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An integrative taxonomic review of the South Asian microhylid genus Uperodon



Based on a recent molecular phylogenetic study, the South Asian microhylid genus Uperodon (subfamily Microhylinae) currently comprises of 12 valid species that are largely restricted to India and Sri Lanka. Considering the revised generic-level status of its various members, here we review the taxonomy of all known species in this genus and clarify their nomenclatural status and geographical distribution, by integrating evidence from genetics, adult and tadpole morphology, breeding ecology, and bioacoustics. Our molecular analyses of a mitochondrial 16S rRNA gene fragment combined with external and internal morphological studies also revealed a distinct new species in the genus. This species, formally described as Uperodon rohani sp. nov., is endemic to Sri Lanka and widely distributed at lower elevations in the island. For nomenclatural stability of various previously known members, the following actions are also undertaken: (1) redescription of the poorly-defined species Ramanella anamalaiensis Rao (= Uperodon anamalaiensis) and Hylaedactylus montanus Jerdon (= Uperodon montanus); (2) neotype designation for Ramanella anamalaiensis Rao (= Uperodon anamalaiensis), Ramanella minor Rao, Ramanella mormorata Rao (= Uperodon mormorata), and Ramanella triangularis rufeventris Rao; (3) lectotype designation for Callula variegata Stoliczka (= Uperodon variegatus); and (4) synonymization of Ramanella minor Rao with Uperodon anamalaiensis.



Amphibia, Amphibians, bioacoustics, endemism, mitochondrial DNA, natural history, neotype, lectotype, new species, tadpoles, Western Ghats-Sri Lanka biodiversity hotspot


Abraham, S.K., Easa, P.S., Jahas, S.S. & Shaji, C.P. (2001) Amphibian fauna of Wayanad, Kerala. Zoos’ Print Journal, 16 (4), 457–461.

Andrews, M.I, George, S. & Joseph, J. (2005) Amphibians in protected areas of Kerala. Fauna of Protected Areas 20. Zoo’s Print Journal, 20 (4), 1823–1831.

Altig, R. & Johnston, G.F. (1989) Guilds of anuran larvae: relationships among developmental modes, morphologies, and habitats. Herpetological monographs, 3, 81–109.

Altig, R. & McDiarmid, R.W. (1999a) Body plan: development and morphology. In: McDiarmid, R.W. & Altig, R. (Eds.), Tadpoles: The Biology of Anuran larvae. The University of Chicago Press, Chicago, pp. 24–51.

Altig, R. & McDiarmid, R.W. (1999b) Diversity: familial and generic characterizations. In: McDiarmid, R.W. & Altig, R. (Eds.), Tadpoles: The Biology of Anuran larvae. The University of Chicago Press, Chicago, pp. 295–337.

Bauer, A.M. (1998) South Asian herpetological specimens of historical note in the Zoological Museum, Berlin. Hamadryad, 23 (2), 133–149.

Bee, M.A., Suyesh, R. & Biju, S.D. (2013) Vocal behavior of the Ponmudi Bush Frog (Raorchestes graminirupes): repertoire and individual variation. Herpetologica, 69 (1), 22–35.

Biju, S.D. (2001) A synopsis to the frog fauna of the Western Ghats, India. ISBC - Occasional Publication, 1, 1–24.

Biju, S.D. & Bossuyt, F. (2006) Two new species of Philautus (Anura, Ranidae, Rhacophorinae) from the Western Ghats, India. Amphibia-Reptilia, 27 (1), 1–9.

Biju, S.D., Garg, S., Gururaja, K.V., Shouche, Y. & Walujkar, S.A. (2014a) DNA barcoding reveals unprecedented diversity in Dancing Frogs of India (Micrixalidae, Micrixalus): a taxonomic revision with description of 14 new species. Ceylon Journal of Science (Biological Sciences), 43 (1), 37–123.

Biju, S.D., Garg, S., Mahony, S., Wijayathilaka, N., Senevirathne, G. & Meegaskumbura, M. (2014b) DNA barcoding, phylogeny and systematics of Golden-backed frogs (Hylarana, Ranidae) of the Western Ghats-Sri Lanka biodiversity hotspot, with the description of seven new species. Contributions to Zoology, 83 (4), 269–335.

