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Phylogeny of the Cyrtodactylus irregularis species complex (Squamata: Gekkonidae) from Vietnam with the description of two new species



The number of described species of bent-toed geckos of the Cyrtodactylus irregularis species complex in Vietnam has increased from one to eight in the last six years. We combined morphological and molecular analyses to explore phylogenetic relationships among all described species in the group. The phylogeny required the description of two new species, Cyrtodactylus phuocbinhensis sp. nov. and Cyrtodactylus taynguyenensis sp. nov. Further, the tree resolved two additional undescribed clades that may also be new species. The species C. bugiamapensis and C. ziegleri were found to require redefinition. Cyrtodactylus phuocbinhensis sp. nov. is characterized by a series of enlarged femoral scales separated from preanal scales while Cyrtodactylus taynguyenensis sp. nov. does not possess enlarged femoral scales. Both new species are distinguished from other congeners by a combination of the following characters: small subcaudal scales, not transversely enlarged; presence (C. phuocbinhensis sp. nov.) or absence (C. taynguyenensis sp. nov.) of enlarged femoral scales; number of preanal pores; and dorsal pattern. Genetic distances between described species and new species were 16.5% and 2.0% in COI and RPL35, respectively, for C. phuocbinhensis sp. nov., and these distances were 18.8% and 2.2% for C. taynguyenensis sp. nov., respectively.


bent-toed geckos, COI, RPL35, matrilineal genealogy

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Bain, R.H. & Hurley, M.M. (2011) A biogeographic synthesis of the amphibians and reptiles of Indochina. Bulletin of the American Museum of Natural History, 360, 1–138.

Che, J., Chen, H.M., Yang, J.X., Jin, J.Q., Jiang, K., Yuan, Z.Y., Murphy, R.W. & Zhang, Y.P. (2012) Universal COI primers for DNA barcoding amphibians. Molecular Ecology Resources, 12, 247–258.

Crawford, A.J., Cruz, C., Griffith, E., Ross, H., Ibáñez, R, Lips, K.R, Driskell, A.C., Bermingham, E. & Crump, P. (2013) DNA barcoding applied to ex situ tropical amphibian conservation program reveals cryptic diversity in captive populations. Molecular Ecology Resources, 13 (6), 1005–1018.

David, P., Teynié, A. & Ohler, A. (2004) A new species of Cyrtodactylus Gray 1827 (Reptilia: Squamata: Gekkonidae) from southern Laos. The Raffles Bulletin of Zoology, 52, 621–627.

Felsenstein, J. (2004) Inferring phylogenies. Sinauer Associates, Sunderland, MA, 580 pp.

Francis, C.M., Borisenko, A.V., Ivanova, N.V., Eger, J.L., Lim, B.K., Guillén-Servent, A., Kruskop, S.V., Mackie, I. & Hebert, P.D.N. (2010) The role of DNA barcodes in understanding and conservation of mammal diversity in Southeast Asia. PLoS ONE, 5, e12575.

Geissler, P., Nazarov, R., Orlov, N.L., Böhme, W., Phung, M.T., Nguyen, Q.T. & Ziegler, T. (2009) A new species of the Cyrtodactylus irregularis complex (Squamata: Gekkonidae) from southern Vietnam. Zootaxa, 2161, 20–32.

Goloboff, P.A., Farris, J.S. & Nixon, K.C. (2008) TNT, a free program for phylogenetic analysis. Cladistics, 24, 774–786.

Grismer, L.L., Wood, P.L. & Lim, K.K.P. (2012) Cyrtodactylus majulah, a new species of bent-toed gecko (Reptilia: Squamata: Gekkonidae) from Singapore and the Riau Archipelago. The Bulletin of Zoology, 60, 487–499.

Hanke, M. & Wink, M. (1994) Direct DNA sequencing of PCR-amplified vector inserts following enzymatic degradation of primer and dNTPs. Biotechniques, 17, 858–860.

