DOI: 10.3724/SP.J.1245.2011.00001

Asian Herpetological Research 2011/2:1 PP.1-11

The Systematic Status of Rhacophorid Frogs

The frog family Rhacophoridae recently underwent notable reorganization and taxonomic rearrangement. Several new genera were erected to reflect evolutionary history, including Ghatixalus, Gracixalus, Feihyla, Liuixalus and Raorchestes. Herein, we reviewed the systematics and phylogeny of the rhacophorids, the definition of the family Rhacophoridae and its tribes, and the taxonomic history and diagnoses of the genera of this family. Also, we suggest future directions for research.

Key words:progress, prospect, faunal survey, systematics, amphibian, China

ReleaseDate:2014-07-21 15:50:43

Ahl E. 1931. Amphibia Anura III. Polypedatidae. Das Tierreich, Berlin: Lieferung, 55, xvi, 1-477

Amphibia Web. 2010. Information on amphibian biology and conservation [web application]. Available at http://amphibiaweb. org (accessed 2010)

Bahir M. M., Meegaskumbura M., Manamendra-Arachchi K., Schneider C. J., Pethiyagoda R. 2005. Reproduction and terrestrial direct development in Sri Lankan shrub frogs (Ranidae: Rhacophorinae: Philautus). Raffles Bull Zool (Suppl.), 12: 339-350

Bain R. H., Nguyen Q. T. 2004. Herpetofaunal diversity of Ha Giang Province in northeastern Vietnam, with descriptions of two new species. Am Mus Nov, 3453: 1-42

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. Zool J Linn Soc, 155: 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. Org Diver Evol, 8: 267-276

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

Bordoloi S., Bortamuli T., Ohler A. 2007. Systematics of the genus Rhacophorus (Amphibia, Anura): Identity of red-webbed forms and description of a new species from Assam. Zootaxa, 1653: 1-20

Bossuyt F., Brown R. M., Hillis 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. Syst Biol, 55: 579-594

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. Proc Natl Acad Sci, 97: 6585-6590

Bossuyt F., Milinkovitch M. C. 2001. Amphibians as indicators of early Tertiary ’Out-of-India’ dispersal of vertebrates. Science, 292: 93-95

Bossuyt F., Roelants K. 2009. Anura. In S. B. Hedges and S. Kumar (Eds.). Timetree of Life, New York: Oxford University Press, 357-364

Boulenger G. A. 1903. On a new frog from upper Burma and Siam. Ann Mag Nat Hist, 12: 219

Brown R. M., Linkem C. W., Siler C. D., Sukumaran J., Esselstyn J. A., Diesmos A. C., Iskandabr D. T., Bickford D., Evans B. J., McGuire J. A., Grismer L., Supriatna J., Andayani N. 2010. Phylogeography and historical demography of Polypedates leucomystax in the islands of Indonesia and the Philippines: Evidence for recent human-mediated range expansion? Mol Phylogenet Evol, 57: 598-619

Channing A. 1989. A re-evaluation of the phylogeny of Old World treefrogs. S Afr J Zool, 24: 116-131

Delorme M., Dubois A., Grosjean S., Ohler A. 2005. Une nouvelle classification générique et subgénérique de la tribu des Philautini (Amphibia, Anura, Ranidae, Rhacophorinae). Bull Mens Soc Linn Lyon, 74: 165-171

Dring J. C. M. 1979. Amphibians and reptiles from northern Trengganu, Malaysia, with descriptions of two new geckos: Cnemaspis and Cyrtodactylus. Bull Br Mus, 34: 181-241

Dring J. C. M. 1983. Some new frogs from Sarawak. Amphib-Rept, 4: 103-115

Dring J. C. M. 1987. Bornean treefrogs of the genus Philautus (Rhacophoridae). Amphib-Rept, 8: 19-47

Dubois A. 1981. Liste des genres et sous-genres nominaux de Ranoidea (Amphibiens Anoures) du monde, avec identification de leurs especes-types: consequences nomenclaturales. Monit Zool Ital (N. S.) (Suppl), 15: 225-284

Dubois A. 1987. Miscellanea taxinomica batrachologica (I). Alytes, 5: 7-95

Dubois A. 1992. Notes sur la classification des Ranidae (Amphibiens, Anoures). Bull Mens Soc Linn Lyon, 61: 305-352

