Skip to main content Skip to main navigation menu Skip to site footer
Responsive image
Issue: Vol. 5590 No. 4: 25 Feb. 2025
Type: Article
Published: 2025-02-25
DOI: 10.11646/zootaxa.5590.4.3
Page range: 507-530
Abstract views: 297
PDF downloaded: 2

Taxonomic revision of the South Asian allies of Hipposideros galeritus Cantor, 1846

Wildlife Biology and Taxonomy Lab; Department of Zoology; Osmania University; Hyderabad 500007; Telangana State; India; Systematics; Ecology & Conservation Laboratory; Zoo Outreach Organization; 3A2 Varadharajulu Nagar; FCI Road; Ganapathy; Coimbatore 641006; Tamil Nadu; India
Department of Agricultural Biology; Faculty of Agriculture; University of Ruhuna; Mapalana; Kamburupitiya; Sri Lanka
Ecology and Conservation Lab; School of Biological Sciences; University of Reading; Whiteknights; Reading; United Kingdom
Department of Biological Sciences; Faculty of Applied Sciences; Rajarata University of Sri Lanka; Mihintale; Sri Lanka
Department of Zoology and Environment Sciences; Faculty of Science; University of Colombo; Colombo; Sri Lanka
Department of Zoology and Environment Sciences; Faculty of Science; University of Colombo; Colombo; Sri Lanka
Centre for Biodiversity and Conservation Studies; Osmania University; Hyderabad 500007; Telangana State; India
Department of Agricultural Biology; Faculty of Agriculture; University of Ruhuna; Mapalana; Kamburupitiya; Sri Lanka
Princess Maha Chakri Sirindhorn Natural History Museum; Prince of Songkla University; Hat Yai; Songkhla 90110; Thailand
Mammal & Osteology Section; Zoological Survey of India; M-Block; New Alipore; Kolkata 700053; West Bengal; India
Department of Zoology and Environment Sciences; Faculty of Science; University of Colombo; Colombo; Sri Lanka
Wildlife Biology and Taxonomy Lab; Department of Zoology; Osmania University; Hyderabad 500007; Telangana State; India; Centre for Biodiversity and Conservation Studies; Osmania University; Hyderabad 500007; Telangana State; India
Mammalia Chiroptera Hipposideros brachyotus Hipposideros srilankaensis new species India Sri Lanka

Abstract

Hipposideros galeritus was described in 1846, with subsequent studies suggesting four subspecies across South and Southeast Asia. Our study indicates that the Indian and Sri Lankan populations previously considered subspecies of H. galeritus are, in fact, distinct species in need of taxonomic revisions. Based on the morphometric analysis, structure of the baculum, molecular phylogenetics, and echolocation call analysis, the Indian, Sri Lankan, and Southeast Asian populations of Hipposideros galeritus are distinct. We provide a detailed description of H. brachyotus Dobson, 1874 and describe a new species—H. srilankaensis sp. nov.—from Sri Lanka. Key morphological differences were found in the noseleaf, ear shape, and cranial features between the Indian (H. brachyotus), Sri Lankan (H. srilankaensis sp. nov.), and Southeast Asian populations (H. galeritus s.l.). Substantial genetic distances were found between H. galeritus populations, suggesting cryptic diversity that is yet to be resolved.

 

References

  1. Agarwal, I., Biswas, S., Bauer, A.M., Greenbaum, E., Jackman, T.R., De Silva, A. & Batuwita, S. (2017) Cryptic species, taxonomic inflation, or a bit of both? New species phenomenon in Sri Lanka as suggested by a phylogeny of dwarf geckos (Reptilia, Squamata, Gekkonidae, Cnemaspis). Systematics and Biodiversity, 15 (5), 427–439. https://doi.org/10.1080/14772000.2017.1282553
  2. Bahir, M. & Ng, P. (2005) Descriptions of ten new species of freshwater crabs (Crustacea: Brachyura: Parathelphusidae: Ceylonthelphusa, Mahatha, Perbrinckia) from Sri Lanka. Raffles Bulletin of Zoology, Supplement 12, 47–75.
  3. Bates, P.J.J. & Harrison, D.L. (1997) The Bats of the Indian Subcontinent. Harrison Zoological Museum Publications, Sevenoaks, 258 pp.