Biju, S.D., Senevirathne, G., Garg, S., Mahony, S., Kamei, R.G., Thomas, A., Shouche, Y., Raxworthy, C.J., Meegaskumbura, M. & Van Bocxlaer, I. (2016) Frankixalus, a new rhacophorid genus of tree hole breeding frogs with oophagous tadpoles. PLoS ONE, 11 (1), e0145727.

Blackburn, D.C., Siler, C.D., Diesmos, A.C., McGuire, J.A., Cannatella, D.C. & Brown, R.M. (2013) An adaptive radiation of frogs in a Southeast Asian island archipelago. Evolution, 67 (9), 2631–2646.

Bossuyt, F. & Milinkovitch, M.C. (2000) Convergent adaptive radiations in Madagascan and Asian ranid frogs reveal covariation between larval and adult traits. Proceedings of the National Academy of Sciences, 97 (12), 6585–6590.

Bowatte, G. & Meegaskumbura, M. (2012) Morphology and ecology of tadpoles of Ramanella obscura (Anura: Microhylidae). Ceylon Journal of Science, Biological Sciences, 40 (2), 109–120.

Boulenger, G.A. (1882) Catalogue of the Batrachia Salientia s. Ecaudata in the Collection of the British Museum. Second edition. Taylor and Francis, London, 503 pp.

Boulenger, G.A. (1890) The fauna of British India, including Ceylon and Burma. Reptilia and Batrachia. Taylor and Francis, London, 541 pp.

Cannatella, D. (1999) Architecture: cranial and axial musculoskeleton. In: McDiarmid, R.W. & Altig, R. (Eds.), Tadpoles: The Biology of Anuran Larvae. University of Chicago Press, Chicago, pp. 52–91.

Chanda, S.K., Das, I. & Dubois, A. (2000) Catalogue of amphibian types in the collection of the Zoological Survey of India. Hamadryad, 25 (2), 100–128.

Chandrasegaran, V. & Ranawana, K.B. (2002) Study of Breeding Ecology of Ramanella obscura (Anura: Microhylidae) in natural tree holes, at Gannoruwa Forest Reserve, Sri Lanka. Proceedings of the Annual Research Sessions, University of Peradeniya, Sri Lanka, 7, 122.

Charif, R.A., Waack, A.M. & Strickman, L.M. (2010) Raven Pro 1.4 User’s Manual. Cornell Lab of Ornithology, Ithaca, 367 pp.

Daniels, R.J.R. (2005) Amphibians of Peninsular India. Universities Press (India), Hyderabad, 268 pp.

Das, I. & Whitaker, R. (1997) A redescription of Ramanella mormorata Rao, 1937 (Anura, Microhylidae). Alytes, 15 (3), 127–132.

de Sá, R.O., Streicher, J.W., Sekonyela, R., Forlani, M.C., Loader, S.P., Greenbaum, E., Richards, S. & Haddad, C.F. (2012) Molecular phylogeny of microhylid frogs (Anura: Microhylidae) with emphasis on relationships among New World genera. BMC Evolutionary Biology, 12 (1), 241.

Deuti, K. & Goswami, B.C.B. (1995) A field-guide to the Amphibians of West Bengal plains. WWF-India, Calcutta, 53 pp.

Dey, M. (2001) A study on the ecological status of the anurans of Barak Valley, Assam, India. Thesis submitted to Assam University, Silchar, 291 pp.

Donnelly, M.A., de Sa, R.O. & Guyer, C. (1990) Description of the tadpoles of Gastrophryne pictiventris and Nelsonophryne aterrima (Anura: Microhylidae), with a review of morphological variation in free-swimming microhylid larvae. American Museum Novitates, 2976, 1–19.

Dubois, A. (1984) Note preliminaire sur le groupe de Rana limnocharis Gravenhorst, 1829 (Amphibiens, Anoures). Alytes, 3 (4), 143–159.

Duellman, W.E. & Trueb, L. (1986) Biology of the Amphibians. McGraw-Hill, New York, 670 pp.

Duméril, A.M.C. & Bibron, G. (1841) Erpétologie Générale ou Histoire Naturelle complete des Reptiles. Roret, Paris, 430 pp.

Dutta, S.K. (1997) Amphibians of India and Sri Lanka, (Checklist and Bibliography). Odyssey Publishing House, Bhubaneswar, 342 pp.

