimens collected from across the Philippine islands and compare them to representatives from the Southeast Asian mainland and adjacent islands. Our results show that the wide-spread M. cyclotis represents a radiation, beginning around 11 mya, of five distinct Murina species endemic to the Philippines, including three single-island endemics. These five new M. cyclotis group species are mostly allopatric in distribution, except for two species—the smallest and largest that occur in the Philippines—that co-occur on Mindanao Island. Our study documents Murina suilla, a species occurring in Indonesia, Malaysia and Thailand, and only occurring on Mindoro and Palawan Islands within the Philippines. Additionally, we describe a new species that is sister to M. suilla and is widespread across the Philippine archipelago. Overall, our study has increased the number of Philippine Murina species from two to seven. As the first phylogeographic analysis of a laryngeal echolocating bat genus within the Philippines, our study provides evidence that the diversification of understory insectivorous bats largely echoes that of fruit bats (Pteropodidae), which is characterized by endemic Philippine lineages that are primarily confined to known biogeographic regions that constitute sets of modern islands that were connected during the last ice age, or to individual oceanic islands. We hypothesize that subsequent phylogeographic investigations of other bat taxa may reveal similar, previously unrecognized, endemic radiations within the highly biodiverse Philippines.
References
- Almeida, F.C., Giannini, N.P., DeSalle, R. & Simmons, N.B. (2011) Evolutionary relationships of the old world fruit bats (Chiroptera, Pteropodidae): Another star phylogeny? BMC Evolutionary Biology, 11, e281. https://doi.org/10.1186/1471-2148-11-281
- Almeida, F.C., Simmons, N.B. & Giannini, N.P. (2020) A species-level phylogeny of Old World fruit bats with a new higher-level classification of the family Pteropodidae. American Museum Novitates, 3950, 1–24. https://doi.org/10.1206/3950.1
- Apan, A., Suarez, L.A., Maraseni, T. & Castillo, J.A. (2017) The rate, extent and spatial predictors of forest loss (2000–2012) in the terrestrial protected areas of the Philippines. Applied Geography, 81, 32–42. https://doi.org/10.1016/J.APGEOG.2017.02.007
- Balete, D.S. (2010) Food and roosting habits of the lesser false vampire bat, Megaderma spasma (Chiroptera: Megadermatidae) in a Philippine lowland forest. Asia Life Sciences, Supplement 4, 111–129.
- Balete, D.S., Alviola, P.A., Duya, M.R.M., Duya, M.V., Heaney, L.R. & Rickart, E.A. (2011) Chapter 3: The mammals of the Mingan Mountains, Luzon: evidence for a new center of mammalian endemism. Fieldiana Life and Earth Sciences, 2011, 75–87. https://doi.org/10.3158/2158-5520-2.1.75
- Balete, D.S. & Heaney, L.R. (1997) Density, biomass, and movement estimates for murid rodents in mossy forest on Mount Isarog, Southern Luzon, Philippines. Ecotropica, 1997, 91–100.
- Balete, D.S., Heaney, L.R. & Rickart, E.A. (2013) The Mammals of Mt. Irid, Southern Sierra Madre, Luzon Island, Philippines. National Museum of the Philippines: Journal of Natural History, 2013, 15–21.