Heidrich, A., Rösler, H., Vu, N.T., Böhme, W. & Ziegler, T. (2007) Another new Cyrtodactylus (Squamata: Gekkonidae) from Phong Nha–Ke Bang National Park, central Truong Son, Vietnam. Zootaxa, 1445, 35–48.

Hillis, D.M. & Bull, J.J. (1993) An empirical test of bootstrapping as a method for assessing confidence in phylogenetic analysis. Systematic Biology, 42, 182–192.

Johnson, C.B., Quah, E.S.H., Anuar, S., Muin, M.A., Wood, P.L.J.R., Grismer, J.L., Greer, L.F., Chan, K.O., Ahmad, N., Bauer, A.M. & Grismer, L.L. (2012) Phylogeography, geographic variation, and taxonomy of the bent-toed gecko Cyrtodactylus quadrivirgatus Taylor, 1962 from Peninsular Malaysia with the description of a new swamp dwelling species. Zootaxa, 3406, 39–58.

Kimura, M. (1980) A simple method for estimating evolutionary rates of base substitutions through comparative studies of nucleotide sequences. Journal of Molecular Evolution, 16, 111–120.

Murphy, R.W., Crawford, A.J., Bauer, A.M., Che, J., Donnellan, S.C., Fritz, U., Haddad, C.F.B., Nagy, Z.T., Poyarkov, N.A., Vences, M., Wang, W.-Z. & Zhang, Y.-P. (2013) Cold Code: the global initiative to DNA barcode amphibians and nonavian reptiles. Molecular Ecology Resources, 13, 161–167.

Nazarov, A.R., Orlov, N.L., Nguyen, N.S. & Ho, T.C. (2008) Taxonomy of naked-toes geckos Cyrtodactylus irregularis complex of South Vietnam and description of a new species from Chu Yang Sin Natural Park (Krong Bong District, Dac Lac Province, Vietnam). Russian Journal of Herpetology, 15, 141–156.

Nazarov, R., Poyarkov, N.A., Orlov, N.L., Phung, T.M., Nguyen, T.T., Hoang, D.M. & Ziegler, T. (2012) Two new cryptic species of the Cyrtodactylus irregularis complex (Squamata: Gekkonidae) from southern Vietnam. Zootaxa, 3302, 1–24.

Ngo, V.T. & Bauer, A.M. (2008) Descriptions of two new species of Cyrtodactylus Gray 1827 (Squamata: Gekkonidae) endemic to southern Vietnam. Zootaxa, 1715, 27–42.

Ngo, V.T. & Grismer, L.L. (2012) A new endemic species of Cyrtodactylus (Squamata: Gekkonidae) from Tho Chu Island, southeastern Vietnam. Zootaxa, 3228, 48–60.

Nylander, J.A.A. (2004) MrModeltest v2. Program distributed by the author. Evolutionary Biology Centre, Uppsala University. [Software]

Orlov, N.L., Poyarkov, N.A., Vassilieva, A.B., Ananjeva, N.B., Nguyen, T.T., Nguyen, N.S. & Geissler, P. (2012) Taxonomic notes on rhacophorid frogs (Rhacophorinae: Rhacophoridae: Anura) of southern part of Annamite Mountains (Truong Son, Vietnam), with description of three new species. Russian Journal of Herpetology, 19, 23–64.

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

Rösler, H., Vu, N.T., Nguyen, Q.T., Ngo, V.T. & Ziegler, T. (2008) A new Cyrtodactylus (Squamata: Gekkonidae) from central Vietnam. Hamadryad, 33, 48–63.

Sambrook, J., Fritsch, E.F. & Maniatis, T. (1989) Molecular Cloning: A Laboratory Manual. Cold Spring Harbor Laboratory Press, Cold Spring Harbor, New York, 1626 pp.

Siler, C.D., Oask, J.R., Esselstyn, J.A., Diesmos, A.C. & Brown, R.M. (2010) Phylogeny and biogeography of Philippine bent-toed geckos (Gekkonidae: Cyrtodactylus) contradict a prevailing model of Pleistocene diversification. Molecular Phylogenetics and Evolution, 55, 699–710.

