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Article
Published: 2019-07-16

Geographic structure of genetic variation in the Parachute Gecko Ptychozoon lionotum Annandale, 1905 across Indochina and Sundaland with descriptions of three new species

Herpetology Laboratory, Department of Biology, La Sierra University, 4500 Riverwalk Parkway, Riverside, California 92515, USA.
Department of Biological Sciences & Museum of Natural History, Auburn University, Auburn, Alabama 36849, USA.
Herpetology Laboratory, Department of Biology, La Sierra University, 4500 Riverwalk Parkway, Riverside, California 92515, USA.
School of Biological Sciences, Universiti Sains Malaysia, 11800 Minden, Penang, Malaysia.
Wild Earth Allies, #77a, St. Betong, Bayab Village, Sk. Phnom Penh Thmei, Kh. Sen Sok, Phnom Penh, Cambodia.
National University of Laos, Faculty of Natural Sciences, Department of Biology, P.O. Box 2273, Dong Dok Campus, Vientiane, Laos.
National University of Laos, Faculty of Natural Sciences, Department of Biology, P.O. Box 2273, Dong Dok Campus, Vientiane, Laos.
National University of Laos, Faculty of Natural Sciences, Department of Biology, P.O. Box 2273, Dong Dok Campus, Vientiane, Laos.
Section of Research & Collections, North Carolina Museum of Natural Sciences, Raleigh, NC, USA.
Sam Noble Oklahoma Museum of Natural History and Department of Biology, University of Oklahoma, Norman, Oklahoma, 73072-7029, USA.
Global Genome Initiative, National Museum of Natural History, Smithsonian Institution, Washington, DC, 20013, USA.
Department of Ecology and Evolutionary Biology and Biodiversity Institute, University of Kansas, Dyche Hall, 1345 Jayhawk Blvd, Lawrence, Kansas 66045-7561, USA.
Department of Ecology and Evolutionary Biology and Biodiversity Institute, University of Kansas, Dyche Hall, 1345 Jayhawk Blvd, Lawrence, Kansas 66045-7561, USA.
Reptilia Asia Gekkonidae gliding integrative taxonomy phylogeography species complex

Abstract

An integrative taxonomic analysis of the Ptychozoon lionotum group across its range in Indochina and Sundaland recovers P. lionotum sensu lato Annandale, 1905 as paraphyletic with respect to P. popaense Grismer, Wood, Thura, Grismer, Brown, & Stuart, 2018a and composed of four allopatric, genetically divergent, ND2 mitochondrial lineages. Multivariate and univariate analyses of continuous and discrete morphological and color pattern characters statistically and discretely diagnose each lineage from one another and together, with maximum likelihood and Bayesian inference analyses, provide the foundation for the recognition of each lineage as a new species—hypotheses corroborated with a Generalized Mixed Yule Coalescent species delimitation analysis. Ptychozoon cicakterbang sp. nov. ranges throughout Peninsular Malaysia to Pulau Natuna Besar, Indonesia; P. kabkaebin sp. nov. is endemic to northern and central Laos; and P. tokehos sp. nov. ranges from southern Thailand south of the Isthmus of Kra northward to Chiang Mai, fringing the Chao Phraya Basin and ranging southward through Cambodia to southern Vietnam. Ptychozoon lionotum sensu stricto ranges from northwestern Laos through southern Myanmar to eastern India. The phylogeographic structure within each species varies considerably with P. lionotum s.s. showing no genetic divergence across its 1,100 km range compared to P. cicakterbang sp. nov. showing upwards of 8.2% sequence divergence between syntopic individuals. Significant phylogeographic structure exists within P. tokehos sp. nov. and increased sampling throughout Thailand may require additional taxonomic changes within this species.