- Balete, D.S., Heaney, L.R. & Rickart, E.A. (2020) A comparison of native and non-native small mammals in adjacent karst and volcanic soil regions in NE Luzon, Philippines. Philippine Journal of Systematic Biology, 14, 1–16. https://doi.org/10.26757/pjsb2020b14011
- Balete, D.S., Quidlat, R.S. & Ibañez, J.C. (2006) Non-volant mammals of Mt. Hamiguitan, eastern Mindanao, Philippines. BANWA Archives, 3, 65–80. [2004–2013]
- Balete, D.S., Rickart, E.A., Heaney, L.R., Alviola, P.A., Duya, M.V., Duya, M.R.M., Sosa, T. & Jansa, S.A. (2012) Archboldomys (Muridae: Murinae) reconsidered: a new genus and three new species of shrew mice from Luzon Island, Philippines. American Museum Novitates, 3754, 1–60. https://doi.org/10.1206/3754.2
- Balete, D.S., Rickart, E.A., Heaney, L.R. & Jansa, S.A. (2015) A new species of Batomys (Muridae, Rodentia) from southern Luzon Island, Philippines. Proceedings of the Biological Society of Washington, 128, 22–39. https://doi.org/10.2988/0006-324X-128.1.22
- Bickham, J.W., Wood, C.C. & Patton, J.C. (1995) Biogeographic implications of cytochrome b sequences and allozymes in sockeye (Oncorhynchus nerka). Journal of Heredity, 86, 140–144. https://doi.org/10.1093/oxfordjournals.jhered.a111544
- Birdlife International (2014) Important Bird and Biodiversity Areas in Danger. Available from: https://www.datazone.birdlife.org/sowb/sowbpubs (accessed 9 June 2025)
- Chen, J., Liu, T., Deng, H., Ning, X. & Zhou, J. (2017) A new species of Murina bats was discovered in Guizhou Province, China. Cave Research, 2, 1–10.
- Daipan, B.P.O. (2021) Patterns of forest cover loss in the terrestrial Key Biodiversity Areas in the Philippines: critical habitat conservation priorities. Journal of Threatened Taxa, 13 (13), 20019–20032. https://doi.org/10.11609/jott.6904.13.13.20019-20032
- Duya, M.R.M., Alviola, P.A., Duya, M.V., Balete, D.S. & Heaney, L.R. (2007) Report on a survey of mammals of the Sierra Madre Range, Luzon Island, Philippines. Banwa, 4 (1), 41–68.
- Eger, J.L. & Lim, B.K. (2011) Three new species of Murina from southern China (Chiroptera: Vespertilionidae). Acta Chiropterologica, 13, 227–243. https://doi.org/10.3161/150811011X624730
- Esselstyn, J. (2007) A new species of stripe-faced fruit bat (Chiroptera: Pteropodidae: Styloctenium) from the Philippines. Journal of Mammalogy, 88, 951–958. https://doi.org/10.1644/06-MAMM-A-294R.1
- Esselstyn, J.A. & Brown, R.M. (2009) The role of repeated sea-level fluctuations in the generation of shrew (Soricidae: Crocidura) diversity in the Philippine Archipelago. Molecular Phylogenetics and Evolution, 53, 171–181. https://doi.org/10.1016/j.ympev.2009.05.034
- Esselstyn, J.A., Evans, B.J., Sedlock, J.L., Khan, F.A.A. & Heaney, L.R. (2012) Single-locus species delimitation: A test of the mixed yule-coalescent model, with an empirical application to Philippine round-leaf bats. Proceedings of the Royal Society B: Biological Sciences, 279, 3678–3686. https://doi.org/10.1098/rspb.2012.0705
- Esselstyn, J.A., Garcia, H.J.D., Saulog, M.G. & Heaney, L.R. (2008) A new species of Desmalopex (Pteropodidae) from the Philippines, with a phylogenetic analysis of the teropodini. Journal of Mammalogy, 89, 815–825. https://doi.org/10.1644/07-MAMM-A-285.1
- Esselstyn, J.A., Maher, S.P. & Brown, R.M. (2011) Species interactions during diversification and community assembly in an island radiation of shrews. PLoS ONE, 6, e21885. https://doi.org/10.1371/journal.pone.0021885
- Esselstyn, J.A., Widmann, P. & Heaney, L.R. (2004) The mammals of Palawan Island, Philippines. Proceedings of the Biological Society of Washington, 117, 271–302.
- Francis, C.M. (2019) Field Guide to the Mammals of South-east Asia. 2nd Edition. Bloomsbury Publishing Plc, London, 408 pp.
