Skip to main content Skip to main navigation menu Skip to site footer
Type: Articles
Published: 2011-12-09
Page range: 22–38
Abstract views: 80
PDF downloaded: 1

A new species of Gracixalus (Anura: Rhacophoridae) with a hyperextended vocal repertoire from Vietnam

Australian Museum, 6 College St, Sydney, NSW, 2010, Australia
Institute of Ecology and Biological Resources, 18 Hoang Quoc Viet, Hanoi, Vietnam
Vietnam National Museum of Nature, 18 Hoang Quoc Viet, Cau Giay, Hanoi, Vietnam
Biology Faculty, Vinh University, 182 Le Duan St, Vinh City, Vietnam
Institute of Ecology and Biological Resources, 18 Hoang Quoc Viet, Hanoi, Vietnam
Amphibia Anura Nghe An Province Rhacophoridae Southeast Asia Vietnam


We describe a new species of small rhacophorid frog from north central Vietnam. Gracixalus quangi sp. nov. is morphologically and genetically most similar to G. gracilipes and G. supercornutus, but is distinguished from these species and all other rhacophorids in Vietnam and adjoining countries by a combination of their small size (males Gracixalus quangi sp. nov. has a non-stereotypical, hyperextended vocal repertoire, with calls highly variable in structure, duration, amplitude and frequency (dominant frequency 4.1–4.7 kHz). Like G. gracilipes and G. supercornutus, G. quangi sp. nov. deposits egg clutches in clumps on leaves overhanging shallow forest pools or puddles. The new species is known from montane evergreen forest in Pu Hoat Proposed Nature Reserve in western Nghe An Province, between ~600–1,300 m elevation. Preliminary molecular (mtDNA) data recovered two strongly supported clades within frogs currently considered to belong to genus Gracixalus, with the new species nested within a monophyletic clade consisting of G. gracilipes, G. supercornutus, and G. quyeti.


  1. Ahl, E. (1927) Zur Systematik der asiatischen Arten der Froschgattung Rhacophorus. Sitzungsberichte der Gesellschaft Naturforschender Freunde zu Berlin, 1927, 35–47.

    Alberch, P. & Gale, E.A. (1985) A developmental analysis of evolutionary trend: Digital reduction in Amphibians. Evolution, 39, 8–23.

    Annandale, N. (1912) Zoological results of the Abor Expedition, 1911–1912. I. Amphibia. Records of the Indian Museum, 8, 7–36.

    Bain, R.H., Nguyen, Q.T. & Doan K.V. (2009) A new species of the genus Theloderma Tschudi, 1838 (Anura: Rhacophoridae) from northwestern Vietnam. Zootaxa, 2191, 58–68.

    Bain, R.H. & Nguyen, Q.T. (2004) Herpetofaunal diversity of Ha Giang Province in Northeastern Vietnam, with descriptions of two new species. American Museum Novitates, 3453, 1–42.

    Boettger, O. (1895) Neue Frösche und Schlangen von den Liukiu-Inseln. Zoologischer Anzeiger, 18, 266–270.

    Boulenger, G.A. (1886) First report on additions to the batrachian collection in the Natural-History Museum. Proceedings of the Zoological Society of London, 1886, 411–416.

    Boulenger, G.A. (1887) An account of the batrachians obtained in Burma by M.L. Fea of the Genoa Civic Museum. Annali del Museo Civico di Storia Naturale di Genova. Serie 2, 5, 418–424.

    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 Genov, Serie 2, 13, 304–347.

    Bourret, R. (1937) Notes herpétologiques sur lIndochine française. XIV. Les batraciens de la collection du Laboratoire des Sciences Naturelles de lUniversité. Descriptions de quinze especes ou variétés nouvelles. Annexe au Bulletin Général de l'Instruction Publique. Hanoi, 1937, 5–56.

    Brown, W.C. & Alcala, A.C. (1994) Philippine frogs of the family Rhacophoridae. Proceedings of the Californian Academy of Sciences, 48, 185–220.

    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.

    Cocroft, R.B. & Ryan, M.J. (1995) Patterns of advertisement call evolution in toads and chorus frogs. Animal Behaviour, 49, 283–303.

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

    Duellman, W.E. & Trueb, L. (1986) Biology of Amphibians. The John Hopkins University Press, Baltimore and London. 670 pp.

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

    Fei, L., Ye, C.-y. & Jiang, J.-p. (2010a) Colored Atlas of Chinese Amphibians. Sichuan Publishing Group/Sichuan Publishing House of Science and Technology. 519 pp.