 

References

  1. Acinas, S.G., Klepac-Ceraj, V., Hunt, D.E., Pharino, C., Ceraj, I., Distel, D.L. & Polz, M.F. (2004) Fine-scale phylogenetic architecture of a complex bacterial community. Nature, 430, 551–554.

    https://doi.org/10.1038/nature02649

    Agarwal, I., Bauer, A.M., Jackman, T.R. & Karanth, K.P. (2014) Insights into Himalayan biogeography from geckos: a molecular phylogeny of Cyrtodactylus (Squamata: Gekkonidae). Molecular Phylogenetics and Evolution, 80, 145–155. https://doi.org/10.1016/j.ympev.2014.07.018

    Annandale, N. (1905) Notes on some Oriental geckos in the Indian Museum, Calcutta, with descriptions of new forms. Annals & Magazine of Natural History, Series 7, 15, 26–32.

    https://doi.org/10.1080/03745480509443635

    Barraclough, T.G., Birky, C.W. Jr. & Burt, A. (2003) Diversification in sexual and asexual organisms. Evolution, 57, 2166–2172.

    https://doi.org/10.1554/02-339

    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, 2, 304–345.

    Boulenger, G.A. (1899) Description of three new reptiles and a new batrachian from Mt. Kina Balu, North Borneo. Annals and Magazine of Natural History, 4, 451–453.

    https://doi.org/10.1080/00222939908678228

    Boulenger, G.A. (1912) A vertebrate fauna of the Malay Peninsula from the Isthmus of Kra to Singapore including the adjacent islands. Reptilia and Batrachia. Taylor & Francis, London, 294 pp.

    https://doi.org/10.5962/bhl.title.10813

    Bourret, R. (2009) Les Lézards de L’Indochine. Edition Chimaira, Frankfurt am Main, 624 pp. [2009 reprinting]

    Brown, R.M. (1999) New species of parachute gecko (Squamata: Gekkonidae: genus Ptychozoon) from northeastern Thailand and central Vietnam. Copeia, 1999, 990–1001.

    https://doi.org/10.2307/1447974

    Brown, R.M., Diesmos, A.C. & Oliveros, C.H. (2011) New flap-legged forest gecko (Genus Luperosaurus) from the Northern Philippines. Journal of Herpetology, 45 (2), 202–210.

    https://doi.org/10.1670/10-123.1

    Brown, R.M, Ferner, J.W. & Diesmos, A.C. (1997) Definition of the Philippine parachute gecko, Ptychozoon intermedium Taylor 1915 (Reptilia: Squamata: Gekkonidae): redescription, designation of a neotype, and comparisons with related species. Herpetologica, 53, 357–373.

    Brown, R.M., Siler, C.D., Grismer, L.L., Das, I. & McGuire, J.A. (2012) Phylogeny and cryptic diversification in Southeast Asian flying geckos. Molecular Phylogenetics and Evolution, 65, 351–361.

    https://doi.org/10.1016/j.ympev.2012.06.009

    Brown, R.M., Supriatna, J. & Ota, H. (2000) Discovery of a new species of Luperosaurus (Squamata: Gekkonidae) from Sulawesi, with a phylogenetic analysis of the genus, and comments on the status of Luperosaurus serraticaudus. Copeia, 2000, 191–209.

    https://doi.org/10.1643/0045-8511(2000)2000[0191:doanso]2.0.co;2

    Cangelosi, R. & Goriely, A. (2007) Component retention in principal component analysis with application of cDNA microarray data. Biology Direct, 2, 2.

    https://doi.org/10.1186/1745-6150-2-2

    Cantor, T.E. (1847) Catalogue of reptiles inhabiting the Malayan Peninsula and islands, collected or observed by Theodore Cantor, Esq., M.D. Bengal Medical Services. Journal of the Asiatic Society of Bengal, 16, 607–656 + 897–952 + 1026–1078.

    https://doi.org/10.5962/bhl.title.5057

    Chan-ard, T., Grossmann, W., Gumprecht, A. & Schultz, K.-D. (1999) Amphibians and Reptiles of Peninsular Malaysia and Thailand. Bushmaster Publications, Wuerselen, 240 pp.

    https://doi.org/10.15560/10.2.335

    Chiari, Y. & Claude, J. (2012) Morphometric identification of individuals when there are more shape variables than reference specimens: A case study in Galápagos tortoises. Comptes Rendus Biologies, 355, 62–68.

    https://doi.org/10.1016/j.crvi.2011.10.007

    Cox, M.J., van Dijk, P.P., Nabhitabhata, J. & Thirakhupt, K. (1998) A Photographic Guide to Snakes and Other Reptiles of Peninsular Malaysia, Singapore and Thailand. New Holland Publishers (UK) Ltd., London, 144 pp.

    https://doi.org/10.2307/1447416

    Coyne, J.A. & Orr, H.A. (1998) The evolutionary genetics of speciation. Philosophical Transactions of the Royal Society of London B, 353, 287–305.

    https://doi.org/0.1098/rstb.1998.0210

    Das, I., Dattagupta, B. & Gayen, N.C. (1998) History and catalogue of reptile types in the collection of the Zoological Survey of India. Journal of South Asian Natural History, 3, 121–172.

