Abstract
A new species of the rhacophorid frog genus Theloderma is described from the forested shoreline of the Nam Lik Reservoir, Vientiane Province, Laos. The new species differs from its congeners by having the combination of males with SVL 17.0–20.6; pearly asperities on dorsum; no vomerine teeth; disc diameter of finger III ca. 40% of tympanum diameter; uniformly gray venter; light brown dorsum with darker brown and black markings; and a uniformly bronze iris with small black reticulations. Molecular phylogenetic analysis of mitochondrial DNA sequence data infers that the new species is most closely related to T. lateriticum from northern Vietnam. Evidence for the monophyly of Theloderma is reviewed.
References
Ahl, E. (1927) Zur Systematik der asiatischen Arten der Froschgattung Rhacophorus. Sitzungsberichte der Gesellschaft Naturforschender Freunde zu Berlin, 1927, 35–47.
Annandale, N. (1912) Zoological results of the Abor Expedition, 1911–1912. I. Amphibia. Records of the Indian Museum, 8, 7–36.
http://dx.doi.org/10.5962/bhl.part.1186Arinin, I.Y. & Ryabov, S.A. (2006) Captive breeding of Theloderma gordoni Taylor, 1962. Russian Journal of Herpetology, 13, 242–246.
Bain, R.H., Nguyen, T.Q. & Doan, K.V. (2009) A new species of the genus Theloderma Tschudi, 1838 (Anura: Rhacophoridae) from northwestern Vietnam. Zootaxa, 2191, 58–68.
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, e0145727.
http://dx.doi.org/10.1371/journal.pone.0145727Boulenger, G.A. (1882) Description of a new genus and species of frogs of the family Ranidae. Annals and Magazine of Natural History, Series 5, 10, 35.
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.
Darriba, D., Taboada, G.L., Doallo, R. & Posada, D. (2012) jModelTest 2: more models, new heuristics and parallel computing. Nature Methods, 9, 772.
http://dx.doi.org/10.1038/nmeth.2109Dever, J.A., Nguyen, H. & Wilkinson, J.A. (2016) Rediscovery and redescription of Theloderma phrynoderma (Ahl, 1927) (Anura: Rhacophoridae) from Myanmar. Copeia, 103, 402–415.
http://dx.doi.org/10.1643/CH-14-130Frost, D.R. (2016) Amphibian Species of the World: an Online Reference. Version 6.0 (21 April 2016). American Museum of Natural History, New York, USA. Available from: http://research.amnh.org/herpetology/amphibia/index.html (accessed 28 Jun. 2016)
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. Bulletin of the American Museum of Natural History, 297, 1–370.
http://dx.doi.org/10.1206/0003-0090(2006)297[0001:TATOL]2.0.CO;2Gawor, A., Chapuis, S., Pham, C.T., Nguyen, T.Q., Schmitz, A. & Ziegler, T. (2012) Larval morphology of two species of the genus Theloderma (Tschudi, 1838) from Vietnam (Anura: Rhacophoridae: Rhacophorinae). Zootaxa, 3395, 59–64.
Gistel, J. (1848) Naturgeschichte des Thierreichs. Für höhere Schulen. Hoffmann, Stuttgart, 216 pp.
Global Energy Observatory (2016) Available from: http://GlobalEnergyObservatory.org/ (accessed 21 Apr. 2016)
Grosjean, S., Delorme, M., Dubois, A. & Ohler, A. (2008) Evolution of reproduction in the Rhacophoridae (Amphibia, Anura). Journal of Zoological Systematics and Evolutionary Research, 46, 169–176.
http://dx.doi.org/10.1111/j.1439-0469.2007.00451.xHertwig, S.T., Das, I., Schweizer, M., Brown, R. & Haas, A. (2011) Phylogenetic relationships of the Rhacophorus everetti-group and implications for the evolution of reproductive modes in Philautus (Amphibia: Anura: Rhacophoridae). Zoologica Scripta, 41, 29–46.
http://dx.doi.org/10.1111/j.1463-6409.2011.00499.xHertwig, 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.
http://dx.doi.org/10.1016/j.ympev.2013.04.001Jiang, J.-P., Fei, L. & Huang, Y.-Z. (2009) Fauna Sinica. Amphibia. Vol. 2. Anura. Chinese Academy of Science, Science Press, Beijing, 957 pp. [pp. 708–711, In: Fei, L., Hu, S.-Q., Ye, C.-Y. & Huang, Y.-Z.]
Jiang, K., Yan, F., Wang, K., Zou, D.-H., Li, C. & Che, J. (2016) A new genus and species of treefrog from Medog, southeastern Tibet, China (Anura, Rhacophoridae). Zoological Research, 37, 15–20.
Katoh, K. & Standley, D.M. (2013) MAFFT Multiple Sequence Alignment software version 7: improvements in performance and stability. Molecular Biology and Evolution, 30, 772–780.
http://dx.doi.org/10.1093/molbev/mst010Li, C., Hou, M., Yu, G.-H., Yan, F., Li, B.-Z., Jiang, K., Li, P.-P. & Orlov, N.L. (2016) Rediscovery of Theloderma moloch (Amphibian: Anura: Rhacophoridae) from southeast Tibet, China after 102 years. Russian Journal of Herpetology, 23, 41–54.
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.
http://dx.doi.org/10.1016/j.ympev.2008.03.023Li, 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.
http://dx.doi.org/10.1016/j.ympev.2009.06.023Li, 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.
http://dx.doi.org/10.1073/pnas.1300881110Liu, C.-c. & Hu, S.-q. (1962) A herpetological report of Kwangsi. Acta Zoologica Sinica, 14 (Supplement), 73–104.