- Francis, C.M. & Eger, J.L. (2012) A review of tube-nosed bats (Murina) from Laos with a description of two new species. Acta Chiropterologica, 14, 15–38. https://doi.org/10.3161/150811012X654231
- Gene Codes Corporation (2007) Sequencher. Version 4.8. Gene Codes Corporation, Ann Arbor, Michigan. [program]
- Giannini, N.P. & Simmons, N.B. (2003) A phylogeny of megachiropteran bats (Mammalia: Chiroptera: Pteropodidae) based on direct optimization analysis of one nuclear and four mitochondrial genes. Cladistics, 19, 496–511. https://doi.org/10.1111/j.1096-0031.2003.tb00385.x
- Giljum, S., Maus, V., Kuschnig, N., Luckeneder, S., Tost, M., Sonter, L.J. & Bebbington, A.J. (2022) A pantropical assessment of deforestation caused by industrial mining. Proceedings of the National Academy of Sciences of the United States of America, 119 (38), e2118273119. https://doi.org/10.1073/pnas.2118273119
- Golroudbary, S.R., Kraslawski, A., Wilson, B.P. & Lundström, M. (2022) Assessment of environmental sustainability of nickel required for mobility transition. Frontiers in Chemical Engineering, 4, 978842. https://doi.org/10.3389/fceng.2022.978842
- Goodman, S.M. & Ingle, N.R. (1993) Sibuyan Island in the Philippines–threatened and in need of conservation. Oryx, 27, 174–180. https://doi.org/10.1017/S0030605300027988
- Gracia, A.G., Mohagan, A.B., Burlat, J.C., Yu, W.L., Mondalo, J., Acma, F.M., Lumista, H.P., Calising, R. & Tanalgo, K.C. (2021) Conservation ecology of birds in Mt. Hilong-hilong, a Key Biodiversity Area on Mindanao Island, the Philippines. Journal of Threatened Taxa, 13 (5), 18110–18121. https://doi.org/10.11609/jott.6760.13.5.18110-18121
- Győrössy, D., Csorba, G., Szabadi, K.L., Estók, P., Tu, V.T., Thong, V.D., Furey, N.M., Huang, J.C.C., Tuanmu, M.N., Fukui, D., Zsebők, S. & Görföl, T. (2024) The calls of Vietnamese bats: a major step toward the acoustic characterization of Asian bats. Scientific Reports, 14 (1), 1–19. https://doi.org/10.1038/s41598-024-72436-6
- Hall, R. (1998) The plate tectonics of SE Asia and the distribution of land and sea. In: Hall, R. & Holloway, J.D. (Eds.), Biogeography and Geological Evolution of SE Asia. Backhuys, Leiden, pp. 99–131.
- Hall, R. (2012) Late Jurassic-Cenozoic reconstructions of the Indonesian region and the Indian Ocean. Tectonophysics, 570–571, 1–41. https://doi.org/10.1016/j.tecto.2012.04.021
- Hall, R. (2013) The palaeogeography of Sundaland and Wallacea since the Late Jurassic. Journal of Limnology, Supplement 2, 72, 1–17. https://doi.org/10.4081/jlimnol.2013.s2.e1
- He, H., Xiao, N. & Zhou, J. (2015) A new species of Murina from China (Chiroptera: Vespertilionidae). Cave Research, 2, 1–6.
- Heaney, L.R. (1986) Biogeography of mammals in SE Asia: estimates of rates of colonization, extinction and speciation. Biological Journal of the Linnean Society, 28, 127–165. https://doi.org/10.1111/j.1095-8312.1986.tb01752.x
- Heaney, L.R. (1991) An analysis of patterns of distribution and species richness among Philippine fruit bats (Pteropodidae). Bulletin of the American Museum of Natural History, 206, 145–167.