    Das, I. & Vijayakumar, S.P. (2009) New species of Ptychozoon (Sauria: Gekkonidae) from the Nicobar Archipelago, Indian Ocean. Zootaxa, 2095, 8–20.

    de Queiroz, K. (2007) Species concepts and species delimitation. Systematic Biology, 56, 879–886.

    https://doi.org/10.1080/10635150701701083

    Dring, J.C.M. (1979) Amphibians and reptiles from northern Trengganu, Malaysia, with descriptions of two new geckos: Cnemaspis and Cyrtodactylus. Bulletin of the British Museum (Natural History), 34, 181–241.

    Drummond, A.J., Ashton, B., Buxton, S., Cheung, M., Cooper, A., Duran, C., Field, M., Heled, J., Kearse, M., Markowitz, S., Moir, R., Stones-Havas, S., Sturrock, S., Thierer, T. & Wilson, A. (2011) Geneious. Version 5.6. Available from: http://www.geneious.com/ (accessed 20 May 2019)

    Drummond, A.J., Suchard, M.A., Xie, D. & Rambaut, A. (2012) Bayesian phylogenetics with BEAUti and BEAST 1.7. Molecular Biology and Evolution, 29, 1969–1973.

    https://doi.org/10.1093/molbev/mss075

    Dudley, R., Byrnes, G., Yanoviak, S.P., Borrell, B., Brown, R.M. & McGuire, J.A. (2007) Gliding and the functional origins of flight: biomechanical novelty of necessity? Annual Review of Ecology, Evolution, and Systematics, 38, 179–201.

    https://doi.org/10.1146/annurev.ecolsys.37.091305.110014

    Fea, L. (1897) Viaggio di Leonardo Fea in Birmania e regioni vicine. LXXVI. Riassunto generale dei risultati zoologici. Annali del Museo Civico di Storia Naturale di Genova, Series 2, 42, 383–660.

    https://doi.org/10.5962/bhl.part.16764

    Fisher-Reid, M.C. & Wiens, J.J. (2011) What are the consequences of combining nuclear and mitochondrial data for phylogenetic analysis? Lessons from Plethodon salamanders and 13 other vertebrate clades. BMC Evolutionary Biology, 11, 1–20. https://doi.org/10.1186/1471-2148-11-300

    Fitzsimons, J.A. (2017) Ecological notes on the three-banded parachute gecko Ptychozoon trinotaterra in southern Vietnam. Russian Journal of Herpetology, 24, 327–328.

    https://doi.org/10.1146/annurev.ecolsys.37.091305.110014

    Fontaneto, D., Herniou, E.A., Boschetti, C., Caprioli, M., Melone, G., Ricci, C. & Barraclough, T.G. (2007) Independently evolving species in asexual bdelloid rotifers. PLoS Biology, 5, e87.

    https://doi.org/10.1371/journal.pbio.0050087

    Fujisawa, T. & Barraclough, T.G. (2013) Delimiting species using single-locus data and the Generalized Mixed Yule Coalescent approach: a revised method and evaluation on simulated data sets. Systematic Biology, 62 707–724.

    https://doi.org/10.1093/sysbio/syt033

    Gray, J.E. (1827) A synopsis of the genera of saurian reptiles in which some new genera are indicated, and the others reviewed by actual examination. Philosophical Magazine, London, 2, 54–58.

    https://doi.org/10.1080/14786442708675620

    Gray, J.E. (1845) Catalogue of the Specimens of Lizards in the Collection of the British Museum. Trustees of die British Museum, London, xxvii + 289 pp.

    https://doi.org/10.5962/bhl.title.5499

    Grismer, L.L. (2011a) Lizards of Peninsular Malaysia, Singapore and their Adjacent Archipelagos. Edition Chimaira, Frankfurt am Main, 728 pp.