  4. Bates, P.J.J., Rossiter, S.J., Suyanto, A. & Kingston, T. (2007) A new species of Hipposideros (Chiroptera: Hipposideridae) from Sulawesi. Acta Chiropterologica, 9 (1), 13–26. https://doi.org/10.3161/1733-5329(2007)9[13:ANSOHC]2.0.CO;2
  5. Batuwita, S. & Bahir, M.M. (2005) Description of five new species of Cyrtodactylus (Reptilia: Gekkonidae) from Sri Lanka. Raffles Bulletin of Zoology, Supplement 12, 351–380.
  6. Bauer, A.M., Jackman, T.R., Greenbaum, E., Giri, V.B. & de Silva, A. (2010) South Asia supports a major endemic radiation of Hemidactylus geckos. Molecular Phylogenetics and Evolution, 57, 343–352. https://doi.org/10.1016/j.ympev.2010.06.014
  7. Beenaerts, N., Pethiyagoda, R., Ng, P., Yeo, D., Bex, G., Bahir, M. & Artois, T. (2010) Phylogenetic diversity of Sri Lankan freshwater crabs and its Implications for conservation. Molecular Ecology, 19, 183–196. https://doi.org/10.1111/j.1365-294X.2009.04439.x.
  8. Biswas, S. & Pawar, S.S. (2006) Phylogenetic tests of distribution patterns in South Asia: towards an integrative approach. Journal of Biosciences, 31, 95–113. https://doi.org/10.1007/BF02705240
  9. Blanford, W.T. (1887) Critical Notes on the Nomenclature of Indian Mammals. Proceedings of the Zoological Society of London, 55 (4), 620–638. https://doi.org/10.1111/j.1469-7998.1887.tb08156.x
  10. Blanford, W.T. (1888–91) The Fauna of British India, Mammalia. Taylor & Francis, London, 617 pp.
  11. Bossuyt, F., Meegaskumbura, M., Beenaerts, N., Gower, D.J., Pethiyagoda, R., Roelants, K., Mannaert, A., Wilkinson, M., Bahir, M.M., Manamendra-Arachchi, K., Ng, P.K., Schneider, C.J., Oommen, O.V. & Milinkovitch, M.C. (2004) Local endemism within the Western Ghats-Sri Lanka biodiversity hotspot. Science, 306, 479–481. https://doi.org/10.1126/science.1100167
  12. Bruford, M.W., Hanotte, O., Brookfield, J.F.Y. & Burke, T. (1992) Single-locus and multilocus DNA fingerprinting. In: Hoelzel, A.R. (ed.) Molecular genetic analysis of populations: A practical approach. Oxford Univeristy Preess, Oxford, pp. 225–269.
  13. Cantor, T. (1846) Catalogue of Mammalia inhabiting the Malayan Peninsula and islands. Journal of the Asiatic Society of Bengal, 15 (171), 171–203.
  14. Cooper, A., Turney, C., Hughen, K.A., Brook, B.W., McDonald, G.H. & Bradshaw, C.J.A. (2015) Abrupt warming events drove Late Pleistocene Holarctic megafaunal turnover. Science, 349, 602–606. https://doi.org/10.1126/science.aac4315
  15. Cooray, P.G. (1984) The Geology of Sri Lanka (Ceylon). National Museums of Sri Lanka Publication, Colombo, 340 pp. https://doi.org/10.1017/S0016756800018148
  16. Csorba, G., Ujhelyi, P. & Thomas, N. (2003) Horseshoe Bats of the World. Alana Books. Bishop’s Castle, 160 pp.
  17. Darriba, D., Taboada, G.L., Doallo, R. & Posada, D. (2012) jModelTest 2: more models, new heuristics and parallel computing. Nature Methods, 9 (8), 772. https://doi.org/10.1038/nmeth.2109
  18. Debata, S. & Palita, S.K. (2017) Distribution, population status, and threats of nationally threatened Cantor’s leaf-nosed bat Hipposideros galeritus Cantor, 1846 in eastern India. Journal of Asia-Pacific Biodiversity, 10 (3), 426–430. https://doi.org/10.1016/j.japb.2017.05.003
  19. Debata, S., Palei, H.S., Mohapatra, P.P. & Palita, S.K. (2015) Additional records of Cantor’s Leaf-Nosed Bat Hipposideros galeritus Cantor, 1846 (Mammalia: Chiroptera: Hipposideridae) in eastern India: Odisha. Journal of Threatened, Taxa 7 (8), 7477–7479. https://doi.org/10.11609/JoTT.o4247.7477-9
  20. Dobson, G.E. (1874) Descriptions of new species of Chiroptera from India and Yunnan. Journal of the Asiatic Society of Bengal, 63, Pt. 2 (4), 237–238.