- Heaney, L.R., Balete, D.S., Alviola, P. & Rickart, E.A. (2012) Nyctalus plancyi and Falsistrellus petersi (Chiroptera: Vespertilionidae) from Northern Luzon, Philippines: Ecology, phylogeny, and biogeographic implications. Acta Chiropterologica, 14, 265–278. https://doi.org/10.3161/150811012X661602
- Heaney, L.R., Balete, D.S., Dolar, M.L., Alcala, A.C., Dans, A.T.L., Gonzales, P.C., Ingle, N.R., Lepiten, M.V., Oliver, W.L.R., Ong, P.S., Rickart, E.A., Tabaranza, B.R., Jr. & Utzurrum, R.C.B. (1998) A synopsis of the mammalian fauna of the Philippine Islands. Fieldiana Zoology, New Series, 88, 1–61.
- Heaney, L.R., Balete, D.S., Duya, M.R.M., Duya, M.V., Jansa, S.A., Steppan, S.J. & Rickart, E.A. (2016a) Doubling diversity: A cautionary tale of previously unsuspected mammalian diversity on a tropical oceanic island. Frontiers of Biogeography, 8.2, e29667. https://doi.org/10.21425/F5FBG29667
- Heaney, L.R., Balete, D.S., Gee, G.V., Lepiten-Tabao, M.V., Rickart, E.A. & Tabaranza, B.R. (2005) Preliminary report on the mammals of Balbalasang, Kalinga Province, Luzon. Sylvatrop, 13 (2003), 51–62.
- Heaney, L.R., Balete, D.S. & Rickart, E.A. (2016b) The Mammals of Luzon Island: biogeography and natural history of a Philippine fauna. Johns Hopkins University Press, Baltimore, Maryland, 287 pp.
- Heaney, L.R., Balete, D.S., Rickart, E.A., Utzurrum, R.C.B. & Gonzales, P.C. (1999) Mammalian diversity on Mount Isarog, a threatened center of endemism on southern Luzon Island. Fieldiana Zoology, New Series, 95, 1–62. https://doi.org/10.5962/bhl.title.3369
- Heaney, L.R., Dolar, M.L., Balete, D.S., Esselstyn, J.A., Rickart, E.A. & Sedlock, J.L. (2010) Synopsis of Philippine mammals. Available from: http://www.fieldmuseum.org/philippine_mammals/ (accessed 14 October 2024)
- Heaney, L.R., Gonzales, P.C., Utzurrum, R.C.B. & Rickart, E.A. (1991) The mammals of Catanduanes Island: implications for the biogeography of small land-bridge islands in the Philippines. Proceedings of the Asia-Pacific Forum on Karst Ecosystems and World Heritage, 104, 399–415.
- Heaney, L.R., Heideman, P.D., Rickart, E.A., Utzurrum, R.B. & Klompen, J.S.H. (1989) Elevational zonation of mammals in the central Philippines. Journal of Tropical Ecology, 5, 259–280. https://doi.org/10.1017/S0266467400003643
- Heaney, L.R., Kyriazis, C.C., Balete, D.S., Steppan, S.J. & Rickart, E.A. (2018) How small an island? Speciation by endemic mammals (Apomys, Muridae) on an oceanic Philippine island. Journal of Biogeography, 45, 1675–1687. https://doi.org/10.1111/jbi.13352
- Heaney, L.R. & Rickart, E.A. (1990) Correlations of clades and clines: geographic, elevational, and phylogenetic distribution patterns among Philippine mammals. In: Peters, G. & Hutterer, R. (Eds.), Vertebrates in the Tropics. Museum Alexander Koenig, Bonn, pp. 321–332.
- Heaney, L.R. & Roberts, T.E. (2009) New perspectives on the long-term biogeographic dynamics and conservation of Philippine fruit bats. In: Fleming, T.H. & Racey, P. (Eds.), Ecology, Evolution, and Conservation of Island Bats. University of Chicago Press, Chicago, pp. 17–58.