    Grismer, L.L. (2011b) Amphibians and Reptiles of the Seribuat Archipelago. Edition Chimaira, Frankfurt am Main, 129 pp.

    Grismer, L.L., Chav, T., Neang, T., Wood Jr., P.L., Grismer, J.L., Youmans, T.M., Ponce, A., Daltry, J.C. & Kaiser, H. (2007) The herpetofauna of the Phnom Aural Wildlife Sanctuary and checklist of the herpetofauna of the Cardamom Mountains, Cambodia. Hamadryad, 31, 216–241.

    Grismer, L.L. & Davis, H.R. (2018c) Phylogeny and biogeography of bent-toed geckos (Cyrtodactylus Gray) of the Sundaic swamp clade. Zootaxa, 4472 (2), 365–374.

    https://doi.org/10.11646/zootaxa.4472.2.9

    Grismer, L.L., Grismer, J.L., Wood Jr., P.L., Ngo, V.T., Neang, T. & Chan, K.O. (2011) Herpetology on the fringes of the Sunda Shelf: a discussion of discovery, taxonomy, and biogeography. In: Schuchmann, K.-L. (Ed.), Tropical Vertebrates in a Changing World. Bonner Zoologische Monographien, Bonn, pp. 57–97.

    Grismer, L.L., Neang, T., Chav, T. Wood Jr., P.L., Oaks, J.R., Holden, J., Grismer, J.L., Szutz, T.R. & Youmans, T.M. (2008) Additional amphibians and reptiles from the Phnom Samkos Wildlife Sanctuary in the northwestern Cardamom Mountains, Cambodia, with comments on their taxonomy and the discovery of three new species. Raffles Bulletin of Zoology, 56, 161–175.

    Grismer, L.L. & Pan, K.A. (2008) Diversity, endemism, and conservation of the amphibians and reptiles of southern Peninsular Malaysia and its offshore islands. Herpetological Review 39, 270–281.

    Grismer, L.L. & Quah, E.S.H. (2019) An updated and annotated checklist of the lizards of Peninsular Malaysia, Singapore, and their adjacent islands. Zootaxa, 4545 (2), 230–248.

    https://doi.org/10.11646/zootaxa.4545.2.4

    Grismer, L.L., Wood, P.L. Jr., Myint, K.T., Grismer, M.S., Brown, R.M. & Stuart, B.L. (2018a) Geographically structured genetic variation in Ptychozoon lionotum (Squamata: Gekkonidae) and a new species from an isolated volcano in Myanmar. Zootaxa, 4514 (2), 202–214.

    https://doi.org/10.11646/zootaxa.4514.2.4

    Grismer, L.L., Wood. Jr., P.L., Myint, K.T., Zin, T., Quah, E.S.H., Murdoch, M.L., Grismer, M.S., Lin, A., Kyaw, H. & Ngwe, L. (2017) Twelve new species of Cyrtodactylus Gray (Squamata: Gekkonidae) from isolated limestone habitats in east-central and southern Myanmar demonstrate high localized diversity and unprecedented microendemism. Zoological Journal of the Linnean Society, 182, 862–959.

    https://doi.org/10.1093/zoolinnean/zlx057

    Grismer, L.L., Wood Jr., P.P., Quah, E.S.H., Murdoch, M.L., Grismer, M.S., Herr, M.W., Espinoza, R.E., Brown, R.M. & Lin, A. (2018b) A phylogenetic taxonomy of the Cyrtodactylus peguensis group (Reptilia: Squamata: Gekkonidae) with descriptions of two new species from Myanmar. PeerJ, 6, e5575.

    https://doi.org/10.7717/peerj.5575

    Grismer L.L., Wood Jr., P.L., Shahrul, A., Awal, R., Norhayati, A., Muin, M., Sumontha, M.,

    Grismer, J., Chan, K.O., Quah, E.S.H. & Pauwels, O.S.G. (2014) Systematics and natural history of Southeast Asian rock geckos (genus Cnemaspis Strauch, 1887) with descriptions of eight new species from Malaysia, Thailand, and Indonesia. Zootaxa, 3880 (1), 1–147.

    https://doi.org/10.11646/zootaxa.3880.1.1

    Grismer, L.L., Youmans, T.M., Wood Jr. P.L. & Grismer, J.L. (2006) Checklist of the herpetofauna of the Seribuat Archipelago, West Malaysia with comments on biogeography, natural history and adaptive types. Raffles Bulletin of Zoology, 54, 157–180.