Dubois A. 2005. Amphibia Mundi.1.1. An ergotaxonomy of recent amphibians. Alytes, 23: 1-24 Duellman W., Trueb L. 1986. Biology of the Amphibians. Baltimore, MD: Johns Hopkins Press

Dutta S. K. 1997. Amphibians of India and Sri Lanka. Bhubaneswar, India: Odyssey Publishing House

Emerson S. B., Richards C., Drewes R. C., Kjer K. M. 2000. On the relationships among ranoid frogs: A review of the evidence. Herpetologica, 56: 209-230

Fei L. 1999. Atlas of Amphibians of China. Zhengzhou, China: Henan Science and Technology Press (In Chinese)

Fei L., Ye C. Y., Jiang J. P., Xie F., Huang Y. Z. 2005. An Illustrated Key to Chinese Amphibians. Chengdu, China: Sichuan Publishing House of Science and Technology (In Chinese)

Fei L., Hu S. Q., Ye C. Y., Huang Y. Z. 2009. Fauna Sinica, Amphibia, Anura. Vol. 2. Beijing, China: Science Press, 473 pp (In Chinese)

Fei L., Ye C. Y., Jiang J. P. 2010a. Colored Atlas of Chinese Amphibians, Chengdu, China: Sichuan Publishing House of Science and Technology (In Chinese)

Fei, L., Ye C. Y., Jiang J. P. 2010b. A new species of Rhacophoridae from Yunnan, China (Amphibia, Anura). Acta Zootax Sin, 35 (2): 413-417 (In Chinese)

Frost D. R. 2010. Amphibian Species of the World: An Online Reference. Version 5.4. (8 April, 2010) Electronic database accessible at New York, USA: American Museum of Natural History

Frost D. R., Grant T., Faivovich J., Bain R. H., Haas A., Haddad C. F. B., De Sá R. O., Channing A., Wilkinson M., Donnellan S. C., Raxworthy C. J., Campbell J. A., Blotto B. L., Moler P., Drewes R. C., Nussbaum R. A., Lynch J. D., Green D. M., Wheeler W. C. 2006. The amphibian tree of life. Bull Am Mus Nat Hist, 297: 1-370

Grosjean S., Delorme M., Dubois A., Ohler A. 2008. Evolution of reproduction in the Rhacophoridae (Amphibia, Anura). J Zool Syst, 46: 169-176

Gururaja K. V., Aravind N. A., Ali S., Ramachandra T. V., Velavan T. P., Krishnakumar V., Aggarwa R. K. 2007. A new frog species from the Central Western Ghats of India, and its phylogenetic position. Zool Sci, 24: 525-534

Harvey M. B., Pemberton A. J., Smith E. N. 2002. New and poorly known parachuting frogs (Rhacophoridae: Rhacophorus) from Sumatra and Java. Herpetol Monogr, 16: 46-92

Inger R. F. 1966. The systematics and zoogeography of the Amphibia of Borneo. Fieldiana: Zoology, 52: 1-402

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

Iskandar D. T., Colijn E. 2000. Preliminary checklist of Southeast Asian and New Guinean herpetofauna. I. Amphibians. Treubia, 31 (Suppl to part 3): 1-134

Jiang S. P., Hu S. Q., Zhao E. M. 1987. The approach of the phylogenetic relationships and the supraspecific classification of 14 Chinese species of treefrogs (Rhacophoridae). Acta Herpetol Sin, 6: 43-47 (In Chinese)

Kuraishi N., Matsui M., Ota H. 2009. Estimation of the origin of Polypedates leucomystax (Amphibia: Anura: Rhacophoridae) introduced to the Ryukyu Archipelago, Japan. Pac Sci, 63: 317-325

Laurent R. F. 1943. Contribution à l’ostéologie et la systématique des Rhacophoridés non-africains. Bull Mus R Hist Nat Belg, 19(28): 1-16

Laurent R. F. 1946. Mise au point de la taxonomie des Ranidés. Rev Zool Bot Afr, 39: 336-338

Laurent R. F. 1951. Sur la necessite de supprimer la famille des Rhacophoridae mais de creer celle des Hyperoliidae. Rev Zool Bot Afr, 45: 116-122