- Heaney, L.R., Tabaranza, B., Jr. Rickart, E.A., Balete, D.S. & Ingle, N.R. (2006) The mammals of Mt. Kitanglad Nature Park, Mindanao, Philippines. Fieldiana Zoology, 112, 1–63. https://doi.org/10.3158/0015-0754(2006)186[1:TMOMKN]2.0.CO;2
- Helgen, K.M., Kock, D., Gomez, R.K.S.C., Ingle, N.R. & Sinaga, M.H. (2007) Taxonomy and natural history of the Southeast Asian fruit-bat genus Dyacopterus. Journal of Mammalogy, 88, 302–318. https://doi.org/10.1644/06-MAMM-A-276R.1
- Hill, J.E. (1992) Order Chiroptera. In: Corbet, G.B. & Hill, J.E. (Eds.), The mammals of the Indomalayan region: a systematic review. Oxford University Press, Oxford, pp. 54–161.
- Ingle, N.R. & Heaney, L.R. (1992) A key to the bats of the Philippine Islands. Fieldiana Zoology, New Series, 69, 1–44.
- Jones, K.E., Purvis, A., MacLarnon, A., Bininda-Emonds, O.R.P. & Simmons, N.B. (2002) A phylogenetic supertree of the bats (Mammalia: Chiroptera). Biological Reviews, 77, 223–259. https://doi.org/10.1017/S1464793101005899
- Justiniano, R., Schenk, J.J., Balete, D.S., Rickart, E.A., Esselstyn, J.A., Heaney, L.R. & Steppan, S.J. (2015) Testing diversification models of endemic Philippine forest mice (Apomys) with nuclear phylogenies across elevational gradients reveals repeated colonization of isolated mountain ranges. Journal of Biogeography, 42, 51–64. https://doi.org/10.1111/jbi.12401
- Kingston, T., Jones, G., Akbar, Z. & Kunz, T.H. (1999) Echolocation signal design in Kerivoulinae and Murininae (Chiroptera: Vespertilionidae) from Malaysia. Journal of Zoology, 249, 359–374. https://doi.org/10.1017/S0952836999009917
- Kumar, S., Stecher, G. & Tamura, K. (2016) MEGA7: Molecular Evolutionary Genetics Analysis Version 7.0 for Bigger Datasets. Molecular Biology and Evolution, 33, 1870–1874. https://doi.org/10.1093/molbev/msw054
- Lim, B.K., Engstrom, M.D., Bickham, J.W. & Patton, J.C. (2008) Molecular phylogeny of New World sheath-tailed bats (Emballonuridae: Diclidurini) based on loci from the four genetic transmission systems in mammals. Biological Journal of the Linnean Society, 93, 189–209. https://doi.org/10.1111/j.1095-8312.2007.00942.x
- Mallari, N.A.D., Tabaranza, B.R., Jr. & Crosby, M.J. (2001) Key Conservation Sites in the Philippines. Bookmark, Inc., Makati City, 485 pp.
- Mammal Diversity Database (2025) Mammal Diversity Database. Version 2.0.. Zenodo. [data set] https://doi.org/10.5281/zenodo.15007505
- Mou, X., Qian, Y., Li, M., Li, B., Luo, X. & Li, S. (2024) A new species of Murina (Chiroptera: Vespertilionidae) from Yunnan, China. Animals, 14 (16), e2371. https://doi.org/10.3390/ani14162371
- Mou, X., Qian, Y., Wang, W., Zhang, W., Wang, J. & Song, L. (2025) A new ‘cyclotis-morphotype’ species of tube-nosed bat (Chiroptera: Vespertilionidae: Murina) from China. Animals, 15 (1), e75. https://doi.org/10.3390/ani15010075
- Musser, G.G. & Carleton, M.D. (2005) Superfamily Muroidea. In: Wilson, D. & Reeder, D. (Eds.), Mammal Species of the World, A Taxonomic and Geographic Reference. The Johns Hopkins University Press, Baltimore, Maryland, pp. 894–1531.