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

    Fellowes, J.R. & Hau, C-h. (1997) A Faunal Survey of Nine Forest Reserves in Tropical South China, with a Review of Conservation Priorities in the Region. Kadoorie Farm and Botanic Garden, Hong Kong. 152 pp.

    Frost, D.R. (2011) Amphibian Species of the World: an online reference. Version 5.5 (31 January, 2011). American Museum of Natural History, New York, USA Available from (accessed 11 March 2011).

    Glaw, F. & Vences, M. (1997) Anuran eye colouration: definitions, variation, taxonomic implications and possible functions. In: Böhme, W., Bischoff, W. & Ziegler, T., (Eds). Herpetologia Bonnensis. SEH Proceedings, Bonn. pp. 125–138.

    Grismer, L.L., Neang, T. Thou, C, & Holden, J. (2007) A new species of Chiromantis Peters 1854 (Anura: Rhacophoridae) from Phnom Samkos in the northwestern Cardomom Mountains, Cambodia. Herpetologica, 63, 392–400.

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

    Hallowell, E. (1861 "1860") Report upon the Reptilia of the North Pacific Exploring Expedition, under command of Capt. John Rogers U.S.N. Proceedings of the Academy of Natural Sciences of Philadelphia, 12, 480–510.

    Hillis, D.M., & Bull, J.J. (1993) Phylogenetic relationships of the dwarf boas and a comparison of Bayesian and bootstrap measures of phylogenetic support. Systematic Biology, 42, 182–192.

    Hu, S.-q. (1978) In: Hu, S.-q., Fei, L. & Ye., C.-y. Three new amphibian species in China. Materials for Herpetological Research, 4, 20.

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

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

    IUCN. (2001) IUCN Red List Categories and Criteria: Version 3.1. IUCN Species Survival Commission. IUCN, Gland, Switzerland and Cambridge, UK. ii + 30 pp.

    Jiang, J.-p., Fei, L., & Huang, Y.-z. (2009) In: Fei, L., Hu, S.-Q.,Ye, C.-Y, &. Huang, Y.-Z. (2009) Fauna Sinica. Amphibia. Volume 2. Anura. Science Press, Beijing. pp. 708–711.

    Kou, Z.-t. (1990) A new species of genus Philautus (Amphibia: Rhacophoridae) from Yunnan, China. In: Zhao, E.-m. (Eds.), From Water onto Land. A volume Issued to Commemorate the Ninetieth Birthday of the Late Professor Liu Cheng-zhao. Chung-kuo lin yeh ch`u pan she [Forestry Press], Beijing. pp. 210–212.

    Kuramoto, M. (1986) Call structures of the rhacophorid frogs from Taiwan. Scientific report of the Laboratory for Amphibian Biology, Hiroshima University, 8, 45–68.

    Kuramoto, M., & Wang, C.-s. (1987) A new rhacophorid treefrog from Taiwan, with comparisons to Chirixalus eiffingeri (Anura, Rhacophoridae). Copeia, 1987, 931–942.

    Kuhl, H., & Van Hasselt, J.C. (1822) Uittreksels uit breieven van de Heeren Kuhl en van Hasselt, aan de Heeren C. J. Temminck, Th. van Swinderen en W. de Haan. Algemeene Konst-en Letter-Bode, 7, 99–104.

    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.

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

    Liu, C.-c. (1935) Types of vocal sac in the Salentia. Proceedings of the Boston Society of Natural History, 41, 19–40.

    Liu, C.-c., &. Hu, S.-q (1962) A herpetological report of Kwangsi. Acta Zoologica Sinica, 14 (Supplement), 73–104.

    Liu, C.-c. & Hu, S.-q. (1973) In: Liu, C.-c., Hu, S.-q., Fei, L., & Huang, C.-C. On collections of amphibians from Hainan Island. Acta Zoologica Sinica, 19, 393–394.

    Liu, S.-l., & Wu, G.-f. (2004) In: Liu, S.-l., Wang, L.-j., Lu, S.-q., Zhao, H., Che, J., & Wu, G.-f., A new species of the genus Philautus (Anura: Rhacophoridae) and winter herpetological exploration in Hainan Province, China. Sichuan Journal of Zoology, 123, 203–204.

    Matsui, M., & Orlov, N.L. (2004). A new species of Chirixalus from Vietnam (Anura: Rhacophoridae). Zoological Science, 21, 671–676.

    Myers, C.W. & Duellman, W.E. (1982) A new species of Hyla from Cerro Colorado, and other tree frog records and geographical notes from western Panama. American Museum Natural History Novitates, 2752, 1–32.