    Grismer, L.L., Zaharil, D., Muin, M.A., Quah, E.H., Karin, B.R., Anwar, S. & Freitas, E.S, (2019) A new skink of the genus Subdoleseps (Hardwicke & Gray, 1828) from Peninsular Malaysia. Zootaxa. [in press]

    Hartmann, T., Betts, A.B., de Greef, S. & Ihlow, F. (2014) First record of the rare parachute gecko Ptychozoon trinotaterra Brown, 1999 from Cambodia. Cambodia Journal of Natural History, 1, 8–11.

    Herr, M.W. & Lee, D.S. (2016) New provincial record and range extension of the parachute gecko Ptychozoon lionotum Annandale, 1905 in Cambodia, with notes on habitat use. Cambodian Journal of Natural History, 2, 111–113.

    Heyer, W.R. & Pongsapipatana, S. (1970) Gliding speeds of Ptychozoon lionatum (Reptilia: Gekkonidae) and Chrysopelea ornata (Reptilia: Colubridae). Herpetologica, 26, 317–331.

    Hoang, D.T., Chernomor, O., von Haeseler, A., Minh, B.Q. & Le, S.V. (2017) UFBoot2: Improving the Ultrafast Bootstrap Approximation. Molecular Biology and Evolution, 35, 518–522.

    https://doi.org/10.1093/molbev/msx281

    Houttuyn, M. (1782) Het onderscheid der salamanderen van de haagdissen in ‘t algemeen, en van de gekkos in ‘t byzonder aangetoond. Verhandelingen Uitgegeven door het Zeeuwsch Genootschap der Wetenschappen te Vlissingen, Series 1, 9, 305–336.

    Huelsenbeck, J.P., Ronquist, F., Nielsen, R. & Bollback, J.P. (2001) Bayesian Inference of phylogeny and its impact on evolutionary biology. Science, 294, 2310–2314. https://doi.org:10.1126/science.1065889

    Jackson, D.A. (1993) Stopping rules in principle component analysis: a comparison of heuristical and statistical approaches. Ecology, 74, 2204–2214.

    https://doi.org/10.2307/1939574

    Jombart, T., Devillard, S. & Balloux, F. (2010) Discriminant analysis of principal components: a new method for the analysis of genetically structured populations. BMC Genetics, 11, 94.

    https://doi.org/10.1186/1471-2156-11-94

    Katoh, M. &Kuma, M. (2002) MAFTT: a novel method for rapid sequence alignment based on fast Fourier transform. Nucleic Acids Research, 30, 3059–3066. https://doi.org/10.1093/nar/gkf436

    Kluge, A.G. (2001) Gekkonid lizard taxonomy. Hamadryad, 26, 1–209.

    Knowles, L.L. & Carstens, B.C. (2007) Delimiting species without monophyletic gene trees. Systematic Biology, 56, 887–895.

    https://doi.org/10.1080/10635150701701091

    Kuhl, H. & van Hasselt, J.C. (1822) Aus einem Schreiben von Dr. Kuhl und Dr. van Hasselt auf Java. An Professor Th. van Swideren zu Gröningen. Isis von Oken, Jena, 6, 472–476.

    Kumar, S., Stecher, G. & Tamura, K. (2016) MEGA7: Molecular evolutionary genetics analysis Version 7.0 for bigger datasets. Molecular Biology and Evolution, 33, 1870­–1874.

    https://doi.org/10.1093/molbev/msw054

    Kunya, K., Pauwels, O.S.G., Sairum, P. & Taokratok, A. (2011) Squamata, Gekkonidae, Ptychozoon trinotaterra Brown, 1999: rediscovery in Thailand. Check List, 7, 820–822.

    https://doi.org/10.15560/11043

    Leaché, A.D., Koo, M.S., Spencer, C.L., Papenfuss, T.J., Fisher, R.N. & McGuire, J.A. (2009) Quantifying ecological, morphological, and genetic variation to delimit species in the coast horned lizard species complex (Phrynosoma). Proceedings of the National Academy of Sciences, 106, 12418–12423.

    https://doi.org/10.1073/pnas.0906380106

    Leong, T.-M., Grismer, L.L. & Mumpuni (2003) Preliminary checklists of the Anambas and Natuna Islands (South China Sea). Hamadryad, 27, 65–174.