McLeod, D.S. & Ahmad, N. (2007) A new species of Theloderma (Anura: Rhacophoridae) from southern Thailand and peninsular Malaysia. Russian Journal of Herpetology, 14, 65–72.
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, 509–522.
http://dx.doi.org/10.1111/zsc.12121Nguyen, T.T., Le, D.T., Nguyen, S.H.L., Matsui, M. & Nguyen, T.Q. (2014) First record of Philautus petilus Stuart and Heatwole, 2004 (Amphibian: Anura: Rhacophoridae) from Vietnam and its phylogenetic position. Current Herpetology, 33, 112–120.
http://dx.doi.org/10.5358/hsj.33.112Nguyen, T.T., Matsui, M. & Eto, K. (2015) Mitochondrial phylogeny of an Asian tree frog genus Theloderma (Anura: Rhacophoridae). Molecular Phylogenetics and Evolution, 85, 59–67.
http://dx.doi.org/10.1016/j.ympev.2015.02.003Nguyen, S.V., Ho, C.T. & Nguyen, T.Q. (2009) Herpetofauna of Vietnam. Edition Chimaira, Frankfurt am Main, 768 pp.
Orlov, N.L. (1997) Die Theloderma–Arten Vietnams (Rhacophoridae; Anura). Herpetofauna, 19, 5–9.
Orlov, N.L., Dutta, S.K., Ghate, H.V. & Kent, Y. (2006) New species of Theloderma from Kon Tum Province (Vietnam) and Nagaland State (India) [Anura: Rhacophoridae]. Russian Journal of Herpetology, 13, 135–154.
Orlov, N.L. & Ho, C.T. (2005) A new species of Philautus from Vietnam (Anura: Rhacophoridae). Russian Journal of Herpetology, 12, 135–142.
Orlov, N.L. & Rybaltovsky, E.M. (1999) Die Vermehrung von Theloderma corticale (Boulenger, 1903) im Terrarium. Sauria, 21, 17–20.
Poyarkov, N.A. Jr., Orlov, N.L., Moiseeva, A.V., Pawangkhanant, P., Ruangsuwan, T., Vassilieva, A.B., 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.
Pyron, R.A. & Wiens, J.J. (2011) A large-scale phylogeny of Amphibia including over 2800 species, and a revised classification of extant frogs, salamanders, and caecilians. Molecular Phylogenetics and Evolution, 61, 543–583.
http://dx.doi.org/10.1016/j.ympev.2011.06.012Ronquist, 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 mo
http://dx.doi.org/10.1093/sysbio/sys029 del choice across a large model spaces. Systematic Biology, 61, 1–4.Rowley, J.J.L., Chan, S.K.F., Tang, W.S., Speare, R., Skerratt, L.F., Alford, R.A., Cheung, K.S., Ho, C.Y. & Campbell, R. (2007) Survey for the amphibian chytrid Batrachochytrium dendrobatidis in Hong Kong in native amphibians and in the international amphibian trade. Diseases of Aquatic Organisms, 78, 87–95.
http://dx.doi.org/10.3354/dao01861Rowley, J.J.L., Le, D.T.T., Hoang, H.D., Dau, V.Q. & Cao, T.T. (2011) Two new species of Theloderma (Anura: Rhacophoridae) from Vietnam. Zootaxa, 3098, 1–20.
Smith, M.A. (1924) New tree-frogs from Indo-China and the Malay Peninsula. Proceedings of the Zoological Society of London, 1924, 225–234.
http://dx.doi.org/10.1111/j.1096-3642.1924.tb01499.xStuart, B.L. & Heatwole, H.F. (2004) A new Philautus (Amphibia: Rhacophoridae) from northern Laos. Asiatic Herpetological Research, 10, 17–21.
Stuart, B.L., Phimmachak, S., Seateun, S. & Sheridan, J.A. (2013) A new Philautus (Anura: Rhacophoridae) from northern Laos allied to P. abditus Inger, Orlov & Darevsky, 1999. Zootaxa, 3745 (1), 73–83.
http://dx.doi.org/10.11646/zootaxa.3745.1.6Swofford, D.L. (2002) PAUP*: Phylogenetic Analysis Using Parsimony *(and other methods). Version 4.0b10. Sinauer Associates, Sunderland, Massachusetts, USA.
Taylor, E.H. (1962) The amphibian fauna of Thailand. University of Kansas Science Bulletin, 43, 265–599.
http://dx.doi.org/10.5962/bhl.part.13347Tschudi, J.J. von (1838) Classification der Batrachier mit Berücksichtigung der fossilen Thiere dieser Abtheilung der Reptilien. Petitpierre, Neuchâtel, 100 pp.
http://dx.doi.org/10.5962/bhl.title.59545Vassilieva, A.B., Galoyan, E.A., Poyarkov, N.A. Jr. & Geissler, P. (2016) A Photographic Guide to the Amphibians and Reptiles of the Lowland Monsoon Forests of Southern Vietnam. Edition Chimaira, Frankfurt am Main, 324 pp.
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.
http://dx.doi.org/10.1016/S1055-7903(02)00212-9Yu, G., Rao, D., Yang, J. & 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.
http://dx.doi.org/10.1111/j.1096-3642.2008.00404.xYu, G., Rao, D., Zhang, M. & Yang, J. (2009) Re-examination of the phylogeny of Rhacophoridae (Anura) based on mitochondrial and nuclear DNA. Molecular Phylogenetics and Evolution, 50, 571–579.
http://dx.doi.org/10.1016/j.ympev.2008.11.023Ziegler, T. & Nguyen, T.Q. (2008) The amphibian and reptilian breeding station at Hanoi. WAZA (World Association of Zoos and Aquariums) Magazine, 9, 10–12.