    Narins, P.M., Lewis E.R., & McClelland, B.E. (2000) Hyperextended call note repertoire of the endemic Madagascar treefrog Boophis madagascariensis (Rhacophoridae). Journal of Zoology, 250, 283–298.

    Nguyen, Q.T., Hendrix, R., Böhme, W., Vu, N.T. & Ziegler, T. (2008) A new species of the genus Philautus (Amphibia: Anura: Rhacophoridae) from the Truong Son Range, Quang Binh Province, central Vietnam. Zootaxa, 1925, 1–13.

    Ohler, A., Swan, S.R. & Daltry J.C. (2002) A recent survey of the amphibian fauna of the Cardamom Mountains, Southwest Cambodia with descriptions of three new species. Raffles Bulletin of Zoology, 50, 465–481.

    Orlov, N.L., & Ho T.C. (2004) A new species of the genus Philautus from central Vietnam (Anura: Rhacophoridae). Russian Journal of Herpetology, 11, 51–64.

    Orlov, N.L., & Ho, T.C. (2005) A new species of Philautus from Vietnam (Anura: Rhacophoridae). Russian Journal of Herpetology 12,135–142.

    Orlov N.L., Murphy, R.W., Ananjeva, N.B., Ryabov, S.A., & Ho, T.C., (2002) Herpetofauna of Vietnam. A checklist. Part 1. Amphibia. Russian Journal of Herpetology, 9, 81–104.

    Palumbi, S.R., Martin, A., Romano, S., McMillan, W.O., Stice, L. & Grabowski, G. (1991) The simple fool's guide to PCR. Department of Zoology, University of Hawaii, Honolulu. 47 pp.

    Peters, W.C.H. (1871) Über neue Reptilien aus Ostafrica und Sarawak (Borneo), vorzüglich aus der Sammlung des Hrn. Marquis J. Doria zu Genua. Monatsberichte der Königlichen Preussische Akademie des Wissenschaften zu Berlin 1871, 566–581.

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

    Posada, D. & Buckley, T.R. (2004) Model selection and model averaging in phylogenetics: advantages of the AIC and Bayesian approaches over likelihood ratio tests. Systematic Biology, 53, 793–808.

    Rambaut, A, & Drummond, A.J. (2007) Tracer v1.4, Available from

    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.

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

    Savage, J.M. & Heyer, W.R. (1967) Variation and distribution of the tree-frog genus Phyllomedusa in Costa Rica, Central America. Beiträge zur Neotropischen Fauna, 5, 111–131.

    Savage, J.M. & Heyer, W.R. (1997) Digital webbing formulae for anurans: a refinement. Herpetological Review, 28, 131.

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

    Smith, M.A. (1940) The amphibians and reptiles obtained by Mr. Ronald Kaulback in Upper Burma. Records of the Indian Museum, 42, 465–486.

    Smith, M.A. (1953) Description of a new species of frog of the genus Philautus. Annals and Magazine of Natural History, Series 12, 6, 477–478.

    Sukumaran, J. & Holder, M.T. (2010) DendroPy: a Python library for phylogenetic computing. Bioinformatics, 26, 1569–1571.

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

    Wilcox, T.P., Zwickl, D.J., Heath, T.A., & Hillis, D.M. (2002). Phylogenetic relationships of the dwarf boas and a comparison of Bayesian and bootstrap measures of phylogenetic support. Molecular Phylogenetics and Evolution, 25, 361–371.

    Yang, D.-t. & Li, S.-m. (1978) In: Yang, D.-t., C.-y. Su, & Li, S.-m. Amphibians and Reptiles of Gaoligongshan. Kunming, 8, 37–38.

    Ye, C.-y. & Fei, L. (1993) Philautus odontotarsus. In: C. Ye, L. Fei, and S. Hu (Eds), Rare and Economic Amphibians of China. Chengdu, China: Sichuan Publishing House of Science and Technology. pp.320–322.

    Ye, C.-y. & Hu, S.-q. (1984) A new species of Philautus (Anura: Rhacophoridae) from Xizang Autonomous Region. Acta Herpetologica Sinica, 3, 67–69.

    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. Zoological Journal of the Linnean Society, 153, 733–749.

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

    Yu, G.-h., M.-w. Zhang, & Yang, J.-x. (2010) A species boundary within the Chinese Kurixalus odontotarsus species group (Anura: Rhacophoridae): New insights from molecular evidence. Molecular Phylogenetics and Evolution, 56, 942–950.

    Zwickl, D.J. (2006) Genetic algorithm approaches for the phylogenetic analysis of large biological sequence datasets under the maximum likelihood criterion. Ph.D. dissertation, The University of Texas at Austin. 115 pp.