    Lleonart, J., Salat, J. & Torres, G.J. (2000) Removing allometric effects of body size in morphological analysis. Journal of Theoretical Biology, 205, 85–93.

    https://doi.org/10.1006/jtbi.2000.2043

    Macey, J.J., Larson, A., Ananjeva, N.B., Fang, Z. & Papenfuss, T.J. (1997) Two novel gene orders and the role of light-strand replication in rearrangement of the vertebrate mitochondrial genome. Molecular Biology and Evolution, 14, 91–104.

    https://doi.org/10.1093/oxfordjournals.molbev.a025706

    Maddison, W.P. & Maddison, D.R. (2015) Mesquite: a modular system for evolutionary analysis. Version 3.04. Available from: http://mesquiteproject.org (accesed 20 May 2019)

    Malhotra, A., Thorpe, R.S., Mrinalini & Stuart, B.L. (2011) Two new species of pitviper of the genus Cryptelytrops Cope 1860 (Squamata: Viperidae: Crotalinae) from Southeast Asia. Zootaxa, 2757 (1), 1–23.

    https://doi.org/10.11646/zootaxa.2757.1.1

    Manthey, U. & Grossmann, W. (1997) Amphibien & Reptilien Südostasiens. Natur und Tier Verlag, Münster, 512 pp.

    Manthey, U. & Manthey, S. (2017) Amphibien und Reptilian von Laos—ein Reisebericht. Teil 1: Phou Khao Khouay NBCA (Februar 1998). Sauria, 39, 35–55.

    Marcellini, D.L. & Keefer, T.E. (1976) Analysis of the gliding behavior of Ptychozoon lion-atum (Reptilia: Gekkonidae). Herpetologica, 32, 362–66.

    Miller, M.A., Pfeiffer, W. & Schwartz, T. (2010) Creating the CIPRES Science Gateway for inference of large phylogenetic trees. In: Proceedings of the Gateway Computing Environments Workshop (GCE), New Orleans, LA, 14 November 2010, pp. 1–8. https://doi.org/10.1109/GCE.2010.5676129

    Murdoch, M.L., Grismer, L.L., Wood Jr., P.L., Neang, T., Poyarkov, N.A., Ngo, V.T., Nazarov, R., Aowphol, A., Pauwels, O.S.G., Nguyen, H.N. & Grismer, J.L. (2019) Six new species of the Cyrtodactylus intermedius complex (Squamata: Gekkonidae) intermedius complex from the Cardamom Mountains and associated highlands of Southeast Asia. Zootaxa, 4554 (1), 1–62.

    https://doi.org/10.11646/zootaxa.4554.1.1

    Nguyen, L.-T., Schmidt, H.A., von Haeseler, A. & Minh, B.Q. (2015) IQ-TREE: A fast and effective stochastic algorithm for estimating maximum likelihood phylogenies. Molecular Biology and Evolution, 32, 268–274.

    https://doi.org/10.1093/molbev/msu300

    Nguyen, V.S., Ho. T.C. & Nguyen, T.Q. (2009) Herpetofauna of Vietnam. Edition Chimaira, Frankfurt am Main, 768 pp.

    Oksanen, J.G., Blanchet, F.G., Friendly, M., Kindt, R., Legendre, P., McGlinn, D., Minchin, P.R., O’Hara, R.B., Simpson, G.L., Solymos, P., Henry, M., Stevens, H., Szoecs, E. & Wagner, H. (2018) Community ecology package. Available from: https://github.com/vegandevs/vegan (accessed 28 January 2019)

    Pauwels, O.S.G., Laohawat, O.-A., David, P., Bour, R., Dangsee, P., Puangjit, C. & Chimsunchart, C. (2000) Herpetological investigations in Phang-Nga Province, southern peninsular Thailand, with a list of reptile species and notes on their biology. Dumerilia, 4 (2), 123–154.