- Nesi, N., Tsagkogeorga, G., Tsang, S.M., Nicolas, V., Lalis, A., Scanlon, A.T., Riesle-Sbarbaro, S.A., Wiantoro, S., Hitch, A.T., Juste, J., Pinzari, C.A., Bonaccorso, F.J., Todd, C.M., Lim, B.K., Simmons, N.B., McGowen, M.R. & Rossiter, S.J. (2021) Interrogating phylogenetic discordance resolves deep splits in the rapid radiation of Old World fruit bats (Chiroptera: Pteropodidae). Systematic Biology, 70, 1077–1089. https://doi.org/10.1093/sysbio/syab013
- Ochoa, J., Mijares, A.S.B., Piper, P.J., Reyes, M.C. & Heaney, L.R. (2021) Three new extinct species from the endemic Philippine cloud rat radiation (Rodentia, Muridae, Phloeomyini). Journal of Mammalogy, 102, 909–930. https://doi.org/10.1093/jmammal/gyab023
- Peterson, A.T. & Heaney, L.R. (1993) Genetic differentiation in Philippine bats of the genera Cynopterus and Haplonycteris. Biological Journal of the Linnean Society, 49, 203–218. https://doi.org/10.1111/j.1095-8312.1993.tb00900.x
- Petrosky, A.L., Rowsey, D.M. & Heaney, L.R. (2021) Molecular assessment of dietary niche partitioning in an endemic island radiation of tropical mammals. Molecular Ecology, 30, 5858–5873. https://doi.org/10.1111/mec.16158
- Plaza, J.L. & Sanguila, M.B. (2015) Preliminary report on the anurans of Mount Hilong-hilong, Agusan Del Norte, Eastern Mindanao, Philippines. Asian Herpetological Research, 6, 18–33.
- Rickart, E.A., Balete, D.S., Alviola, P.A., Veluz, M.J. & Heaney, L.R. (2016) The mammals of Mt. Amuyao: A richly endemic fauna in the Central Cordillera of northern Luzon Island, Philippines. Mammalia, 80, 579–592. https://doi.org/10.1515/mammalia-2015-0132
- Rickart, E.A., Balete, D.S., Timm, R.M., Alviola, P.A., Esselstyn, J.A. & Heaney, L.R. (2019) Two new species of shrew-rats (Rhynchomys: Muridae: Rodentia) from Luzon Island, Philippines. Journal of Mammalogy, 100, 1112–1129. https://doi.org/10.1093/jmammal/gyz066
- Rickart, E.A., Heaney, L.R., Balete, D.S. & Tabaranza, B.R. (2011) Small mammal diversity along an elevational gradient in northern Luzon, Philippines. Mammalian Biology, 76, 12–21. https://doi.org/10.1016/j.mambio.2010.01.006
- Rickart, E.A., Heaney, L.R., Heideman, P.D. & Utzurrum, R.C.B. (1993) The distribution and ecology of mammals on Leyte, Biliran and Maripipi Islands, Philippines. Fieldiana Zoology, New Series, 72, 1–62. https://doi.org/10.5962/bhl.title.3152
- Rickart, E.A., Mercier, J.A. & Heaney, L.R. (1999) Cytogeography of Philippine bats. Proceedings of the Biological Society of Washington, 112, 453–469.
- Roberts, T.E. (2006a) History, ocean channels, and distance determine phylogeographic patterns in three widespread Philippine fruit bats (Pteropodidae). Molecular Ecology, 15, 2183–2199. https://doi.org/10.1111/j.1365-294X.2006.02928.x
- Roberts, T.E. (2006b) Multiple levels of allopatric divergence in the endemic Philippine fruit bat Haplonycteris fischeri (Pteropodidae). Biological Journal of the Linnean Society, 88, 329–349. https://doi.org/10.1111/j.1095-8312.2006.00616.x
- Rohlf, F.J. (2009) NTSYSpc: numerical taxonomy system. 2nd Edition. Exeter Publishing, Setauket, New York, 49 pp.