Simpson, G.G. (1961) Principles of Animal Taxonomy. Columbia University Press, New York, 247 pp.

Smith, M.A. (1921) New or little-known reptiles and batrachians from southern Annam (Indo-China). Proceedings of the Zoological Society of London, 11, 423–440.

Smith, M.A. (1935) The Fauna of British India, including Ceylon and Burma. Reptilia and Amphibia, Vol. II—Sauria. Taylor and Francis Ltd., London.

Song, H., Buhay, J.E., Whiting, M.F. & Crandall, K.A. (2008) Many species in one: DNA barcoding overestimates the number of species when nuclear mitochondrial pseudogenes are coamplified. Proceedings of the National Academy of Sciences U.S.A., 105, 13486–13491.

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

Tamura, K., Dudley, J., Nei, M. & Kumar, S. (2007) MEGA4: molecular evolutionary genetics analysis (MEGA) software version 4.0. Molecular Biology and Evolution, 24, 1596–1599.

Taylor, E.H. (1963) The lizards of Thailand. University of Kansas Science Bulletin, 44, 687–1077.

Thompson, J.D, Higgins, D.G. & Gibson, T.J. (1994) CLUSTAL W: improving the sensitivity of progressive multiple sequence alignment through sequence weighting, position-specific gap penalties and weight matrix choice. Nucleic Acids Research, 22, 4673–4680.

Vieites, D.R., Wollenberg, K.C., Andreone, F., Köhler J., Glaw, F. & Vences, M. (2009) Vast underestimation of Madagascar’s biodiversity evidenced by an integrative amphibian inventory. Proceedings of the National Academy of Sciences U.S.A., 106, 8267–8272.

Welton, L.J., Siler, C.D, Diesmos, A.C. & Brown, R.M. (2010a) Phylogeny-based species delimitation of southern Philippines bent-toed geckos and a new species of Cyrtodactylus (Squamata: Gekkonidae) from western Mindanao and the Sulu Archipelago. Zootaxa, 2390, 49–68.

Welton, L.J., Siler, C.D., Linkem, C.W., Diesmos, A.C. & Brown, R.M. (2010b) Philippine bent-toed geckos of the Cyrtodactylus agusanensis complex: multilocus phylogeny, morphological diversity, and descriptions of three new species. Herpetological Monographs, 24, 55–85.

Wiley, E.O. & Lieberman, B.S. (2011) Phylogenetics: Theory and Practice of Phylogenetic Systematics. Wiley-Blackwell, Singapore, 432 pp.

Wood, P.L. Jr., Heinicke, M.P., Jackman, T.R. & Bauer, A.M. (2012) Phylogeny of bent-toed geckos (Cyrtodactylus) reveals a west to east pattern of diversification. Molecular Phylogenetics and Evolution, 65, 992–1003.

=He"}gup��� ='margin-top:0cm;margin-right:0cm;margin-bottom: 0cm;margin-left:18.0pt;margin-bottom:.0001pt;text-align:justify;text-indent: -18.0pt;line-height:11.0pt;mso-pagination:none;page-break-after:auto; mso-hyphenate:none;tab-stops:18.0pt 35.0pt 70.0pt 105.0pt 141.0pt 176.0pt 212.0pt 247.0pt 282.0pt 318.0pt 353.0pt 388.0pt 424.0pt'>Jacoby, M. (1892) Description of the new genera and species of the phytophagous Coleoptera obtained by Sign. L. Fea in Burma. Annali del Museo Civico di Storia Naturale di Genova, 32, 869–999.

Kasap, H. & Crowson, R.A. (1985) The studies on the ovipositors and 8th abdominal segments of species of Bruchidae and Chrysomelidae (Coleoptera). Türkiye Bitkiler koruma Dergisi, 9 (3), 131–145.

Kimoto, S. (2005) Systematic Catalog of the Chrysomelidae (Coleoptera) from Nepal and Bhutan. Bulletin of the Kitakyushu Museum of Natural History and Human History Series A, 3, 13–114.