Dutta, S.K. & Manamendra-Arachchi, K. (1996) The Amphibian fauna of Sri Lanka. Wildlife Heritage Trust of Sri Lanka, Colombo, 230 pp.

Frank, N. & Ramus, E. (1995) Complete Guide to Scientific and Common Names of Amphibians and Reptiles of the World. N. G. Publishing Inc, Pottsville, Pennsylvania, 377 pp.

Frost, D.R. (2017) Amphibian species of the world: an online reference. Version 6.0. American Museum of Natural History, New York, USA. Available from: (accessed 24 January 2017)

Ganesh, S.R. (2012) Arboreal behaviour in the Indian Painted Frog Kaloula taprobanica parker, 1934. Herpetotropicos, 8 (1–2), 67–70.

Garg, S. & Biju, S.D. (2016) Molecular and morphological study of Leaping Frogs (Anura, Ranixalidae) with description of two new species. PLoS ONE, 11, e0166326.

Garg, S. & Biju, S.D. (2017) Description of four new species of Burrowing Frogs in the Fejervarya rufescens complex (Dicroglossidae) with notes on morphological affinities of Fejervarya species in the Western Ghats. Zootaxa, 4277 (4), 451–490.

Garg, S., Suyesh, R., Sukeshan, S. & Biju, S.D. (2017) Seven new species of Night Frogs (Anura, Nyctibatrachidae) from the Western Ghats Biodiversity Hotspot of India, with remarkably high diversity of diminutive forms. PeerJ, 5, e3007.

Gosner, K.L. (1960) A simplified table for staging anuran embryos and larvae with notes on identification. Herpetologica, 16 (3), 183–190.

Grillitsch, B., Grillitsch, H., Dubois, A. & Splechtna, H. (1993) The tadpoles of the brown frogs Rana (graeca) graeca and Rana (graeca) italica (Amphibia, Anura). Alytes, 11 (4), 117–139.

Günther, A.C.L.G. (1876 “1875”) Third report on collections of Indian reptiles obtained by the British Museum. Proceedings of the Zoological Society of London, 1875, 567–577.

Günther, A.C.L.G. (1864 “1863”) The Reptiles of British India. Ray Society by R. Hardwicke, London, 452 pp.

Hanken, J. & Wassersug, R.J. (1981) The visible skeleton. Functional Photography, 16 (4), 22–26.

Harpalani, M., Kanagavel, A. & Tapley, B. (2015) Notes on breeding and behaviour in the Anamalai Dot Frog Ramanella anamalaiensis Rao, 1937. Herpetology Notes, 8, 221–225.

Huelsenbeck, J.P., Ronquist, F., Neilsen, R. & Bollback, J.P. (2001) Bayesian inference of phylogeny and its impact on evolutionary biology. Science, 294 (5550), 2310–2314.

Inger, R.F., Shaffer, H.B., Koshy, M. & Bakde, R. (1984) A report on a collection of amphibians and reptiles from the Ponmudi, Kerala, South India. Journal of the Bombay Natural History Society, 81 (2), 406–427.

Janzen, P. & Bopage, M. (2011) The herpetofauna of a small and unprotected patch of tropical rainforest in Morningside, Sri Lanka. Amphibian and Reptile Conservation, 5 (2), 1–13.

Jerdon, T.C. (1854 “1853”) Catalogue of reptiles inhabiting the Peninsula of India. Journal of the Asiatic Society of Bengal, 22, 522–534.

Jerdon, T.C. (1870) Notes on Indian herpetology. Proceedings of the Asiatic Society of Bengal, 1870, 66–85.

Kadadevaru, G.G., Kanamadi, R.D. & Schneider, H. (1998) Mating call of the burrowing frog, Ramanella montana (Jerdon 1859). Journal of Advanced Zoology, 19 (2), 91–93.

Kanamadi, R.D., Hiremath, C.R. & Schneider, H. (1993) The advertisement call of the south Indian frog Ramanella variegata (Microhylidae). Journal of herpetology, 27 (2), 218–219.

Karunarathna, S.D.M.S. & Amarasinghe, A.T. (2009) Mutualism in Ramanella nagaoi Manamendra-Arachchi & Pethiyagoda, 2001 (Amphibia: Microhylidae) and Poecilotheria species (Arachnida: Thereposidae) from Sri Lanka. Taprobanica, 1, 16–18.