- Rowsey, D.M., Duya, M.R.M., Ibañez, J.C., Jansa, S.A., Rickart, E.A. & Heaney, L.R. (2022) A new genus and species of shrew-like mouse (Rodentia: Muridae) from a new center of endemism in eastern Mindanao, Philippines. Journal of Mammalogy, 103, 1259–1277. https://doi.org/10.1093/jmammal/gyac057
- Rowsey, D.M., Heaney, L.R. & Jansa, S.A. (2018) Diversification rates of the “Old Endemic” murine rodents of Luzon Island, Philippines are inconsistent with incumbency effects and ecological opportunity. Evolution, 72, 1420–1435. https://doi.org/10.1111/evo.13511
- Ruedi, M., Stadelmann, B., Gager, Y., Douzery, E.J.P., Francis, C.M., Lin, L.K., Guillén-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. https://doi.org/10.1016/j.ympev.2013.08.011
- Sarmiento, R.T. & Demetillo, M.T. (2017) Rapid assessment on tree diversity of Nickel Mining sites in Carrascal, Surigao del Sur, Philippines. Journal of Biodiversity and Environmental Sciences, 10 (4), 201–207.
- Schmieder, D.A., Kingston, T., Hashim, R. & Siemers, B.M. (2012) Sensory constraints on prey detection performance in an ensemble of vespertilionid understorey rain forest bats. Functional Ecology, 26, 1043–1053. https://doi.org/10.1111/j.1365-2435.2012.02024.x
- Sedlock, J.L. (2001a) Autecology and the conservation of insectivorous bats on Mt. Makiling, Philippines. Silliman Journal, 42, 163–201.
- Sedlock, J.L. (2001b) Inventory of insectivorous bats on Mount Makiling, Philippines using echolocation call signatures and a new tunnel trap. Acta Chiropterologica, 3, 163–178.
- Sedlock, J.L., Heaney, L.R., Balete, D.S. & Ruedi, M. (2020) Philippine bats of the genus Kerivoula (Chiroptera: Vespertilionidae): Overview and assessment of variation in K. pellucida and K. whiteheadi. Zootaxa, 4755 (3), 454–490. https://doi.org/10.11646/zootaxa.4755.3.2
- Sedlock, J.L., Jose, R.P., Vogt, J.M., Paguntalan, L.M.J. & Cariño, A.B. (2014) A survey of bats in a karst landscape in the central Philippines. Acta Chiropterologica, 16, 197–211. https://doi.org/10.3161/150811014X683390
- Sedlock, J.L. & Weyandt, S.E. (2009) Genetic divergence between morphologically and acoustically cryptic bats: Novel niche partitioning or recent contact? Journal of Zoology, 279, 388–395. https://doi.org/10.1111/j.1469-7998.2009.00634.x
- Sedlock, J.L., Weyandt, S.E., Cororan, L., Damerow, M., Hwa, S.H. & Pauli, B. (2008) Bat diversity in tropical forest and agro-pastoral habitats within a protected area in the Philippines. Acta Chiropterologica, 10 (2), 349–358. https://doi.org/10.3161/150811008X414926
- Sikes, R.S. & The Animal Care and Use Committee of the American Society of Mammalogists (2016) 2016 Guidelines of the American Society of Mammalogists for the use of wild mammals in research and education. Journal of Mammalogy, 97, 663–688. https://doi.org/10.1093/jmammal/gyw078
- Simmons, N.B. &Cirranello, A.L. (2024) Bat Species of the World: A Taxonomic and Geographical Database. Version 1.5. [https://batnames.org]