Kimoto, S. & Gressitt, J.L. (1979) Chrysomelidae (Coleoptera) of Thailand, Cambodia, Laos and Vietnam. 1. Sagrinae, Donaciinae, Zeugophorinae, Megalopodinae and Criocerinae. Pacific Insects, 20 (2–3), 191–256.

Komiya, Y. (1986) Notes on the Taiwanese Megalopodinae with description of a new species (Coleoptera, Chrysomelidae). Elytra, Tokyo, 14 (1), 1–9.

Lacordaire, T. (1845) Monographie des coléoptères subpentamères de la famille des phytophages. Tome I. Mémoires de la Société Royale des Sciences de Liége, 3 (1), xiii + 740 pp.

Lawrence, J.F., Beutel, R.G., Leschen, R.A. & Slipinski, A. (2010) Glossary of morphological terms. In: Leschen, R.A., Beutel, R.G. & Lawrence, J.F. (Eds.), Handbook of Zoology, Anthropoda: Insecta, Coleoptera, Beetles, Vol. 2. Morphology and Systematics (Elateroidea, Bostrichiformia, Cucujiformia partim). Walter de Gruyter, Berlin, pp. 9–20.

Marvaldi, A.E., Duckett, C.N., Kier, K.M. & Gillespie, J.J. (2009) Structural alignment of 18S and 28rDNA sequences provides insights into phylogeny of Phytophaga (Coleoptera: Curculionoidea and Chrysomeloidea). Zoologica Scripta, 38, 63–77.

Medvedev, L.N. (2002) Jacoby’s types of Chrysomelidae (Coleoptera) from Burma in the Museo Civico di Storia Naturale “Giacomo Doria”, Genoa. Part 3. Annali del Museo Civico di Storia Naturale “Giacomo Doria”, 94, 249–264.

Medvedev, L.N. (2010) Subfamily Megalopodinae Latreille, 1802. In: Löbl, I. & Smetana, A. (Eds.), Catalogue of Palaearctic Coleoptera, Vol. 6. Apollo Books, Stenstrup, pp. 336–337.

Medvedev, L.N. & Sprecher-Uebersax, E. (1997) Chrysomelidae of Nepal and neighbouring regions. (Coleoptera: Chrysomelidae). Coleoptera, 1, 203–247.

Pic, M. (1914) Nouveaux Coléoptères de diverses familles. Mélanges Exotico-Entomologiques, 10, 7–20.

Pic, M. (1922) Nouveautés diverses. Méanges Exotico-Enomologiques, 36, 1–32.

Reid, C.A.M. (1995) A cladistic analysis of subfamilial relationships in the Chrysomelidae sensu lato (Chrysomeloidea). In: Pakaluk, J., Slipinski, S.A., (Eds.), Biology, phylogeny and classification of Coleoptera: papers celebrating the 80th birthday of Roy A. Crowson. Muzeum i Instytut Zoologii PAN, Warszawa, pp. 559–631.

Reid, C.A.M. (2000) Spilopyrinae Chapuis: a new subfamily in the Chrysomelidae and its systematic placement (Coleoptera). Invertebrate Taxonomy, 14, 837–862.

Silfverberg, H. (2010) Family Megalopodidae Latreille, 1802, subfamily Zeugophorinae Böving & Craighead, 1931. , In: Löbl, I. & Smetana, A. (Eds.), Catalogue of Palaearctic Coleoptera, Vol. 6. Apollo Books, Stenstrup, pp. 334–335.

Snodgrass, R.E. (1935) Principles of Insect Morphology. McGraw-Hill Book Company, New York, 667 pp.

Suzuki, K. (1988) Comparative morphology of the internal reproductive system of the Chrysomelidae (Coleoptera)., In: Jolivet, P., Petitpierre, E. & Hsiao, T.H. (Eds.), Biology of Chrysomelidae. Kluwer Academic Publishers, Dordrecht, pp. 317–355.