  21. Dobson, G.E. (1876) Monograph of the Asiatic Chiroptera and catalogue of the species of bats in the collection of the Indian Museum Calcutta. Printed by Order of the Trustees of the Indian Museum, London, 228 pp.
  22. Donnelly, M., Martin, T.E., Cropper, O., Yusti, E., Arfian, A., Smethurst, R., Fox, C., Pryde, M., Hafirun, Phangurha, J., van der Aar, R.N., Hutchison, A., Karya, A., Analuddin, K., Samsudin & Courtney Jones, S.K. (2021) New species records from Buton Island, South East Sulawesi, including regional range extensions. Journal of Bat Research and Conservation, 14 (1), 14–32. https://doi.org/10.14709/BarbJ.14.1.2021.03
  23. Edgar, R.C. (2004) MUSCLE: multiple sequence alignment with high accuracy and high throughput. Nucleic Acids Research, 32 (5), 1792–97. https://doi.org/10.1093/nar/gkh340
  24. Ellerman, J.R. & Morrison-Scott, T.C.S. (1951) Checklist of palaearctic and Indian mammals 1758 to 1946. Trustees of the British Museum, London, iv + 810 pp.
  25. Faurby, S. & Svenning, J. (2016) Resurrection of the island rule: Human driven extinctions have obscured a basic evolutionary pattern. American Naturalist, 187, 812–820. https://doi.org/10.1086/686268
  26. Foster, J.B. (1964) Evolution of mammals on islands. Nature, 202, 234–235. https://doi.org/10.1038/202234a0
  27. Francis, C.M. & Habersetzer, J. (1998) Interspecific and intraspecific variation in the echolocation call frequency and morphology of horseshoe bats, Rhinolophus and Hipposideros. In: Kunz, T.H. & Racey, P.A. (Eds.), Bat Biology and Conservation, Smithsonian Institution Press, Washington, D.C., pp. 169–179.
  28. Francis, C.M. (2019) Field Guide to the Mammals of Southeast Asia, Second edition. Bloomsbury Publishing, London, 416 pp.
  29. Ghosh, M.K. (2008) Catalogue of Chiroptera in the collection of Zoological Survey of India (Part-II: Microchiroptera). In: Records of the Zoological Survey of India. Occasional Paper No. 281. Published by the Director, Zoological Survey of India, Kolkata, pp. 1–339.
  30. Gray, J.E. (1847) Characters of six new genera of bats not hitherto distinguished. Proceedings of the Zoological Society, London, 1847, 14–16.
  31. Grismer, J.L., Schulte, J.A., Alexander, A., Wagner, P., Travers, S.L., Buehler, M.D., Welton, L.J. & Brown, R.M. (2016) The Eurasian invasion: phylogenomic data reveal multiple Southeast Asian origins for Indian Dragon Lizards. BMC Evolutionary Biology, 16, 43. https://doi.org/10.1186/s12862-016-0611-6
  32. Hall, T.A. (1999) BioEdit: A User-Friendly Biological Sequence Alignment Editor and Analysis Program for Windows 95/98/ NT. Nucleic Acids Symposium Series, 41, 95–98.
  33. Hasegawa, M., Kishino, H. & Yano, T. (1985) Dating of the human-ape splitting by a molecular clock of mitochondrial DNA. Journal of Molecular Evolution, 22, 160–174. https://doi.org/10.1007/BF02101694
  34. Hill, J.E. (1963) A revision of the genus Hipposideros. Bulletin British Museum of Natural History, Zoology, 11, 1–129. https://doi.org/10.1126/science.aac4315
  35. Hughes, A.C., Satasook, C., Bates, P.J.J., Soisook, P., Sritongchuay, T., Jones, G. & Bumrungsri, S. (2010) Echolocation call analysis and presence-only modelling as conservation monitoring tools for rhinolophoid bats in Thailand. Acta Chiropterologica, 12, 311–327. https://doi.org/10.3161/150811010X537891
  36. Ivanova, N.V., Dewaard, J.R. & Hebert P.D.N. (2006) An inexpensive, automation-friendly protocol for recovering high-quality DNA. Molecular Ecology Notes, 6, 998–1002. https://doi.org/10.1111/j.1471-8286.2006.01428.x
  37. Jenkins, P.D. & Hill, J.E. (1981) The status of Hipposideros galeritus Cantor, 1846 and Hipposideros cervinus (Gould, 1854) (Chiroptera: Hipposideridae). Bulletin of the Natural History Museum, Zoology, 41 (5), 279–294.