- Soisook, P., Karapan, S., Satasook, C. & Bates, P.J.J. (2013a) A new species of Murina (Mammalia: Chiroptera: Vespertilionidae) from peninsular Thailand. Zootaxa, 3746 (4), 567–579. https://doi.org/10.11646/zootaxa.3746.4.4
- Soisook, P., Karapan, S., Satasook, C., Thong, V.D., Khan, F.A.A., Maryanto, I., Csorba, G., Furey, N., Aul, B. & Bates, P.J.J. (2013b) A Review of the Murina cyclotis complex (Chiroptera: Vespertilionidae) with descriptions of a new species and subspecies. Acta Chiropterologica, 15, 271–292. https://doi.org/10.3161/150811013X678928
- Soisook, P., Thaw, W.N., Kyaw, M., Oo, S.S.L., Pimsai, A., Suarez-Rubio, M. & Renner, S.C. (2017) A new species of Murina (Chiroptera: Vespertilionidae) from sub-Himalayan forests of northern Myanmar. Zootaxa, 4320 (1), 159–172. https://doi.org/10.11646/zootaxa.4320.1.9
- Son, N.T., Csorba, G., Tu, V.T., Thong, V.D., Wu, Y., Harada, M., Oshida, T., Endo, H. & Motokawa, M. (2015) A new species of the genus Murina (Chiroptera: Vespertilionidae) from the Central Highlands of Vietnam with a review of the subfamily Murininae in Vietnam. Acta Chiropterologica, 17, 201–232. https://doi.org/10.3161/15081109ACC2015.17.2.001
- Steppan, S.J., Zawadzki, C. & Heaney, L.R. (2003) Molecular phylogeny of the endemic Philippine rodent Apomys (Muridae) and the dynamics of diversification in an oceanic archipelago. Biological Journal of the Linnean Society, 80, 699–715. https://doi.org/10.1111/j.1095-8312.2003.00274.x
- Tamura, K., Battistuzzi, F.U., Billing-Ross, P., Murillo, O., Filipski, A. & Kumar, S. (2012) Estimating divergence times in large molecular phylogenies. Proceedings of the National Academy of Sciences of the United States of America, 109, 19333–19338. https://doi.org/10.1073/pnas.1213199109
- Temminck, C.J. (1840) 1835–1841. Monographies de Mammalogie, ou description de quelques genres de Mammifères, dont les espèces ont été observées dans les différens musées de l’Europe. Tome Second. E. d’Ocagne et A. Bertrand, Paris and C.C. van der Hock, Leiden, 392 pp., pls. XXVI–LXX.
- Temminck, C.J. (1853) Esquisses Zoologie sur la Cote de Guine. 1. MammiferesBrill, . Leiden, xvi + 256 pp.
- Wang, X., Han, X., Csorba, G., Wu, Y., Chen, H., Zhao, X., Dong, Z., Yu, W. & Lu, Z. (2025) A new species of tube-nosed bat (Chiroptera: Vespertilionidae: Murina) from Qinghai-Tibet Plateau, China. Journal of Mammalogy, 10(1), 178–186. https://doi.org/10.1093/jmammal/gyae104
- Wang, X., Liang, X., Li, Y., Csorba, G., Hu, Y., He, K., Wu, Y. & Yu, W. (2024) Phylogenetic conflict between species tree and mitochondrial gene trees in Murininae (Mammalia, Chiroptera) and validation of the genus Harpiola. Zoologica Scripta, 54 (3), 305–316. https://doi.org/10.1111/zsc.12708
- Yu, W.H., Csorba, G. & Wu, Y. (2020) Tube-nosed variations—a new species of the genus Murina (Chiroptera: Vespertilionidae) from China. Zoological Research, 41, 70–77. https://doi.org/10.24272/j.issn.2095-8137.2020.009
- Zeng, X., Chen, J., Deng, H., Ning, X. & Zhou, J. (2018) A new species of Murina from China (Chiroptera: Vespertilionidae). Ekoloji, 27, 9–16.