Suzuki, K. (1994) Comparative morphology of the hindwing venation of the Chrysomelidae (Coleoptera). In: Jolivet, P., Cox, M.L. & Petitpierre, E. (Eds.), Novel Aspects of the Biology of Chrysomelidae, Series Entomologica, Vol. 50. Kluwer Academic Publishers, Dordrecht, pp. 337–354.

Suzuki, K. (2003) Systematic position of the subfamilies Megalopodinae and Megascelinae (Chrysomelidae) based on the comparative morphology of internal reproductive system. International Congress of Entomology Proceedings, 21, 105–116.

Tan, J.J., Yu, P.Y., Li, H.X., Wang, S.Y. & Jiang, S.Q. (1980) Economic Insect Fauna of China, Coleoptera: Chrysomeloidea (I). Science Press, Beijing, 213 pp.

Yu, P.Y. (1977) On Temnaspis nankinea (Pic) (Megalopodinae: Chrysomeloidea), a new pest of Fraxinus chinensis Roxb. Acta Entomologica Sinica, 20 (4), 482–484.

Yu, P.Y. & Yang, X.K. (1994) Biological studies on Temnaspis nankinea (Pic) (Chrysomelidae: Megalopodinae). In: Jolivet, P.H., Cox, M.L. & Petitpierre, E. (Eds.), Novel Aspects of the Biology of Chrysomelidae, Series Entomologica, Vol. 50. Kluwer Academic Publishers, Dordrecht, pp. 527–531.

Yu, P.Y. & Liang, H.B. (2002) A check-list of the Chinese Megalopodinae (Coleoptera: Chrysomelidae). Oriental Insects, 36, 117–128.

y, S%z99@��h�rsity of tropical Homoptera, with the first data from Africa. American Entomologist, 45, 213–222.

Nonnotte, P., Guillou, H., Le Gall, B., Benoit, M., Cotten, J. & Scaillet, S. (2008) New K–Ar age determinations of Kilimanjaro volcano in the North Tanzanian diverging rift, East Africa. Journal of Volcanology and Geothermal Research, 173, 99–112.

Ragge, D.R. (1956) A revision of the genera Phaneroptera Serville and Nephoptera Uvarov (Orthoptera: Tettigoniidae), with conclusions of zoogeographical and evolutionary interest. Proceedings of the zoological Society London, 27, 205–283.

Ragge, D.R. (1961) A revision of the genus Ducetia Stål (Orthoptera: Tettigoniidae). Bulletin of the British Museum (Natural History) Entomology, 10 (5), 13–208.

Ragge, D.R. (1968) An index-catalogue of African Phaneropterinae (Orthoptera: Tettigoniidae). Bulletin of the British Museum (Natural History), 22 (3), 76–108.

Ragge, D.R. (1980) A review of the African Phaneropterinae with open tympana (Orthoptera: Tettigoniidae). Bulletin of the British Museum (Natural History) Entomology, 40 (2), 1–192.

Rodgers, W.A. & Homewood, K.M. (1982) The conservation of the East Usambara Mountains, Tanzania: a review of biological values and land use pressures. Biological Journal of the Linnean Society, 24, 285–304.

Schultz, O., Hemp, C., Hemp, A. & Wägele, W. (2007) Molecular phylogeny of the endemic East African flightless grasshoppers Altiusambilla Jago 1981, Usambilla (Sjöstedt 1909) and Rhainopomma Jago 1981 (Orthoptera: Acridoidea). Systematic Entomology, 32 (4), 712–719.

Trauth, M.H., Maslin, M.A., Deino, A. & Strecker, M.R. (2005) Late cenozoic moisture history of East Africa. Science, 309, 2051– 2053.

Vosseler, J. (1909) Die Gattung Myrmecophana Brunner. Ihre hypertelische und Ameisen-Nachahmung. Zoologische Jahrbücher. Abtheilung für Systematik, Geographie und Biologie der Tiere, 27, 157–210.

Wattenwyl, B. von (1891) Additamenta zur Monographie der Phaneropteriden. Verhandlungen der Kaiserlich-Königlichen Zoologisch-Botanischen Gesellschaft in Wien, 41, 1–196, pl. 1–2.


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