Li J. T., Che J., Bain R. H., Zhao E. M., Zhang Y. P. 2008. Molecular phylogeny of Rhacophoridae (Anura): A framework of 10 Asian Herpetological Research Vol. 2 taxonomic reassignment of species within the genera Aquixalus, Chiromantis, Rhacophorus, and Philautus. Mol Phylogenet Evol, 48: 302-312

Li J. T., Che J., Murphy R. W., Zhao H., Zhao E. M., Rao D. Q., Zhang Y. P. 2009. Evolution of reproduction in the Rhacophoridae (Amphibia: Anura) inferred from a phylogenetic analysis of five nuclear and three mitochondrial genes. Mol Phylogenet Evol, 53: 509-522

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

Manzano A. S., Fabrezi M., Vences M. 2007. Intercalary elements, treefrogs, and the early differentiation of a complex system in the Neobatrachia. Anat Rec, 290: 1551-1567

Marmayou J., Dubois A., Ohler A., Pasquet E., Tillier A. 2000. Phylogenetic relationships in the Ranidae. Independent origin of direct development in the genera Philautus and Taylorana. CR Acad Sci Paris, Life Sci, 323: 287-297

Matsui M., Wu G. F. 1994. Acoustic characteristics of treefrogs from Sichuan, China, with comments on systematic relationship of Polypedates and Rhacophorus (Anura, Rhacophoridae). Zool Sci, 11: 485-490

Manamendra-Arachchi K., Pethiyagoda R. 2005. The Sri Lankan shrub-frogs of the genus Philautus Gistel, 1848 (Ranidae: Rhacophorinae), with description of 27 new species. Raff Bull Zool (Suppl), 12: 163-303

Meegaskumbura M., Bossuyt F., Pethiyagoda R., Manamendra-Arachchi K., Bahir M., Milinkovitch M. C., Schneider C. J. 2002. Sri Lanka: An amphibian hot spot. Science, 298: 379

Meegaskumbura M., Manamendra-Arachchi K. 2005. Description of eight new species of shrub-frogs (Ranidae: Rhacophorinae: Philautus) from Sri Lanka. Raffl B Zoo (Suppl), 12: 305-338

Narins P. M., Feng A. S., Yong H. S., Christensen-Dalsgaard J. 1998. Morphological, behavioral, and genetic divergence of sympatric morphotypes of the treefrog Polypedates leucomystax in penninsular Malaysia. Herpetologica, 254: 129-142

Nishioka M., Sumida M., Ohta S., Suzuki H. 1987. Speciation of three allied genera, Buergeria, Rhacophorus, and Polypedates, elucidated by the method of electrophoretic analyses. Scientific Report of the Laboratory for Amphibian Biology. Hiroshima Univ, 9: 53-96

Orlov N. L., Cuc H. T., Truong N. Q. 2004. A new species of the genus Philautus from Central Vietnam (Anura: Rhacophoridae). Russ J Herp, 11: 51-64

Orlov N. L., Murphy R. W., Ananjeva N. B., Ryabov S. A., Cuc H. T. 2002. Herpetofauna of Vietnam, a checklist. Part I. Amphibia. Russ J Herpetol, 9: 81-104

Rao D. Q., Wilkinson J. A., Liu H. N. 2006. A new species of Rhacophorus (Anura: Rhacophoridae) from Guangxi Province, China. Zootaxa, 1258: 17-31

Richards C. M., Nussbaum R. A., Raxworthy C. J. 2000. Phylogenetic relationships within the Madagascan boophids and mantillids as elucidated by mitochondrial ribosomal genes. Afr J Herpetol, 49: 23-32

Richards C. M., Moore W. S. 1998. A molecular phylogenetic study of the Old World treefrog family Rhacophoridae. Herpetol J, 8: 41-46

Roelants K., Jiang J., Bossuyt F. 2004. Endemic ranid (Amphibia: Anura) genera in southern mountain ranges of the Indian subcontinent represent ancient frog lineages: Evidence from molecular data. Mol Phylogenet Evol, 31: 730-740

Smith M. A. 1931. The herpetology of Mt. Kinabalu, North Borneo, 13,455 ft. Bull Raffles Mus, 5: 8-32, pl.1-2

