Skip to main content Skip to main navigation menu Skip to site footer
Type: Article
Published: 2019-10-09
Page range: 381–411
Abstract views: 120
PDF downloaded: 5

Integrative taxonomy of the Asian skinks Sphenomorphus stellatus (Boulenger, 1900) and S. praesignis (Boulenger, 1900) with the resurrection of S. annamiticus (Boettger, 1901) and the description of a new species from Cambodia

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 Institute of Tropical Biodiversity and Sustainable Development, Universiti Malaysia Terengganu, 21030 Kuala Terengganu, Terengganu, Malaysia
Institute of Tropical Biodiversity and Sustainable Development, Universiti Malaysia Terengganu, 21030 Kuala Terengganu, Terengganu, Malaysia
Faculty of Biology, Department of Vertebrate Zoology, Moscow State University, Moscow, Russia. Laboratory of Tropical Ecology, Joint Russian-Vietnamese Tropical Research and Technological Center, Hanoi, Vietnam
Wild Earth Allies, 77a, St. Betong, Bayap Village, Sangkat Phom Penh Thmei, Khan Sen Sok, Phnom Penh, Cambodia.
Zoological Institute, Russian Academy of Sciences, Universitietskaya nab., 1, St. Petersburg 199034, Russia.
Department of Biology, Faculty of Science, Chulalongkorn University, 254 Phayathai Road, Pathumwan, Bangkok, Thailand 10330
Wild Earth Allies, 77a, St. Betong, Bayap Village, Sangkat Phom Penh Thmei, Khan Sen Sok, Phnom Penh, Cambodia.
Reptilia Phylogenetic systematics Indochina Peninsular Malaysia Cambodia Vietnam Scincidae Prey Lang Wildlife Sanctuary

Abstract

Molecular phylogenetic analyses of the sister species Sphenomorphus stellatus and S. praesignis based on the mitochondrial genes 12S and 16S rRNA recover the former as paraphyletic with respect to the latter in that a specimen of S. stellatus from the type locality in Peninsular Malaysia is more closely related to S. praesignis than to Indochinese populations of S. stellatus. Furthermore, the phylogeny indicates that the Indochinese populations represent two species, thus resulting in four major lineages within this clade. These relationships are consistent with multivariate and univariate analyses of morphological and discrete color pattern data which statistically define and diagnose the four lineages and together with the molecular data, provide the foundation for robust, testable, species-level hypotheses. As such, S. stellatus is herein restricted to Peninsular Malaysia; S. annamiticus is resurrected for the circum-continental populations ranging through southeastern Thailand, southern Cambodia, and southern Vietnam; a new species—S. preylangensis sp. nov.—is described from an isolated mountain, Phnom Chi, from the Prey Lang Wildlife Sanctuary in central Cambodia; and the taxonomy of S. praesignis remains unchanged. The description of S. preylangensis sp. nov. underscores the necessity to conserve this remnant of lowland evergreen rainforest in the Prey Lang Wildlife Sanctuary.

 

References

  1. Aljanabi, S.M. & Martinez, I. (1997) Universal and rapid salt-extraction of high quality genomic DNA for PCR-based techniques. Nucleic Acids Research, 25, 4692–4693.

    https://doi.org/10.1093/nar/25.22.4692

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

    https://doi.org/10.1206/360.1

    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

    Boettger, O. (1901) Aufzählung einer Liste von Reptilien und Batrachiern aus Annam. Bericht der Senckenbergischen Naturforschenden Gesellschaft in Frankfurt am Main, 1901, 45­–53

    Boulenger, G.A. (1900) Descriptions of new batrachians and reptiles from Larut Hills, Perak. Annals and Magazine of Natural History, Series 7, 6, 186–193.

    https://doi.org/10.1080/00222930008678356

    Chan-ard, T, Parr, J.W.K. & Nabhitabhata, J. (2015) A Field Guide to the Reptiles of Thailand. Oxford University Press, New York, 214 pp.