  38. Jinggong, E.R., Rajasegaran, P., Morni, M.A., William-Dee, J., Denel, A. & Anwarali Khan, F.A. (2022) Bats of Mount Silabur, Sarawak, and Its Potential for Conservation. Borneo Journal of Resource Science and Technology, 12 (1), 141–156. https://doi.org/10.33736/bjrst.4235.2022
  39. Jukar, A.M., Lyons, S.K., Wagner, P.J. & Uhen, M.D. (2020) Late Quaternary extinctions in the Indian Subcontinent. Palaeogeography, Palaeoclimatology, Palaeoecology, 562, 110137. https://doi.org/10.1016/j.palaeo.2020.110137
  40. Katupotha, J. (1995) Evolution and Geological Significance of late Pleistocene Fossil Shell Beds of the Southern Coastal Zone of Sri Lanka. Natural Resources, Energy and Science Authority of Sri Lanka, Colombo, 63 pp.
  41. Katupotha, J. (2013) Palaeoclimate change during Glacial Periods: Evidence from Sri Lanka. Journal of Tropical Forestry and Environment, 3, 42–54. https://doi.org/10.31357/jtfe.v3i1.1122
  42. Khajuria, H. (1980) Taxonomical and ecological studies on bats of Jabalpur district Madhya Pradesh, India. Part 1 (Families Megadermatidae, Rhinolophidae and Vespertilionidae). Records of the Zoological Survey of India, Miscellaneous Publications, Occasional Paper, No. 19, 1–69.
  43. Kruskop, S.V. (2014) Baculums of chiropterans of Indochina: Rhinolophoids (Chiroptera: Rhinolophidae, Hipposideridae). Plecotus et al. Mammalia, 17 (2014), 3–17.
  44. Krzanowski, A. (1967) The magnitude of islands on the size of bats (Chiroptera). Acta Zoologica Cracoviensia, 12, 281–346.
  45. Kumar, S., Stecher, G., Li, M., Knyaz, C. & Tamura, K. (2018) MEGA X: Molecular Evolutionary Genetics Analysis across Computing Platforms. Molecular Biology and Evolution, 35 (6), 1547–1549. https://doi.org/10.1093/molbev/msy096
  46. Kusuminda, T., Mannakkara, A., Amarasinghe, C.J., Karunarathna, K.A.P.M.K., Patterson, B.D. & Yapa, W.B. (2019) Echolocation call characteristics of Hipposiderid bats in Sri Lanka. Rajarata International Research Conference, Mihintale, Sri Lanka, November 2019, 218 pp.
  47. Kusuminda, T., Mannakkara, A., Gamage, R., Patterson, B.D.& Yapa, W.B. (2022a). Roosting ecology of insectivorous bats in a tropical agricultural landscape. Mammalia, 86 (2), 134-143. https://doi.org/10.1515/mammalia-2021-0056
  48. Kusuminda, T., Mannakkara, A., Ukuwela, K.D.B., Kruskop, S.V., Amarasinghe, C.J., Saikia, U., Venugopal, P., Karunarathna M., Gamage, R., Ruedi, M., Csorba, G., Yapa, W.B. & Patterson, B.D. (2022b) DNA barcoding and morphological analyses reveal a cryptic species of Miniopterus from India and Sri Lanka. Acta Chiropterologica, 24 (1), 1–17. https://doi.org/10.3161/15081109ACC2022.24.1.001
  49. Lajmi, A., Bansal, R., Giri, V.B. & Karanth, P. (2019) Phylogeny and biogeography of the endemic Hemidactylus geckos of the Indian subregion suggest multiple dispersals from Peninsular India to Sri Lanka. Zoological Journal of the Linnean Society, 186, 286–301. https://doi.org/10.1093/zoolinnean/zly047
  50. Leach, W.E. (1816) Systematic catalogue of the specimens of the indigenous mammalia and birds that are preserved in the British Museum: with their localities and authorities, to which are added a list of the described species that are wanting to complete the collection of British Mammalia and Birds. Richard and Arthur Taylor, London, 42 pp.
  51. Lomolino, M.V. (2005) Body size evolution in insular vertebrates: generality of the island rule. Journal of Biogeography, 32, 1683–1699. https://doi.org/10.1111/j.1365-2699.2005.01314.x
  52. Manamendra-Arachchi, K., Batuwita, S. & Pethiagoda, R. (2007) A taxonomic revision of the Sri Lankan day-geckos (Reptilia: Gekkonidae: Cnemaspis), with description of new species from Sri Lanka and southern India. Zeylanica, 7 (1), 9–122.