    Pauwels, O.S.G., Laohawat, O.-A., Naaktae, W., Puangjit, C., Wisutharom, T., Chimsunchart, C. & David, P. (2002) Reptile and amphibian diversity in Phang-nga Province, southern Thailand. Natural History Journal of Chulalongkorn University, 2 (1), 25–30.

    Pawar, S. & Biswas, S. (2001) First record of the Smooth-backed Parachute Gecko Ptychozoon lionotum Annandale 1905 from the Indian mainland. Asiatic Herpetological Research, 9, 101–106.

    Pons, J., Barraclough, T.G., Gomez-Zurita, J., Cardoso, A., Duran, D.P, Hazell, S., Kamoun, S., Sumlin, W.D. & Vogler, A.P. (2006) Sequence-based species delimitation for the DNA taxonomy of undescribed insects. Systematic Biology, 55, 595–609. https://doi.org/10.1080/10635150600852011

    Quah, E.S.H & Anuar, S. (2018) Amphibians and reptiles of the Malaysian-Thai border regions. Journal of Wildlife and Parks, 33, 15–29.

    R Core Team (2015) R: A language and environment for statistical computing. R Foundation for Statistical Computing, Vienna. Available from: http://www.R-project.org (accessed 29 December 2016)

    Rambaut, A. & Drummond, A.J. (2013) TreeAnnotator. Version 1.8.0. MCMC Output Analysis. Available from: http://beast.community/treeannotator (accessed 20 May 2019)

    Rambaut, A., Suchard, M.A., Xie, D. & Drummond, A.J. (2014) Tracer. Vol. 1.6. Available from http//dx.doi.org:beast.bio.ed.ac.uk/Tracer (accessed 20 May 2019)

    Ripley, B., Venables, B., Bates, D.M., Hornik, K., Gebhardt, A. & Firth, D. (2018) Support functions and databases for Venables and Ripley’s MASS. Available from: http://www.stats.ox.ac.uk/pub/MASS4/ (accessed 20 May 2019)

    Ronquist, F., Teslenko, M., van der Mark, P., Ayres, D.L., Darling, A., Höhna, B., Larget, L., Liu, L., Suchard, M.A. & Huelsenbeck, J.P. (2012) Mr. Bayes 3.2: efficient Bayesian phylogenetic inference and model choice across a large model space. Systematic Biology, 61, 539–542.

    https://doi.org/10.1093/sysbio/sys029

    Russell, A.P., Dijkstra, L.D. & Powell, G.L. (2001) Structural characteristics of the patagium of Ptychozoon kuhli (Reptilia: Gekkonidae) in relation to parachuting locomotion. Journal of Morphology, 247, 252–263.

    https://doi.org/10.1002/1097-4687(200103)247:3<252::aid-jmor1015>3.3.co;2-q

    Sabaj, M.H. (2016) Standard symbolic codes for institutional resource collections in herpetology and ichthyology: an Online Reference. Version 6.5 (16 August 2016). American Society of Ichthyologists and Herpetologists, Washington, D.C. Electronically accessible. Available from: http://www.asih.org/ (accessed 20 May 2019)

    Shaw, K.L. (2002) Conflict between nuclear and mitochondrial DNA phylogenies of a recent species radiation: what mtDNA reveals and conceals about modes of speciation in Hawaiian crickets. Proceedings of the National Academy of Sciences of the United States, 99, 16122–16127.

    https://doi.org/10.1073/pnas.242585899

    Sheridan, J.A. & Stuart, B.L. (2018) Hidden species diversity in Sylvirana nigrovittata (Amphibia: Ranidae) highlight the importance of taxonomic revisions in biodiversity conservation. PLoS ONE, 13, e0192766.

    https://doi.org/10.1371/journal.pone.0192766

    Smith, M.A. (1935) Fauna of British India, including Ceylon and Burma. Reptilia and Amphibia. Vol. II. Sauria. Taylor & Francis Ltd., London, 440 pp.

    Stejneger, L. (1902) Ptychozoon kuhli, a new name for P. homalocephalum. Proceedings of the Biological Society of Washington, 15, 37.

    Stejneger, L. (1907) A new geckoid lizard from the Philippine Islands. Proceedings of the United States National Museum, 33, 545–546.

    https://doi.org/10.5479/si.00963801.1576.545

    Stuart, B.L. & Emmett, D.A. (2006) A collection of amphibians and reptiles from the Elephant and Cardamom Mountains, southwestern Cambodia. Fieldiana Zoology, New Series, 109, 1–27.

    https://doi.org/10.3158/0015-0754(2006)109[1:acoaar]2.0.co;2

    Stuart, B.L., Inger, R.F. & Voris, H.K. (2006) High level of cryptic species diversity revealed by sympatric lineages of Southeast Asian forest frogs. Biology Letters, 2, 470–474.

    https://doi.org/10.1098/rsbl.2006.0505

    Sukumaran, J. & Knowles, L.L. (2017) Multispecies coalescent delimits structure, not species. Proceedings of the National Academy of Sciences, 114, 1607–1612.

    https://doi.org/10.1073/pnas.1607921114

    Sumontha, M., Pauwels, O.S.G., Kunya, K., Limlikhitaksorn, C., Ruksue, S., Taokratok, A., Ansermet, M. & Chanhome, L. (2012) A new species of Parachute Gecko (Squamata: Gekkonidae: genus Ptychozoon) from Kaeng Krachan National Park, western Thailand. Zootaxa, 3513 (1), 68–78.

    https://doi.org/10.11646/zootaxa.3513.1.4

    Taylor, E.H. (1915) New species of Philippine lizards. Philippine Journal of Science, 10, 89–109.

    Taylor, E.H. (1922) Additions to the herpetological fauna of the Philippine Islands, I. Philippine Journal of Science, 21, 161–206.

    https://doi.org/10.5962/bhl.part.25389

    Taylor, E.H. (1963) The lizard fauna of Thailand. The University of Kansas Science Bulletin, 54, 687–1077.

    Teynié, A., Nguyen, T.Q., Lorvelec, O., Piquet, A., Lottier, A. & David, P. (2014) Amphibiens et reptiles du Laos: nouvelles données nationales et provincials. Bulletin de la Société Herpétologique de France, 151, 21–52.

    Thorpe, R.S. (1975) Quantitative handling of characters useful in snake systematics with particular reference to interspecific variation in the Ringed Snake Natrix natrix (L.). Biological Journal of the Linnean Society, 7, 27–43.

    https://doi.org/10.1111/j.1095-8312.1975.tb00732.x

    Thorpe, R.S. (1983) A review of the numerical methods for recognizing and analysing racial differentiation. In: Felsenstein, J. (Ed.), Numerical Taxonomy. NATO ASI Series (Series G: Ecological Sciences). Vol. 1. Springer-Verlag, Berlin, pp. 404–423.

    https://doi.org/10.1007/978-3-642-69024-2_43

    Toews, D.P.L.J. & Brelsford, A. (2012) The biogeography of nuclear discordance in animals. Molecular Ecology, 21, 3907–3930.

    https://doi.org/10.1111/j.1365-294x.2012.05664.x

    Turan, C. (1999) A note on the examination of morphometric differentiation among fish populations: the truss system. Turkish Journal of Zoology, 23, 259–263.

    Uetz, P., Freed, P. & Hošek, J. (Eds.) (2019) The Reptile Database. Available from: http://www.reptile-database.org/ (accessed 19 January 2019)

    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. Edition Chimaira, Frankfurt am Main, 324 pp.

    Wang, Y.-Y., Wang, J. & Liu, Z.-Y. (2016) Description of a new species of the genus Ptychozoon (Squamata: Gekkonidae), representing a new national record of this genus from southern Yunnan Province, China. Zootaxa, 4084 (3), 406–420.

    https://doi.org/10.11646/zootaxa.4084.3.6

    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. https://doi.org/10.1016/S1055-7903(02)00244-0

    Young, B.A., Lee, C.E. & Daley, K.M. (2002) On a flap and a foot; aerial locomotion in the “flying” gecko, Ptychozoon kuhli. Journal of Herpetology, 36, 412–418.

    https://doi.org/10.1670/0022-1511(2002)036[0412:OAFAAF]2.0.CO;2