    https://doi.org/10.1108/rr-03-2017-0074

    Chen, J.-M., Poyarkov, N.A., Jr., Suwannapoom, C., Lathrop, A., Wu, Y-H., Zhou, W-W., Yuan, Z-Y., Jin, J-Q., Chen, H-M., Liu, H.-Q., Nguyen, T.Q., Nguyen, S.N., Duong, T.V., Eto, K., Nishikawa, K., Matsui, M., Orlov, N.L., Stuart, B.L., Brown, R.M., Rowley, J.J.L., Murphy, R.W., Wang, Y.-Y. & Che, J. (2018) Large-scale phylogenetic analyses provide insights into unrecognized diversity and historical biogeography of Asian leaf-litter frogs, genus Leptolalax (Anura: Megophryidae). Molecular Phylogenetics and Evolution, 124, 162–171.

    https://doi.org/10.1016/j.ympev.2018.02.020

    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

    Darevsky, I.S. & Nguyen, V.S. (1983) New and little known lizard species from Vietnam. Zoologichesky Zhurnal, 62, 1827–1837. [in Russian] de Queiroz, K. (2007) Species concepts and species delimitation. Systematic Biology, 56, 879–886.

    https://doi.org/10.1080/10635150701701083

    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

    Freitas, E.S., Datta-Roy, A., Karanth, P., Grismer, L.L. & Siler, C.D. (2019) Multilocus phylogeny and a new classification for African, Asian and Indian supple and writhing skinks (Scincidae: Lygosominae). Zoological Journal of the Linnean Society, 186, 1067–1096.

    https://doi.org/10.1093/zoolinnean/zlz001

    Geissler, P., Poyarkov, N.A. Jr., Grismer, L.L., Nguyen, T.Q., An, H.T., Neang, T., Kupfer, A., Ziegler, T., Böhme, W. & Müller, H. (2015a) New Ichthyophis species from Indochina (Gymnophiona, Ichthyophiidae): 1. The unstriped forms with descriptions of three new species and the redescriptions of I. acuminatus Taylor, 1960, I. youngorum Taylor, 1960 and I. laosensis Taylor, 1969. Organisms Diversity & Evolution, 15, 143–174.

    https://doi.org/10.1007/s13127-014-0190-6.

    Geissler, P., Hartmann, T., Ihlow, F., Rödder, D., Poyarkov, N.A. Jr., Nguyen, T.Q., Ziegler, T. & Böhme, W. (2015b) The Lower Mekong: an insurmountable barrier to amphibians in southern Indochina? Biological Journal of the Linnean Society, 114, 905–914.

    https://doi.org/10.1111/bij.12444

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

    Grismer, L.L., Dzukafly, Z., Muin, M.A., Quah, E.S.H., Karin, B.R., Shahrul, A. & Freitas, E.S. (2019a) A new skink of the genus Subdoluseps (Hardwicke & Gray, 1828) from Peninsular Malaysia. Zootaxa, 4609 (2), 358–372.

    https://doi.org/10.11646/zootaxa.4609.2.10

    Grismer, L.L. & Grismer, J.L. (2007) A re-evaluation of the phylogenetic relationships of the Cyrtodactylus condorensis group (Squamata; Gekkonidae) and a suggestd protocol for the characterization of rock-dwelling ecomorphology in Cyrtodactylus. Zootaxa, 4300 (4), 486–504.

    https://doi.org/10.11646/zootaxa.4300.4.2

    Grismer, L.L. & Ngo, V.T. (2007) Four new species of the gekkonid genus Cnemaspis Strauch 1887 (Reptilia: Squamata) from southern Vietnam. Herpetologica, 63, 482–500.

    https://doi.org/10.1655/0018-0831(2007)63[482:FNSOTG]2.0.CO;2

    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. & Quah, E.S.H. (2015) The rediscovery of Sphenomorphus malayanus Doria, 1888 (Squamata: Scincidae) from the Titiwangsa Mountain Range of Peninsular Malaysia and its re-description as S. senja sp. nov. Zootaxa, 3931 (1), 63–70.

    https://doi.org/10.11646/zootaxa.3931.1.4

    Grismer, L.L., Quah, E.S.H., Duzulkafly, Z. & Yambun, P. (2018) On the taxonomy of Lygosoma bampfyldei Bartlett, 1895 (Squamata: Scincidae) with descriptions of new species from Borneo and Peninsular Malaysia and the resurrection of Lygosoma schneideri Werner, 1900. Zootaxa, 4438 (3), 528–550.

    https://doi.org/10.11646/zootaxa.4438.3.6

    Grismer, L.L., Wood Jr., P.L., Grismer, J.L., Quah, E.S.H., Neang, T., Phimmachak, S., Sivongxay, N., Seateun, S., Stuart, B.L., Siler, C.B., Mulcahy, D.G., Anamza, T. & Brown, R.M. (2019b) Geographic structure of genetic variation in the Parachute Gecko Ptychozoon lionotum Annandale, 1905 across Indochina and Sundaland with descriptions of three new species. Zootaxa. [in press]

    https://doi.org/10.11646/zootaxa.4638.2.1

    Grismer, L.L., Wood, Jr., P.L., Quah, E.S.H., Anuar, S., Ngadi, E. B., Nur, A.M. I. & Norhayati, A. (2017) Systematics, ecomorphology, cryptic speciation and biogeography of the lizard genus Tytthoscincus Linkem, Diesmos & Brown (Squamata: Scincidae) from the sky-island archipelago of Peninsular Malaysia. Zoological Journal of the Linnean Society, 183, 635–671.

    https://doi.org/10.1093/zoolinnean/zlx067

    Hartmann, T., Geissler, P., Poyarkov, N.A. Jr., Ihlow, F., Galoyan, E.A., Rödder, D. & Böhme, W. (2013) A new species of the genus Calotes Cuvier, 1817 (Squamata: Agamidae) from southern Vietnam. Zootaxa, 3599 (3), 246–260.

    https://doi.org/10.11646/zootaxa.3599.3.3

    Hartwell, H., Welsh, J.R. & Oliver, L.M. (1998) Stream Amphibians as Indicatorsof Ecosystem Stress: A Case Study from California Redwoods. Ecological Applications, 8, 1118–1132.

    https://doi.org/10.1890/1051-0761(1998)008[1118:saaioe]2.0.co;2

    Hayes, B., Khou, E.H., Neang, T., Furey, N., Sophea, C., Holden, J., Seiha, H., Sarith, P. Pengly, L. & Simpson, V. (2015) Biodiversity assessment of Prey Lang Kratie, Kampong Thom, Stung Treng and Preah Vihear Provinces. Unpublished environmental impact assessment. Conservation International, Greater Mekong and Winrock International USAID Supporting Forests and Biodiversity Project, Phnom Penh,125 pp.

    Hedges, S.B., Nussbaum, R.A. & Maxson, L.R. (1993). Caecilian phylogeny and biogeography inferred from mitochondrial DNA sequences of the 12S rRNA and 16S rRNA genes (Amphibia: Gymnophiona). Herpetological Monographs, 7, 64–76.

    https://doi.org/10.2307/1466952

    Hillis, D.M. (2019) Species delimitation in herpetology. Journal of Herpetology, 53, 3–12.

    https://doi.org/10.1670/18-123

    Hoang, D.T., Chernomor, O., von Haeseler, A., Minh, B.Q. & Vinh, L.S. (2018) Ufboot2: Improving the ultrafast bootstrap approximation. Molecular Biology and Evolution, 35 (2), 518–522.