Kirtisinghe P. (1957) The Amphibia of Ceylon. Published by the author, Colombo, 112 pp.

Kirtisinghe, P. (1958) Some hitherto undescribed anuran tadpoles. Ceylon Journal of Science, 1 (2), 171–176.

Krishna, S.N. & Bosch, J. (2007) The breeding behaviour and advertisement calls of the tree-hole breeding frog Ramanella montana (Microhylidae) in the Western Ghats, S. India. Acta Zoologica Sinica, 53 (3), 575–578.

Krishna, S.N., Krishna, S.B. & Vijayalaxmi, K.K. (2004) Breeding ecology of a rare microhylid, Ramanella montana, in the forests of Western Ghats, India. Current Science, 87 (1), 80–82.

Kurabayashi, A., Matsui, M., Belabut, D.M., Yong, H.S., Ahmad, N., Sudin, A., Mitsuru, K., Amir, H. & Sumida, M. (2011) From Antarctica or Asia? New colonization scenario for Australian-New Guinean narrow mouth toads suggested from the findings on a mysterious genus Gastrophrynoides. BMC Evolutionary Biology, 11 (1), 175.

Kuramoto, M. & Dubois, A. (2009) Bioacoustic studies on three frog species from the Western Ghats, South India. Current Herpetology, 28 (2), 65–70.

Manamendra-Arachchi, K. & Pethiyagoda, R. (2001) Ramanella nagaoi, a new tree-hole frog (Microhylidae) from southern Sri Lanka. Journal of South Asian Natural History, 5 (2), 121–133.

Matsui, M., Hamidy, A., Belabut, D.M., Ahmad, N., Panha, S., Sudin, A., Khonsue, W., Oh, H.S., Yong, H.S., Jiang, J.P. & Nishikawa, K. (2011) Systematic relationships of Oriental tiny frogs of the family Microhylidae (Amphibia, Anura) as revealed by mtDNA genealogy. Molecular phylogenetics and evolution, 61 (1), 167–176.

Meegaskumbura, M. (1999) The life history of the red Ramanella (Ramanella obscura), a frog endemic to Sri Lanka. Loris, 22 (1), 22–28.

Meegaskumbura, M. (2001) The ecology of Ramanella obscura (Anura: Microhylidae) in a home-garden habitat and a mid country forest (Gannoruwa). Lyriocephalus, 4, 36–39.

Nelson, C.E. (1973) Mating calls of the Microhylinae: descriptions and phylogenetic and ecological considerations. Herpetologica, 29 (2), 163–176.

Parker, H.W. (1934) A Monograph of the Frogs of the Family Microhylidae. Trustees of the British Museum, London, 208 pp.

Peloso, P.L.V., Frost, D.R., Richards, S.J., Rodrigues, M.T., Donnellan, S.C., Matsui, M., Raxworthy, C.J., Biju, S.D., Lemmon, E.M., Lemmon, A.R. & Wheeler, W.C. (2016). The impact of anchored phylogenomics and taxon sampling on phylogenetic inference in Narrow-mouthed Frogs (Anura, Microhylidae). Cladistics, 32 (2), 113–140.

Peters, W.C.H. (1864 “1863”) Fernere mittheilungen über neue batrachier. Monatsberichte der Königlichen Preussische Akademie des Wissenschaften zu Berlin, 1863, 445–470.

Posada, D. & Crandall, K.A. (1998) Modeltest: testing the model of DNA substitution. Bioinformatics, 14 (9), 817–818.

Pyron, R.A. & Wiens, J.J. (2011) A large-scale phylogeny of Amphibia including over 2800 species, and a revised classification of advanced frogs, salamanders, and caecilians. Molecular Phylogenetics and Evolution, 61 (2), 543–583.

Ramaswami, L.S. (1936) The cranial morphology of the genera Kaloula Gray and Phrynella Boulenger (Anura). Proceedings of the Zoological Society of London, 106 (4), 1137–1155.

Rao, C.R.N. (1918) Notes on tadpoles of Indian Engystomatidae. Records of Indian Museum, 15, 41–45.

Rao, C.R.N. (1937) On some new forms of Batrachia from S. India. Proceedings of the Indian Academy of Sciences, Section B, 6, 387–427.

Rao, C.R. & Ramanna, B.S. (1925) On a new Genus of the Family Engystomatidæ (Batrachia). Journal of Zoology, 95 (2), 587–597

Ronquist, F. & Huelsenbeck, J.P. (2003) MrBayes 3: Bayesian phylogenetic inference under mixed models. Bioinformatics, 19 (12), 1572–1574.

Schneider, J.G. (1799) Historia amphibiorum naturalis et literarariae. Fasciculus primus. Continens Ranas, Calamitas, Bufones, Salamandras et Hydros in genera et species Descriptos notisque suis distinctos. Friederici Frommanni, Jena, 266 pp.

Sclater, W.L. (1892) List of the Batrachia in the Indian Museum. Taylor and Francis, London, 43 pp.

Siliwal, M. & Ravichandran, B. (2008) Commensalism in microhylid frogs and mygalomorph spiders. Zoos’ Print, 23 (8), 13.

Silvestro, D. & Michalak, I. (2012) raxmlGUI: a graphical front-end for RAxML. Organisms Diversity & Evolution, 12 (4), 335–337.

Simon, C., Frati, F., Beckenbach, A., Crespi, B., Liu, H. & Flook, P. (1994) Evolution, weighting and phylogenetic utility of mitochondrial gene sequences and a compilation of conserved polymerase chain reaction primers. Annals of the Entomological Society of America, 87 (6), 651–701.

Somaweera, R. (2006) Defensive posture in Kaloula taprobanica Parker (Microhylidae: Amphibia). Journal of Bombay Natural History Society, 103 (1), 108.

Stamatakis, A., Hoover, P. & Rougemont, J. (2008) A rapid bootstrap algorithm for the RAxML Web Servers. Systematic biology, 57 (5), 758–771.

Stoliczka, F. (1872) Observations on Indian Batrachia. Proceedings of the Asiatic Society of Bengal, 1872, 101–113.

Subramanian, K.A., Dinesh, K.P. & Radhakrishnan, C. (2013) Atlas of endemic amphibians of Western Ghats. Zoological Survey of India, Kolkata, 246 pp.

Swofford, D.L. (2002) PAUP*: Phylogenetic Analysis Using Parsimony (* and other methods). Version 4.0b10. Sinauer Association Inc., Sunderland, Massachusetts.

Tamura, K., Stecher, G., Peterson, D., Filipski, A. & Kumar, S. (2013) MEGA6: Molecular Evolutionary Genetics Analysis Version 6.0. Molecular biology and evolution, 30 (12), 2725–2729.

Taylor, W.R. & Van Dyke, G.C. (1985) Revised procedures for staining and clearing small fishes and other vertebrates for bone and cartilage study. Cybium, 9 (2), 107–119.

Tiedemann, F., Häupl, M. & Grillitsch, H. (1994) Katalog der Typen der Herpetologischen Sammlung nach dem Stand vom 1. Jänner 1994. Teil I: Amphibia. Kataloge Band 9 der wissenschaftlichen Sammlungen des Naturhistorischen Museums in Wien. Vertebrata. Heft 3. Naturhistorisches Museum, Wien, 42 pp.

Trueb, L. (1993) Patterns of cranial diversity among the Lissamphibia. In: Hanken, J. and Hall, B.K. (Eds), The skull. Vol. 2. Patterns of structural and systematic diversity. The University of Chicogo Press, Chicago, pp. 255–343.

Ukuwela, K.D.B., Silva, E.I.L. & Athukorala, N.P. (2010) The tadpole of Ramanella palmata (Anura: Microhylidae), a frog endemic to Sri Lanka. Zootaxa, 2628, 63–65.

Van Bocxlaer, I., Roelants, K., Biju, S.D., Nagaraju, J. & Bossuyt, F. (2006) Late Cretaceous vicariance in Gondwanan amphibians. PloS ONE, 1 (1), e74.

Vijaykumar, S.P., Zachariah, A. & Biju, S.D. (2011) Lost Amphibians of India. Available from: new2_rediscovered.php (accessed 24 January 2017)

Wells, K.D. (2007) The Ecology and Behavior of Amphibians. University of Chicago Press, Chicago, 1400 pp.

Wijayathilaka, N., Garg, S., Senevirathne, G., Karunarathna, N., Biju, S.D. & Meegaskumbura, M. (2016) A new species of Microhyla (Anura: Microhylidae) from Sri Lanka: an integrative taxonomic approach. Zootaxa, 4066 (3), 331–342.



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