  53. Meiri, S., Cooper, N. & Purvis, A. (2008) The island rule: made to be broken? Conservation Genetics, 275, 141–148. https://doi.org/10.1098/rspb.2007.1056
  54. Molinari, J. (2023) A global assessment of the ‘island rule’ in bats based on functionally distinct measures of body size. Journal of Biogeography, 50, 1179–1190. https://doi.org/10.1111/jbi.14624
  55. Monadjem, A. (2019) Family Hipposideridae (Old World leaf-nosed bats). In: Wilson, D.E., Mittermeier, R.A. (Eds.), Handbook of the Mammals of the World. Vol. 9. Bats. Lynx Ediciones, Barcelona, pp. 210–227.
  56. Pearson, D.L. & Ghorpade, K. (1989) Geographical distribution and ecological history of tiger beetles (Coleoptera: Cicindelidae) of the Indian subcontinent. Journal of Biogeography, 1, 333–344. https://doi.org/10.2307/2845225
  57. Pethiyagoda, R. & Sudasinghe, H. (2021) The Ecology and Biogeography of Sri Lanka: A context for freshwater fishes. WHT Publications (Private) Limited, Colombo, xiv + 258 pp.
  58. Pfenninger, M. & Schwenk, K. (2007) Cryptic animal species are homogenously distributed among taxa and biogeographical regions. BMC Evolutionary Biology, 7, 121. https://doi.org/10.1186/1471-2148-7-121
  59. Phauk, S., Phen S. & Furey, N.M. (2013) Cambodian bat echolocation: a first description of assemblage call parameters and assessment of their utility for species identification. Cambodian Journal of Natural History, 2013, 16–26.
  60. Phillips, W.W.A. (1923) Further notes on some Ceylon bats. Journal Bombay Natural History Society, 29, 154–156.
  61. Phillips, W.W.A. (1980) Manual of the mammals of Sri Lanka. Part I. Wildlife and Nature Protection Society of Sri Lanka, 1980, 1–116.
  62. Pyron, R.A., Burbrink, F.T. & Wiens, J.J. (2013) A phylogeny and revised classification of Squamata, including 4161 species of lizards and snakes. BMC Evolutionary Biology, 13: 93. https://doi.org/10.1186/1471-2148-13-93
  63. R Core Team (2023) R: A language and environment for statistical computing. R Foundation for Statistical Computing, Vienna. Available from: https://www.R-project.org/ (accessed 8 January 2024)
  64. Rambaut, A., Suchard, M.A. & Drummond, A.J. (2013) Tracer. Version 1.6. Software. Available from: https://tree.bio.ed.ac.uk/software/tracer/ (accessed 8 January 2024)
  65. Rambaut, A. (2018) Figtree. Version 1.4.4. Institute of Evolutionary Biology, University of Edinburgh, Edinburgh. [program]
  66. Ripley, S.D. & Beehler, B.M. (1990) Patterns of speciation in Indian birds. Journal of Biogeography, 1, 639–648. https://doi.org/10.2307/2845145
  67. Ronquist, F., Teslenko, M., van der Mark, P., Ayres, D.L., Darling, A., Höhna, S., Larget, B., Liu, L., Suchard, M.A. & Huelsenbeck, J.P. (2012) MrBayes 3.2: efficient Bayesian phylogenetic inference and model choice across a large model space. Systematic Biology, 61 (3), 539–542. https://doi.org/10.1093/sysbio/sys029
  68. Safi, K., Meiri, S. & Jones, K.E. (2013) Evolution of body size in bats. In: Smith, F.A. & Lyons, S.K. (Eds.), Animal body size: linking pattern and process across space and time, and taxonomic group. University of Chicago Press, Chicago, Illinois, pp. 95–115. https://doi.org/10.7208/chicago/9780226012285.003.0005
  69. Sazali, S.N. & Aziz, N.S.A. (2012) Morphometric analysis of Cantor’s Roundleaf bat Hipposideros galeritus 1846 from several localities in Sarawak, Malaysia. Journal of Tropical Biology and Conservation, 9 (1), 97–104. https://doi.org/10.51200/jtbc.v9i.235
  70. Sclater, P.L. (1901) The Mammals of South Africa. Rodentia, Chiroptera, Insectivora, Cetacea, Edentata. Vol. 2. R. H. Porter Publications, London, 515 pp.