Trépanier T. L., Lathrop A., Murphy R. W. 1999. Rhacophorus leucomystax in Vietnam with an acoustic analysis of advertisement and release calls. Asiatic Herpetol Res, 8: 102-106

Van der Meijden A., Vences M., Hoegg S., Meyer A. 2005. A previously unrecognized radiation of ranid frogs in southern Africa revealed by nuclear and mitochondrial DNA sequences. Mol Phylogenet Evol, 37: 674-685

Vences M., Glaw F. 2001. When molecules claim for taxonomic changes: New proposals on the classification of Old World treefrogs (Amphibia, Anura, Ranoidea). Spixiana, 24: 85-92

Vitt L. J., Caldwell J. P. 2009. Herpetology: An Introductory Biology of Amphibians and Reptiles, 3rd Edition. Burlington, Massachusetts: Academic Press

Wake D. B. 1966. Comparative osteology and evolution of the lungless salamanders, family Plethodontidae. Mem S Calif Acad Sci, 4: 1-111

Wake D. B. 1987. Adaptive radiation of salamanders in Middle American cloud forests. Ann Missouri Bot Gard, 74: 242-264

Wiens J. J., Sukumaran J., Pyron R. A., Brown R. M. 2009. Evolutionary and biogeographic origins of high tropical diversity in Old World frogs (Ranidae). Evolution, 63: 1217-1231

Wilkinson J. A. 1997. Historical biogeography of rhacophorid frogs from Japan, the Ryukyu Archipelago, and Taiwan inferred from a phylogenetic perspective. Ph. D. Dissertation, Texas Tech University, Lubbock, Texas.

Wilkinson J. A., Drewes R. C. 2000. Character assessment, genus level boundaries, and phylogenetic analyses of the family Rhacophoridae: A review and present day status. Cont Herpetol, 2: 1-24

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. Mol Phylogenet Evol, 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). Proc Calif Acad Sci, 54: 17-26

Yang Y. J., Lin Y. S., Wu J. L., Hui C. F. 1994. Variation in mitochondrial DNA and population structure of the Taipei treefrog Rhacophorus taipeianus in Taiwan. Mol Eco, 3: 219-228

Yang Y. J., Lin Y. S. 1997. Molecular phylogenetics and biogeography of the genus Rhacophorus in Taiwan. American Society of Ichthyologists and Herpetologists, 77th Annual Meeting, Seattle Washington

Yu G. H., Rao D. Q., Yang J. X., Zhang M. W. 2007. Non-monophyly of Rhacophorus rhodopus, Theloderma, and Philautus albopunctatus inferred from mitochondrial 16S rRNA gene sequences. Zool Res, 28: 437-442 (In Chinese)

Yu G. H., Rao D. Q., Yang J. X., Zhang M. W. 2008. Phylogenetic relationships among Rhacophorinae (Rhacophoridae, Anura, Amphibia), with an emphasis on the Chinese species. Zool J Linn, 53: 733-749

Yu G. H., Rao D. Q., Zhang M. W., Yang J. X. 2009. Reexamination of the phylogeny of Rhacophoridae (Anura) based on mitochondrial and nuclear DNA. Mol Phylogenet Evol, 50: 571-579

Yu G. H., Zhang M. W., Yang J. X. 2010. A species boundary within the Chinese Kurixalus odontotarsus species group (Anura: Rhacophoridae): New insights from molecular evidence. Mol Phylogenet Evol, 56(3): 942-50

Zhang P., Chen Y. Q., Zhou H., Liu Y. F., Wang X. L., Papenfuss T. J., Wake D. B., Qu L. H. 2006. Phylogeny, evolution, and biogeography of Asiatic Salamanders (Hynobiidae). Proc National Acad Sci, 103: 7360-7365

Zhao E. M., Adler K. 1993. Herpetology of China. Oxford Ohio: Society for the Study of Amphibians and Reptiles, in Cooperation with Chinese Society for the Study of Amphibians and Reptiles

Zhao E. M., Chang H. W., Zhao H., Adler K. 2000. Revised checklist of Chinese Amphibia and Reptilia. Sich J Zool, 19: 196-207 (In Chinese)