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

    Honda, M., Ota, H., Murphy, R.W. & Hikida, T. (2006) Phylogeny and biogeography of the water skinks of the genus Tropidophorus (Reptilia: Scincidae): a molecular approach. Zoological Scripta, 35, 85–95.

    https://doi.org/10.1111/j.1463-6409.2005.00215.x

    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

    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

    Kalyaanamoorthy, S., Minh, B.Q., Wong, T.K., von Haeseler, A. & Jermiin, L.S. (2017) ModelFinder: fast model selection for accurate phylogenetic estimates. Nature Methods, 14, 587.

    https://doi.org/10.1038/nmeth.4285

    Karin, B.R., Freitas, E.S., Shonleben, S., Grismer, S.S., Bauer, A.M. & Das, I. (2018) Unrealized diversity in an urban rainforest: a new species of Lygosoma (Squamata: Scincidae) from western Sarawak, Malaysia (Borneo). Zootaxa, 4370 (4), 345–362.

    https://doi.org/10.11646/zootaxa.4370.4.2

    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

    Kearse, M., Moir, R., Wilson, A., Stones-Havas, S., Cheung, M., Sturrock, S., Buxton, S., Cooper, A., Markowitz, S., Duran, C., Thierer, T., Ashton, B., Meintjes, P. & Drummond, A. (2012) Geneious Basic: an integrated and extendable desktop software platform for the organization and analysis of sequence data. Bioinformatics, 28, 1647–1649.

    https://doi.org/10.1093/bioinformatics/bts199

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

    https://doi.org/10.1080/10635150701701091

    Kocher, T.D., Thomas, W.K., Meyer, A., Edwards, S.V., Pääbo, S., Villablanca, F.X. & Wilson, A.C. (1989) Dynamics of mitochondrial DNA evolution in animals: Amplification and sequencing of conserved primers. Proceedings of the National Academy of Sciences, 86, 6196–6200.

    https://doi.org/10.1073/pnas.86.16.6196

    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

    Leaché, A.D., Koo, M.S., Spencer, C.S., 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

    Linkem, C.W. (2013) Molecular Phylogenetics and Biogeography of Sphenomorphini (Squamata: Scincidae). Unpublished PhD Thesis, Department of Ecology and Evolutionary Biology, University of Kansas, Lawrence, Kansas, 256 pp.

    Lim, B.L. (1967) Further comments on rare snakes. Federated Museum Journal of Malaya, 12, 123–126.

    https://doi.org/10.1177/003693306701200308

    Maddison, W.P. & Maddison, D.R. (2015) Mesquite: a modular system for evolutionary analysis. Version 3.04. Available from: http://mesquiteproject.org (accessed 18 September 2019)

    https://doi.org/10.1093/sysbio/42.2.218

    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

    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, Louisiana, 14 November 2010, pp. 1–8. https//dx.doi.org/10.1109/GCE.2010.5676129

    Minh, Q., Nguyen, M.A.T. & von Haeseler, A. (2013) Ultrafast approximation for phylogenetic bootstrap. Molecular Biology and Evolution, 30, 1188–1195. https://doi.org/10.1093/molbev/mst024

    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

    Myers, N., Mittermeier, R.A. & Mittermeier, C.G. (2000) Biodiversity hotspots for conservation priorities. Nature, 403, 853–858.

    https://doi.org/10.1038/35002501

    Neang, T., Chan, S. & Poyarkov, N.A. Jr. (2018) A new species of smooth skink (Squamata: Scincidae: Scincella) from Cambodia. Zoological Research, 38 (3), 214–233.

    https://doi.org/10.24272/j.issn.2095-8137.2018.008

    Nguyen, H.N., Lu, C.-W., Chu, J.-H, Grismer, L.L., Hung, C.-M. & Lin, S.-M. (2018) Historical demography of four gecko species specializing in boulder cave habitat: Implications in the evolutionary dead end hypothesis and conservation. Molecular Ecology, 28, 772–784.

    https://doi.org/10.1111/mec.14985

    Nguyen, T.Q., Schmitz, A., Nguyen, T.T., Orlov, N.L., Böhme, W. & Ziegler (2011) Review of the genus Sphenomorphus Fitzinger, 1843 (Squamata; Sauria: Scincidae) in Vietnam, with description of a new species from northern Vietnam and southern China and the first record of Sphenomorphus mimicus Taylor, 1962 from Vietnam. Journal of Herpetology, 45, 145–154.

    https://doi.org/10.1670/09-068.1

    Nguyen, S.-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.

    Norsham, Y., Norhayati, A., Shariff, F., Nordin, M. & Lim, B.L. (2005) Pre-logging survey on vertebrate species diversity at Sungai Weng sub-catchment, Ulu Muda Forest Reserve, Kedah. 2: Reptilian fauna. Malayan Nature Journal, 57, 145–154.

    Poyarkov, N.A., Jr., Duong, T.V., Orlov, N.L., Gogoleva, S.S., Vassilieva, A.B., Nguyen, L.T., Nguyen, V.H.D., Nguyen, S.N., Che, J. & Mahony S. (2017) Molecular, morphological and acoustic assessment of the genus Ophryophryne (Anura, Megophryidae) from Langbian Plateau, southern Vietnam, with description of a new species. ZooKeys, 672, 49–120.

    https://doi.org/10.3897/zookeys.672.10624

    Poyarkov, N.A., Jr., Geissler, P., Gorin, V.A., Dunayev, E.A., Hartmann, T. & Suwannapoom, C. (2019) Counting stripes: revision of the Lipinia vittigera complex (Reptilia, Squamata, Scincidae) with description of two new species from Indochina. Zoological Research, 40, 358–393.

    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 1 June 2016)

    Rambaut, A., Suchard, M.A., Xie, D. & Drummond, A.J. (2014) Tracer. Version 1.6. Available from: http://beast.bio.ed.ac.uk/Tracer/ (accessed 18 September 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 18 September 2019)

    Ronquist, F., Teslenko, M., van der Mark, P., Ayres, D.S., Darling, A., Höhna, B., Larget, S., Liu, S., 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

    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

    Siler, C.D., Heitz, B.B., Davis, D.R., Freitas, E.S., Aowphol, A., Termprayoon, K. & Grismer, S.S. (2018) New Supple Skink, Genus Lygosoma (Reptilia: Squamata: Scincidae), from Indochina and redescription of Lygosoma quadrupes (Linnaeus, 1766). Journal of Herpetology, 52, 332–347.

    https://doi.org/10.1670/16-064

    Smith, M.A. (1935) The fauna of British India, including Ceylon and Burma. Reptiles and Amphibia. Vol. II. Sauria. Taylor and Francis, London, 440 pp.

    Stuart, B.L. & Emmett, A.E. (2006) A collection of amphibians and reptiles from the Cardamom Mountains, southwestern Cambodia. Fieldiana Zoology, 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

    Sumarli, A., Grismer, L.L., Wood, P.L., Jr., Ahmad, A.B., Rizal, S., Ismail, L.H., Izam, N.A.M., Ahmad, M. & Linkem, C.W. (2016) The first riparian skink (Genus: Sphenomorphus Strauch, 1887) from Peninsular Malaysia and its relationship to other Indochinese and Sundaic species Zootaxa, 4173 (1), 029–044.

    https://doi.org/10.11646/zootaxa.4173.1.3

    Taylor, E.H. (1935) A taxonomic study of the cosmopolitan lizards of the genus Eumeces with an account of the distribution and relationships of its species. The University of Kansas Science Bulletin, 23, 1–643.

    https://doi.org/10.2307/1436337

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

    Uetz, P., Freed, P. & Hošek, J. (Eds.) (2019) The Reptile Database. Available from: http://www.reptile-database.org, (accessed 27 June 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, 319 pp.

    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