  71. Simmons, N.B. (2005) Chiroptera. In: Wilson, D.E. & Reeder, D.A.M. (Eds.) Mammal Species of the World: A Taxonomic and Geographic Reference. 3rd Edition. Johns Hopkins University Press, Baltimore, Maryland, pp. 312–529.
  72. Srinivasulu, B. & Srinivasulu, A. (2023) A new species of the Miniopterus australis species complex (Chiroptera: Miniopteridae) from the Western Ghats, India. Zootaxa, 5296 (2), 233–249. https://doi.org/10.11646/zootaxa.5296.2.5
  73. Srinivasulu, B. & Srinivasulu, C. (2017) A first record of three hitherto unreported species of bats from Kerala, India with a note on Myotis peytoni (Mammalia: Chiroptera: Vespertilionidae). Journal of Threatened Taxa, 9 (5), 10216–10222. https://doi.org/10.11609/jott.3324.9.5.10216-10222
  74. Srinivasulu, B., Srinivasulu, C. & Kaur, H. (2015) Echolocation calls of four species of leaf-nosed bats (Genus Hipposideros) from Central Peninsular India. Current Science, 108 (6), 1055–1057.
  75. Srinivasulu, C. & Srinivasulu, B. (2005) A review of chiropteran diversity of Bangladesh. BAT NET-CCINSA Newsletter, 6, 6–11.
  76. Srinivasulu, C. & Srinivasulu, B. (2012) South Asian mammals: their diversity, distribution, and status. Springer New York, New York, xi + 467 pp. https://doi.org/10.1007/978-1-4614-3449-8
  77. Srinivasulu, C. (2004) Cantor’s Round Leaf Bat Hipposideros galeritus Cantor, 1846: An addition to chiropteran diversity of Andhra Pradesh, India. BAT NET-CCINSA Newsletter, 5 (1), 4–5.
  78. Sudasinghe, H., Adamson, E.A., Ranasinghe, R.T., Meegaskumbura, M., Ikebe, C. & Britz, R. (2020a) Unexpected species diversity within Sri Lanka’s snakehead fishes of the Channa marulius group (Teleostei: Channidae). Zootaxa, 4747 (1), 113–132. https://doi.org/10.11646/zootaxa.4747.1.4
  79. Sudasinghe, H., Ranasinghe, R.H.T., Pethiyagoda, R., Meegaskumbura, M. & Britz, R. (2020b) Genetic diversity and morphological stasis in the Ceylon Snakehead, Channa orientalis (Teleostei: Channidae). Ichthyological Research, 2020, 1–14. https://doi.org/10.1007/s10228-020-00768-0
  80. Talavera, G. & Castresana, J. (2007) Improvement of phylogenies after removing divergent and ambiguously aligned blocks from protein sequence alignments. Systematic Biology, 56, 564–577. https://doi.org/10.1080/10635150701472164
  81. Tate, G.H.H. (1941) A review of the genus Hipposideros with special reference to Indo-Australian species. Bulletin of the American Museum of Natural History, 78, 353–393.
  82. Thong, V.D. (2015) Bats of Cat Tien National Park: diversity, echolocation and taxonomic remarks. Tap Chi Sinh Hoc, 37, 336–343. https://doi.org/10.15625/0866-7160/v37n3.7418
  83. Tingga, R.C.T., Anwarali, F.A., Ridwan, A.R.M., Senawi, J. & Abddulah, M.T. (2012) Small mammals from Kuala Atok, Taman Negara Pahang, Malaysia. Sains Malaysiana, 41 (6), 659–669.
  84. Topal, G. (1958) Morphological studies on the os penis of bats in the Carpathian Basin. Annales historico-naturales Musei nationalis hungarici, 1958, Series Nova, 9, 331–342.
  85. Van Valen, L. (1973) Body size and numbers of plants and animals. Evolution, 27, 27–35. https://doi.org/10.1111/j.1558-5646.1973.tb05914.x
  86. Wu, W., Ng, W.L., Yang, J.X., Li, W.M. & Ge, X.J. (2018) High cryptic species diversity is revealed by genome-wide polymorphisms in a wild relative of banana, Musa itinerans, and implications for its conservation in subtropical China. BMC Plant Biology, 18, 194. https://doi.org/10.1186/s12870-018-1410-6
  87. Yapa, W.B. (2017) A field guide to the bats of Sri Lanka. Dilmah Ceylon Tea Company PLC, Colombo, 142 pp.