Abstract
An integrative taxonomic analysis based on morphology, color pattern, and the mitochondrial gene ND2 recovered four new species of Hemiphyllodactylus Bleeker that are endemic to the Shan Plateau or Salween Basin in eastern Myanmar. Hemiphyllodactylus ngwelwini sp. nov. from the Shan Plateau is part of the earlier described “eastern Myanmar clade” renamed herein as the north lineage and H. kyaiktiyoensis sp. nov. and H. pinlaungensis sp. nov. of the Shan Plateau and H. zwegabinensis sp. nov. of the Salween Basin compose an entirely new Burmese clade herein referred to as the south lineage. Although the north and south lineages come within 46 km of one another on the Shan Plateau, they are not sister lineages but sequentially separated by two lineages from Yunnan, China and another from northwestern Thailand. Hemiphyllodactylus zwegabinensis sp. nov. is the first species of this genus to be recorded from the Salween Basin and is known only from a wind-blown cloud forest on the top of the insular, karstic mountain Zwegabin in Kayin State. All other Burmese species except for H. typus, are endemic to the various localities throughout the Shan Plateau. These four new species bring the total number of Hemiphyllodactylus in Myanmar to at least 10 which is certainly an extreme underestimate of the diversity of this genus given that we discover new species at every upland locality we survey.
References
Agarwal, A., El-Ghazawi, T., El-Askary, H. & Le-Moigne, J. (2007) Efficient hierarchical-PCA dimension reduction for hyperspectral imagery. 2007 IEEE International Symposium on Signal Processing and Information Technology, Giza, Egypt, 15–18 December 2007, 9863589.
https://doi.org/10.1109/ISSPIT.2007.4458191
Agarwal, I, Khandekar, A., Varad, B.G. & Ramakrishnan, U. & Karanth, K.P. (2019) The hills are alive with geckos! A radiation of a dozen species on sky islands across peninsular India (Squamata: Gekkonidae, Hemiphyllodactylus) with the description of three new species. Organisms, Diversity, & Evolution, 19, 341–361.
https://doi.org/10.1007/s13127-019-00392-5
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. (1903) Descriptions of new lizards in the collection of the British Museum. The Annals and Magazine of Natural History, Series 7, 12 (70), 429–435.
https://doi.org/10.1080/00222930308678877
Cangelosi, R. & Goriely, A. (2007) Component retention in principal component analysis with application of cDNA microarray data. Biology Direct, 2 (2), 1–21.
https://doi.org/10.1186/1745-6150-2-2
Cobos, A.L., Grismer, L.L., Wood Jr., P.L., Quah, E.S.H., Anuar, S. & Muin, M.A. (2016) Phylogenetic relationships of geckos of the Hemiphyllodactylus harterti group, a new species from Penang Island, Peninsular Malaysia, and a likely case of true cryptic speciation. Zootaxa, 4107 (3), 367–380.
https://doi.org/10.11646/zootaxa.4107.3.5
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/10.1098/rstb.1998.0210
De Queiroz, K. (2007) Species concepts and species delimitation. Systemtic 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
Eliades, S.J. Phimmachak, S., Sivongxay, N., Siler, C.D. & Stuart, B.L. (2019) Two new species of Hemiphyllodactylus (Reptilia: Gekkonidae) from Laos. Zootaxa, 4577 (1), 131–147.
https://doi.org/10.11646/zootaxa.4577.1.8
Grismer, L.L., Wood Jr., P.L., Anuar, S., Muin, M.A., Quah, E.S.H., McGuire, J.A., Brown, R.A., Ngo, V.T. & Pham, H.T. (2013) Integrative taxonomy uncovers high levels of cryptic species diversity in Hemiphyllodactylus Bleeker, 1860 (Squamata: Gekkonidae) and the description of a new species from Peninsular Malaysia. Zoological Journal of the Linnean Society, 169, 849–880.
https://doi.org/10.1111/zoj.12064
Grismer, L.L., Riyanto, A., Djoko, I.T. & McGuire, J.A. (2014a) A new species of Hemiphyllodactylus Bleeker, 1860 (Squamata: Gekkonidae) from Pulau Enggano, southwestern Sumatra, Indonesia. Zootaxa, 3821 (4), 485–495.
https://doi.org/10.11646/zootaxa.3821.4.7
Grismer, L.L., Wood Jr., PL. & Cota, M. (2014b) A new species of Hemiphyllodactylus Bleeker, 1860 (Squamata: Gekkonidae) from northwestern Thailand. Zootaxa, 3760 (1), 67–68.
https://doi.org/10.11646/zootaxa.3760.1.4
Grismer, L.L. Wood Jr., P.L., Zug, G.R., Thura, M.K., Grismer, M.S., Murdoch, M.L., Quah, E.S.H. & Lin, A. (2018) Two more new species of Hemiphyllodactylus Bleeker (Squamata: Gekkonidae) from the Shan Hills of eastern Myanmar (Burma). Zootaxa, 4483 (2), 295–316.
https://doi.org/10.11646/zootaxa.4483.2.4
Grismer, L.L., Wood Jr., P.L., Thura, M.K., Zin, T., Quah, E.S.H., Murdoch, M.L., Grismer, M.S., Li, A., Kyaw, H. & Ngwe, L. (2017b) Phylogenetic taxonomy of Hemiphyllodactylus Bleeker, 1860 (Squamata: Gekkonidae) with descriptions of three new species from Myanmar. Journal of Natural History, 52 (13–16), 881–915.
https://doi.org/10.1080/00222933.2017.1367045
Guo, W., Kaiya, Z., Jie, Y. & Peng, L. (2015) A new species of Hemiphyllodactylus Bleeker, 1860 (Squamata: Gekkonidae) from western Yunnan, China. Zootaxa, 3974 (3), 377–390.
https://doi.org/10.11646/zootaxa.3974.3.5
Heinicke, M.P., Greenbaum, E., Jackman, T.R. & Bauer, A.M. (2011) Phylogeny of a trans-Wallacean radiation (Squamata, Gekkonidae, Gehyra) supports a single early colonization of Australia. Zoologica Scripta, 40, 584–602.
https://doi.org/10.1111/j.1463-6409.2011.00495.x
Hillis, D.M. (2019) Species delimitation in herpetology. Journal of Herpetololgy, 53, 3–12.
https://doi.org/10.1670/18-123
Hoang, D.T., Chernomor, O., von Haeseler, A., Minh, B.Q. & Le, S.V. (2017) UFBoot2: Improving the Ultrafast Bootstrap Approximation. Moloecular Biology and Evolution, 35, 518–522.
https://doi.org/10.1093/molbev/msx281
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
Jombart, T., Collins, C. 2015. A tutorial for discriminant analysis of principal components (DAPC) using adegenet 2.0.0. Available from: http://adegenet.r-forge.r-project.org/files/tutorial-dapc-pdf (accessed 11 February 2020)
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
Knowles, L.L. & Carstens, B.C. (2007) Delimiting species without monophyletic gene trees. Systematic Biology, 56, 887–895.
https://doi.org/10.1080/10635150701701091
Kumar, S., Stecher, G. & Tamura, K. (2016) MEGA7: Molecular evolutionary genetics analaysis 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.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
Lleonart, J., Salat, J. & Torres, G.J. (2000) Removing allometric effects of body size in morphological analysis. Journal of Theoretical Biology, 205, 85–93.
Macey, J.J., Larson, A., Ananjeva, N.B., Fang, Z. & Papenfuss, T.J. (1997) Two
https://doi.org/10.1006/jtbi.2000.2043novel 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
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://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
Ngo, V.T., Grismer, L.L., Thai, P.H. & Wood Jr., P.L. (2014) A new species of Hemiphyllodactylus Bleeker, 1860 (Squamata: Gekkonidae) from Ba Na-Nui Nature reserve, central Vietnam, Zootaxa, 3760 (4), 539–552.
https://doi.org/10.11646/zootaxa.3760.4.3
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, T.Q., Botoc, A., Le, M.D., Nophaseud, L., Zug, G., Bonkowski, M. & Ziegler, T. (2014) A new species of Hemiphyllodactylus (Reptilia: Gekkonidae) from northern Laos. Zootaxa, 3827 (1), 45–56.
https://doi.org/10.11646/zootaxa.3827.1.4
Nguyen, T.Q., Tanja, L., Minh, D.L., Ha, T.D., Michael, B. & Ziegler, T. (2013) A new species of Hemiphyllodactylus (Reptilia: Gekkonidae) from northern Vietnam. Zootaxa, 3736 (1), 89–98.
https://doi.org/10.11646/zootaxa.3736.1.5
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 19 November 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
Sabaj, M.H. (2016) Standard symbolic codes for institutional resource collections in herpetology and ichthyology: an Online Reference. Version 6.5. Available from: http://www.asih.org/ (accessed 23 November 2019)
Siler, C.D., Oaks, 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.
https://doi.org/10.1016/j.ympev.2010.01.027
Smith, M.A. (1935) Fauna of British India, including Ceylon and Burma. Reptilia and Amphibia. Vol. II Sauria. Taylor & Francis Ltd, London, 440 pp.
https://doi.org/10.2307/1436417
Sukprasert, A., Sutthiwises S., Lauhachinda V. & Taksintum W. (2018) Two new species of Hemiphyllodactylus (Squamata: Gekkonidae) from Thailand. Zootaxa, 4369 (3), 363–376.
https://doi.org/10.11646/zootaxa.4369.3.4
Sung, Y-H., Wing-Ho, L., Ng, N.-H., Zhang, Y. & Yang, J-H (2018) A new species of Hemiphyllodactylus (Squamata: Gekkonidae) from Hong Kong. Zootaxa, 4392 (2), 361–373.
https://doi.org/10.11646/zootaxa.4392.2.8
Taylor, E.H. (1963) The lizards of Thailand. The University of Kansas Science Bulletin, 44, 687–1077.
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 analyzing 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
Trifinopoulos, J., Nguyen, L.-T., von Haeseler, A. & Minh, B.Q. (2016) W-IQ-TREE: a fast online phylogenetic tool for maximum likelihood analysis. Nucleic Acids Research, 44, W232–W235.
https://doi.org/10.1093/nar/gkw256
Turan, C. (1999) A note on the examination of morphometric differentiation among fish populations: The Truss System. Turkish Journal of Zoology, 23, 259–263.
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
Yan, J., Yubo, L., Guo, W., Peng, Li. & Kaiya, Z. (2016) A new species of Hemiphyllodactylus Bleeker, 1860 (Squamata: Gekkonidae) from Guizhou, China. Zootaxa, 4117 (4), 543–554.
https://doi.org/10.11646/zootaxa.4117.4.6
Zhou, K.-Y., Liu, Y.-Z. &Yang, G.-P. (1981) Three new subspecies of Hemiphyllodactylus yunnanensis (Boulenger) from China. Acta Zootaxonomica Sinica, 6, 202–209. [in Chinese, English translation by H. Ota (1996) in Smithsonian Herpetological Information Service, 110, 1–8, 1 pl.]
Zug, G.R. (2010) Speciation and dispersal in a low diversity taxon: the Slender geckos Hemiphyllodactylus (Reptilia, Gekkonidae). Smithsonian Contributions to Zoology, 631, 1–70.
https://doi.org/10.5479/si.00810282.631
Agarwal, A., El-Ghazawi, T., El-Askary, H. & Le-Moigne, J. (2007) Efficient hierarchical-PCA dimension reduction for hyperspectral imagery. 2007 IEEE International Symposium on Signal Processing and Information Technology, Giza, Egypt, 15–18 December 2007, 9863589.
https://doi.org/10.1109/ISSPIT.2007.4458191
Agarwal, I, Khandekar, A., Varad, B.G. & Ramakrishnan, U. & Karanth, K.P. (2019) The hills are alive with geckos! A radiation of a dozen species on sky islands across peninsular India (Squamata: Gekkonidae, Hemiphyllodactylus) with the description of three new species. Organisms, Diversity, & Evolution, 19, 341–361.
https://doi.org/10.1007/s13127-019-00392-5
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. (1903) Descriptions of new lizards in the collection of the British Museum. The Annals and Magazine of Natural History, Series 7, 12 (70), 429–435.
https://doi.org/10.1080/00222930308678877
Cangelosi, R. & