Skip to main content Skip to main navigation menu Skip to site footer
Type: Article
Published: 2020-11-12
Page range: 1–55
Abstract views: 658
PDF downloaded: 11

New insights on the systematics and reproductive behaviour in tree frogs of the genus Feihyla, with description of a new related genus from Asia (Anura, Rhacophoridae)

Systematics Lab, Department of Environmental Studies, University of Delhi, Delhi, India
Systematics Lab, Department of Environmental Studies, University of Delhi, Delhi, India
Andaman and Nicobar Regional Centre, Zoological Survey of India, Port Blair, Andaman and Nicobar Islands, India
Andaman and Nicobar Regional Centre, Zoological Survey of India, Port Blair, Andaman and Nicobar Islands, India
Department of Biology, Faculty of Science, Chulalongkorn University, Phyathai Road, Pathumwan, Bangkok, Thailand.
Chengdu Institute of Biology, Chinese Academy of Sciences, Chengdu, Sichuan, China.
Systematics Lab, Department of Environmental Studies, University of Delhi, Delhi, India
Systematics Lab, Department of Environmental Studies, University of Delhi, Delhi, India
Museum Zoologicum Bogoriense, Research Center for Biology, Indonesian Institute of Sciences, Cibinong, West Java, Indonesia.
National Centre for Cell Science, Savitribai Phule University of Pune Campus, Ganeshkhind, Pune, Maharashtra, India.
Chiromantis Chirixalus egg attendance gel-nesting generic reallocation integrative taxonomy larval morphology molecular phylogeny Rohanixalus gen. nov. species groups


The taxonomy of the Asian tree frog genus Feihyla has been in a state of flux ever since its proposal in 2006. Allocation of species to Feihyla remains confusing, particularly with respect to the closely related genus Chirixalus (formerly Chiromantis sensu lato). At the same time, several Chirixalus species are known only from cursory descriptions and remain poorly studied. In this study, we review the systematics of the genus Feihyla and clarify the generic placement of its members along with all other species currently assigned to Chirixalus. Based on integrative evidence gathered from new collections, examination of types and original descriptions, morphological comparisons, phylogenetic relationships inferred from a multi-gene (three mitochondrial + two nuclear) 1,937 bp dataset, as well as reproductive modes including egg-laying, nesting behaviour, and clutch morphology, our results show that the six species formerly attributed to Feihyla represent three morphologically and phylogenetically distinct groups: (1) the type species of the genus, F. palpebralis, along with F. fuhua constitute the Feihyla palpebralis group or Feihyla sensu stricto; (2) F. inexpectata and F. kajau represent the Feihyla vittiger group and are closely related to the Feihyla palpebralis group; (3) ‘F. hansenae’ and ‘F. vittata’ constitute another distinct and reciprocally monophyletic lineage, more closely related to Chirixalus + Chiromantis. In light of long-standing taxonomic confusions and unresolved phylogenetic relationships, we propose recognition of a new genus to accommodate ‘F. hansenae’ and ‘F. vittata’ in order to stabilise the classification of several Asian rhacophorid species that have been frequently confused and transferred within the Chirixalus–Chiromantis–Feihyla complex. Based on integrative evidence, eight species previously attributed to the genus Chirixalus or Feihyla are formally transferred to Rohanixalus gen. nov., two Chirixalus members are allocated to Feihyla, and one synonymised with Rhacophorus bipunctatus. In addition, we report the first member of the tree frog family Rhacophoridae from the Andaman Islands of India—Rohanixalus vittatus, along with description of its male advertisement call, reproductive behaviour including parental care by the female, and larval morphology. Extended distributions are also provided for Rohanixalus species across Northeast India. The study further reveals the presence of potentially undescribed diversity in the new genus. Altogether, the revised classification and novel insights presented herein will facilitate a better working taxonomy for four phylogenetically distinct but morphologically related groups of Old World tree frogs.


  1. Ahl, E. (1931) Amphibia, Anura III, Polypedatidae. Das Tierreich, 55, i–xvi + 1–477.

    Ao, M., Bordoloi, S. & Ohler, A. (2003) Amphibian fauna of Nagaland with nineteen new records from the State including five new records for India. Zoos’ Print Journal, 18 (6), 1117–1125.

    Aowphol, A., Rujirawan, A., Taksintum, W., Arsirapot, S. & McLeod, D.S. (2013) Re-evaluating the taxonomic status of Chiromantis in Thailand using multiple lines of evidence (Amphibia: Anura: Rhacophoridae). Zootaxa, 3702 (2), 101–123.

    Bain, R.H., Nguyen, Q.T. & Doan, V.K. (2007) New herpetofaunal records from Vietnam. Herpetological Review, 38, 107–117.

    Banerjee, A. (2014) Jelly secretion by a foam-nesting tree frog Chiromantis simus (Anura: Rhacophoridae): An unreported behaviour. Alytes, 31, 77–82.

    Bee, M.A., Suyesh, R. & Biju, S.D. (2013) The vocal repertoire of Pseudophilautus kani, a shrub frog (Anura: Rhacophoridae) from the Western Ghats of India. Bioacoustics, 22 (1), 67–85.

    Biju, S.D. & Bossuyt, F. (2003) New frog family from India reveals an ancient biogeographical link with the Seychelles. Nature, 425, 711–714.

    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. & Bossuyt, F. (2009) Systematics and phylogeny of Philautus. Gistel, 1848 (Anura, Rhacophoridae) in the Western Ghats of India, with descriptions of 12 new species. Zoological Journal of the Linnean Society, 155 (2), 374–444.

    Biju, S.D., Roelants, K. & Bossuyt, F. (2008) Phylogenetic position of the montane treefrog Polypedates variabilis Jerdon, 1853 (Anura: Rhacophoridae), and description of a related species. Organisms Diversity & Evolution, 8 (4), 267–276.

    Biju, S.D., Shouche, Y.S., Dubois, A., Dutta, S.K. & Bossuyt, F. (2010) A ground-dwelling rhacophorid frog from the highest mountain peak of the Western Ghats of India. Current Science, 98 (8), 1119–1125.

    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 (Bioligical 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.

    Biswas, S. (2000) Notes on the breeding habits of Chirixalus dudhwaensis Ray 1992, from Uttar Pradesh, north India. Hamadryad, 25 (2), 200–203.

    Bossuyt, F. & Dubois, A. (2001) A review of the frog genus Philautus Gistel, 1848 (Amphibia, Anura, Ranidae, Rhacophorinae). Zeylanica, 6, 1–112.

    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.

    Bossuyt, F., Brown, R.M., Hills, D.M., Cannatella, D.C. & Milinkovitch, M.C. (2006) Phylogeny and biogeography of a cosmopolitan frog radiation: Late Cretaceous diversification resulted in continent-scale endemism in the family Ranidae. Systematic Biology, 55 (4), 579–594.

    Bourret, R. (1939) Notes herpétologiques sur l’Indochine française. XX. Liste des Reptiles et batraciens actuellement connus en Indochine Française. Annexe au Bulletin Général de l’Instruction Publique, 1939, 49–60.

    Bourret, R. (1942) Les Batraciens de L’Indochine. Mémoires de l’Institute Océanographique de l’Indochine, 6, 1–547.

    Boulenger, G.A. (1887) On new batrachians from Malacca. Annals and Magazine of Natural History, 19, 345–348.

    Boulenger, G.A. (1893) Concluding report on the reptiles and batrachians obtained in Burma by Signor L. Fea dealing with the collection made in Pegu and the Karin Hills in 1887–88. Annali del Museo Civico di Storia Naturale di Genova, 13 (2), 304–347.

    Boulenger, G.A. (1897) Descriptions of new Malay frogs. Annals and Magazine of Natural History, Series 5, 19 (113), 106–108.

    Chan, K.O., Grismer, L.L., Anuar, S., Quah, E., Grismer, L.J., Wood, P.L. Jr., Muin, M.A. & Ahmad, N. (2011) A new species of Chiromantis Peters 1854 (Anura: Rhacophoridae) from Perlis State Park in extreme northern peninsular Malaysia with additional herpetofaunal records for the park. Russian Journal of Herpetology, 18, 253–259.

    Chan, K.O., Grismer, L.L. & Brown, R.M. (2018) Comprehensive multi-locus phylogeny of Old World tree frogs (Anura: Rhacophoridae) reveals taxonomic uncertainties and potential cases of over- and underestimation of species diversity. Molecular Phylogenetics and Evolution, 127, 1010–1019.

    Chanda, S.K. & Ghosh, A.K. (1989) A new frog of the genus Philautus Gistel, from the proposed Namdapha Biosphere Reserve, Arunachal Pradesh, northeast India. Journal of the Bombay Natural History Society, 86, 215–217.

    Chanda, S.K. &. Sarkar, A.K. (1997) The validity of Philautus shyamrupus Chanda and Ghosh, 1989 (Anura: Rhacophoridae). Hamadryad, 22 (1), 46–47.

    Charif, R.A., Waack, A.M. & Strickman, L.M. (2010) Raven Pro 1.4 User’s Manual. Cornell Laboratory of Ornithology, Ithaca, New York, 379 pp.

    Chen, J.M., Prendini, E., Wu, Y.H., Zhang, B.L., Suwannapoom, C., Chen, H.M., Jin, J.Q., Lemmon, E.M., Lemmong, A.R., Stuart, B.L., Raxworthy, C.J., Murphy, R.W., Yuan, Z.Y. & Jing, C. (2020) An integrative phylogenomic approach illuminates the evolutionary history of Old World tree frogs (Anura: Rhacophoridae). Molecular Phylogenetics and Evolution, 145, 106724. [Early view available online on 24 December 2019]

    Cochran, D.M. (1927) New reptiles and batrachians collected by Dr. Hugh M. Smith in Siam. Proceedings of the Biological Society of Washington, 40, 179–192.

    Das, I. (1999) Biogeography of the amphibians and reptiles of the Andaman and Nicobar Islands, India. In: Ota, H. (Ed.), Tropical Island herpetofauna. Origin, current diversity and current status. Elsevier, Amsterdam, pp. 43–77.

    Das, I., Hedeir, H., Pui, Y.M., Hertwig, S.T. & Haas, A. (2016) Larval external morphology and development in Feihyla kajau (Dring, 1983) (Amphibia: Anura: Rhacophoridae). Raffles Bulletin of Zoology, 64, 319–328.

    Delorme, M., Dubois, A., Grosjean, S. & Ohler, A. (2005) Une nouvelle classification générique et subgénérique de la tribu des Philautini (Amphibia, Anura, Ranidae, Rhacophorinae). Bulletin Mensuel de la Societe Linneenne de Lyon, 74 (5), 165–171.

    Deuti, K. & Dutta, S.K. (2002) First record of Boulenger’s tree frog Chirixalus vittatus (Anura: Rhacophoridae) from Mizoram, northeast India. Journal of the Bombay Natural History Society, 99, 126–127.

    Dring, J. (1983) Some new frogs from Sarawak. Amphibia-Reptilia, 4, 103–115.

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

    Dubois, A. (1999) South Asian Amphibia: a new frontier for taxonomists. Journal of South Asian Natural History, 4 (1), 1–11.

    Fei, L. (1999) Atlas of Amphibians of China. Henan Press of Science and Technology, Zhengzhou, 432 pp. [in Chinese]

    Fei, L., Hu, S.Q., Ye, C.Y. & Huang, Y.Z. (2009) Fauna Sinica. Amphibia Vol. 2 Anura. Science Press, Beijing, 957 pp.

    Fei, L., Ye, C.Y. & Jiang, J.P. (2010) A new species of Rhacophoridae from Yunnan, China (Amphibia, Anura). Acta Zootaxonomica Sinica, 35, 413–417.

    Fei, L., Ye, C.Y. & Jiang, J.P. (2012) Coloured Atlas of Chinese Amphibians and their Distributions. Sichuan Publishing House of Science & Technology, Sichuan, 620 pp.

    Frost, D.R., Grant, T., Faivovich, J., Bain, R.H., Haas, A., Haddad, C.F.B., de S, R.O., Channing, A., Wilkinson, M., Donnellan, S.C., Raxworthy, C.J., Campbell, J.A., Blotto, B.L., Moler, P.E., Drewes, R.C., Nussbaum, R.A., Lynch, J.D., Green, D.M. & Wheeler, W.C. (2006) The amphibian tree of life. Bulletin of the American Museum of Natural History, 297, 1–370.[0001:TATOL]2.0.CO;2

    Frost, D.R. (2020) Amphibian species of the world: an online reference. Version 6.0. American Museum of Natural History, New York. Available from: (accessed 25 April 2020)

    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., Senevirathne, G., Wijayathilaka, N., Phuge, S., Deuti, K., Manamendra-Arachchi, K., Meegaskumbura, M. & Biju, S.D. (2018) An integrative taxonomic review of the South Asian microhylid genus Uperodon. Zootaxa, 4384 (1), 1–88.

    Garg, S., Suyesh, R., Das, A., Jiang, J., Wijayathilaka, N., Amarasinghe, A.A.T., Alhadi, F., Vineeth, K.K., Aravind, N.A., Senevirathne, G., Meegaskumbura, M. & Biju, S.D. (2019) Systematic revision of Microhyla (Microhylidae) frogs of South Asia: a molecular, morphological, and acoustic assessment. Vertebrate Zoology, 69 (1), 1–71.

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

    Goutte, S., Dubois, A., Howard, S.D., Marquez, R., Rowley, J.J.L., Dehling, J.M., Grandcolas, P., Rongchuan, X. & Legendre, F. (2016) Environmental constraints and call evolution in torrent-dwelling frogs. Evolution, 70 (4), 811–826.

    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, 117–139.

    Grismer, L.L., Thy, N., Chav, T. & Holden, J. (2007) A new species of Chiromantis Peters 1854 (Anura: Rhacophoridae) from Phnom Samkos in the northwestern Cardamom Mountains, Cambodia. Herpetologica, 63 (3), 392–400.[392:ANSOCP]2.0.CO;2

    Grosjean, S. & Dubois, A. (2011) Description of the advertisement calls of nine species of Fejervarya Bolkay, 1915 and Minervarya Dubois, Ohler and Biju, 2001 from China, India and Nepal. Alytes, 27, 117–141.

    Grosjean, S., Ohler, A. & Chuaynkern, Y. (2015) Improving biodiversity assessment of anuran amphibians using DNA barcoding of tadpoles. Case studies from Southeast Asia. Comptes rendus biologies, 338 (5), 351–361.

    Grosjean, S. & Inthara, C. (2016) Molecular identifications and descriptions of the tadpoles of Rhacophorus kio Ohler & Delorme, 2006 and Rhacophorus rhodopus Liu & Hu, 1960 (Amphibia: Anura: Rhacophoridae). Zoosystema, 38 (2), 267–282.

    Harikrishnan, S., Chandramouli, S.R. & Vasudevan, K. (2012) A survey of herpetofauna on Long Island, Andaman and Nicobar Islands, India. Herpetological Bulletin, 119, 19–29.

    Harikrishnan, S. & Vasudevan, K. (2018) Amphibians of the Andaman & Nicobar Islands: distribution, natural history, and notes on taxonomy. Alytes, 36 (1–4), 238–265.

    Hasan, M.K., Mahony, S. & Kabir, V. (2010) Geographic distribution: Chiromantis doriae. Herpetological Review, 41, 103.

    Hertwig, S.T., Das, I., Schweizer, M., Brown, R. & Haas, A. (2012) Phylogenetic relationships of the Rhacophorus everetti-group and implications for the evolution of reproductive modes in Philautus (Amphibia: Anura: Rhacophoridae). Zoologica Scripta, 41 (1), 29–46.

    Hertwig, S.T., Schweizer, M., Das, I. & Haas, A. (2013) Diversification in a biodiversity hotspot—The evolution of Southeast Asian rhacophorid tree frogs on Borneo (Amphibia: Anura: Rhacophoridae). Molecular Phylogenetics and Evolution, 68, 567–581.

    Huelsenbeck, J.P. & Ronquist, F. (2001) MRBAYES: Bayesian inference of phylogenetic trees. Bioinformatics, 17, 754–755.

    Inger, R.F., Orlov, N.L. & Darevsky, I.S. (1999) Frogs of Vietnam: A report on new collections. Fieldiana Zoology, 92, 1–46.

    Kiyasetuo & Khare, M.K. (1986) A new record of frog, Chiromantis vittatus (Boulenger, 1887) from northeastern hills of India. Zoologica Orientalis, 3, 47–49.

    Kusrini, M.D., Lubis, M.I., Darmawan, B. & Rahman, L.N. (2017) Morphological and ecological observation on Chiromantis vittiger (Anura: Rhacophoridae) in Mount Halimun-Salak National Park, Indonesia. Treubia, 44, 47–66.

    Kuramoto, M., Joshy, S.H., Kurabayashi, A. & Sumida, M. (2008 [2007]) The genus Fejervarya (Anura: Ranidae) in central Western Ghats, India, with descriptions of four new cryptic species. Current Herpetology, 26, 81–105.[81:TGFARI]2.0.CO;2

    Li, J.T., Che, J., Bain, R.H., Zhao, E.M. & Zhang, Y.P. (2008) Molecular phylogeny of Rhacophoridae (Anura): a framework of taxonomic reassignment of species within the genera Aquixalus, Chiromantis, Rhacophorus and Philautus. Molecular Phylogenetics and Evolution, 48, 302–312.

    Li, J.T., Che, J., Murphy, R.W., Zhao, H., Zhao, E.M., Rao, D.Q. & Zhang, Y.P. (2009) New insights to the molecular phylogenetics and generic assessment in the Rhacophoridae (Amphibia: Anura) based on five nuclear and three mitochondrial genes, with comments on the evolution of reproduction. Molecular Phylogenetics and Evolution, 53, 509–522.

    Li, J.T., Li, Y., Klaus, S., Rao, D.Q., Hillis, D.M & Zhang, Y.P. (2013) Diversification of rhacophorid frogs provides evidence for accelerated faunal exchange between India and Eurasia during the Oligocene. Proceedings of the National Academy of Sciences of the United States of America, 110, 3441–3446.

    Li, P.-P., Zhao, E.-M. & Dong, B.-J. (2010) Amphibians and Reptiles of Tibet. Science Press, Beijing, XII + 251 pp.

    Liem, S.S. (1970) The morphology, systematics and evolution of the Old World treefrogs (Rhacophoridae and Hyperoliidae). Fieldiana Zoology, 57, 1–145.

    Lyu, Z.T., Dai, K.Y., Li, Y., Wan, H., Liu, Z.Y., Qi, S., Lin, S.M., Wang, J., Li, Y.L., Zeng, Y.J., Li, P.P., Pang, H. & Wang, Y.Y. (2020) Comprehensive approaches reveal three cryptic species of genus Nidirana (Anura, Ranidae) from China. ZooKeys, 914, 127–159.

    Mani, M.S. (1974) Ecology and Biogeography of India. Dr. W. Junk Publishers, The Hague, 527 pp.

    Mathew, R. & Sen, N. (2010 [2009]) Description of a new species of Leptobrachium Tschudi, 1838 (Amphibia: Anura: Megophryidae) from Meghalaya, India. Records of the Zoological Survey of India, 109, 91–108.

    Mathew, R. & Sen, N. (2009) Studies on little known amphibians of Northeast India. Records of the Zoological Survey of India, Occasional Papers, 293, 1–64.

    Matsui, M., Shimada, T. & Sudin, A. (2014) First record of the tree-frog genus Chiromantis from Borneo with the description of a new species (Amphibia: Rhacophoridae). Zoological Science, 31, 45–51.

    McDiarmid, R.W. & Altig, R. (1999) Tadpoles: The Biology of Anuran larvae. The University of Chicago Press, Chicago, xiv + 444 pp.

    Meegaskumbura, M., Bossuyt, F., Pethiyagoda, R., Manamendra-Ararchchi, K., Bahir, M., Milinkovitch, M.C. & Schneider, C.J. (2002) Sri Lanka: an amphibian hotspot. Science 298, 379.

    Meegaskumbura, M., Meegaskumbura, S., Bowatte, G., Manamendra-Arachchi, K., Pethiyagoda, R., Hanken, J. & Schneider, C.J. (2010) Taruga (Anura: Rhacophoridae), A new genus of foam-nesting tree frogs endemic to Sri Lanka. Ceylon Journal of Science (Biological Sciences), 39, 75–94.

    Meegaskumbura, M., Senevirathne, G., Biju, S.D., Garg, S., Meegaskumbura, S., Pethiyagoda, R., Hanken, J. & Schneider, C.J. (2015) Patterns of reproductive mode evolution in Old World tree frogs (Anura, Rhacophoridae). Zoologica Scripta, 44 (5), 509–522.

    Nguyen, V.S., Ho, T.C. & Nguyen, Q.T. (2005) Danh Luc Ech Nhai Va Bo Sat Viet Nam [A Checklist of the Amphibians and Reptiles of Vietnam]. Nha Xuat Ban Hong Nghiep, Hanoi. [in Vietnamese, unkown pagination]

    Poo, S. & Bickford, D.P. (2013) The adaptive significance of egg attendance in a South-East Asian tree frog. Ethology, 119, 671–679.

    Poo, S. & Low, M.R. (2018) Feihyla hansenae (Hansen’s Bush Frog). Multimale amplexus. Herpetological Review, 49 (3), 516–517.

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

    Poyarkov, N.A., Orlov, N.L., Moiseeva, A.V., Galoyan, E.A., Nguyen, T.T. & Gogoleva, S.S. (2015) Sorting out moss frogs: mtDNA data on taxonomic diversity and phylogenetic relationships of the Indochinese species of the genus Theloderma (Anura, Rhacophoridae). Russian Journal of Herpetology, 22, 241–280.

    Rangasamy, V., Sivaperuman, C., Gokulakrishnan, G. & Prathapan, P. (2018) Herpetofauna of Andaman and Nicobar Islands. In: Sivaperuman, C. & Venkatraraman, K. (Eds.), Indian Hotspots: Vertebrate Faunal Diversity, Conservation and Management. Vol. 2. Springer, Singapore, pp. 37–56.

    Richards, C.M. & Moore, W. (1996) A molecular phylogeny of the Old World tree frog family Rhacophoridae. Journal of Herpetology, 8, 41–46.

    Riyanto, A. & Kurniati, H. (2014) Three new species of Chiromantis Peters 1854 (Anura: Rhacophoridae). Russian Journal of Herpetology, 21, 65–73.

    Romer, J.D. (1951 “1949”) Herpetological observations in Assam and Bengal (1944). Journal Bombay Natural History Society, 48 (2), 374–376.

    Roonwal, M.L. & Kripalani, M.B. (1966 “1961”) A new frog, Philautus cherrapunjiae (family Ranidae) from Assam, India, with field observation on its behavior and metamorphosis. Records of the Indian Museum, 59, 325–333.

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

    Rowley, J.J.L., Tran, D.T.A., Frankham, G.J., Dekker, A.H., Le.D. T.T., Nguyen, T.Q., Dau, V.Q. & Hoang, H.D. (2015) Undiagnosed cryptic diversity in small, microendemic frogs (Leptolalax) from the Central Highlands of Vietnam. PLoS ONE, 10 (5), e0128382.

    Sarkar, A.K. (1990) Taxonomic and ecological studies on the amphibians of Andaman and Nicobar Islands, India. Records of the Zoological Survey of India, 86, 103–117.

    Sarkar, A.K. & Ray, S. (2006) Amphibia. In: Alfred, J.R.B. (Ed.), Fauna of Arunachal Pradesh. State Fauna Series. 13 (1). Zoological Survey of India, Kolkata, pp. 285–316.

    Sheridan, J.A. & Ocock, J.F. (2008) Parental care in Chiromantis hansenae (Anura: Rhacophoridae). Copeia, 2008, 733–736.

    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.

    Smith, M.A. (1924) New tree-frogs from Indo-China and the Malay Peninsula. Proceedings of the Zoological Society of London, 1924, 225–234.

    Sorokin, A. & Steigerwald, E.C. (2018) Evidence for parental care in Feihyla kajau (Anura: Rhacophoridae). Phyllomedusa, 17 (1), 127–130.

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

    Stuart, B.L. (2005) New frog records from Laos. Herpetological Review, 36, 473–479.

    Stuart, B.L. & Emmett, D.A. (2006) A collection of amphibians and reptiles from the Cardomom Mountains, southwestern Cambodia. Fieldiana Zoology, 109, 1–27.[1:TNSOFA]2.0.CO;2

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

    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, E.H. (1962) The amphibian fauna of Thailand. University of Kansas Science Bulletin, 43, 265–599.

    Thorell, T. (1887) Viaggio di L. Fea in Birmania e regioni vicine - II. Primo saggio sui Ragni Birmani. Annali del Museo Civico di Storia Naturale di Genova, Series 2, 5 (25), 5–417.

    Vassilieva, A.B., Galoyan, E.A., Poyarkov, N.A. & Geissler, P. (2016) A Photographic Field Guide to the Amphibians and Reptiles of the Lowland Monsoon Forests of Southern Vietnam. Frankfurt Contributions to Natural History. Vol. 36. Edition Chimaira, Frankfurt, 319 pp.

    Wan, J.P., Sung, Y.H., Chan, B.P.L. & Lu, G. (2012) Chiromantis vittatus (Two-striped Pigmy Tree Frog). Parental Care. Herpetological Review, 43, 630–631.

    Wang, Y.Y., Zhao, J., Yang, J.H., Zhou, Z.X., Chen, G.L. & Liu, Y. (2014) Morphology, molecular genetics, and bioacoustics support two new sympatric Xenophrys (Amphibia: Anura: Megophryidae) species in Southeast China. PLoS ONE, 9, e93075.

    Wilkinson, J.A., Drewes, R.C. & Tatum, O.L. (2002) A molecular phylogenetic analysis of the family Rhacophoridae with an emphasis on the Asian and African genera. Molecular Phylogenetics and Evolution, 24, 265–273.

    Wilkinson, J.A., Win, H., Thin, T., Lwin, K.S., Shein, A.K. & Tun, H. (2003) A new species of Chirixalus (Anura: Rhacophoridae) from western Myanmar (Burma). Proceedings of the California Academy of Sciences, 54 (2), 17–26.

    Wostl, E., Riyanto, A., Hamidy, A., Kurniawan, N., Smith, E.N. & Harvey, M.B. (2017) A taxonomic revision of the Philautus (Anura: Rhacophoridae) of Sumatra with the description of four new species. Herpetological Monographs, 31 (1), 98–141.

    Yu, G., Rao, D., Yang, J.X. & Zhang, M. (2008) Phylogenetic relationships among Rhacophorinae (Rhacophoridae, Anura, Amphibia), with an emphasis on the Chinese species. Zoological Journal of the Linnean Society, 153, 733–749.

    Yu, G., Rao, D., Yang, J. & Zang, M. (2009) Re-examination of the phylogeny of Rhacophoridae (Anura) based on mitochondrial and nuclear DNA. Molecular Phylogenetics and Evolution, 50, 571–579.

    Yodthong, S., Siler, C.D., Prasankok, P. & Aowphol, A. (2014) Phylogenetic patterns of the Southeast Asian Tree Frog Chiromantis hansenae in Thailand. Asian Herpetological Research, 5, 179–196.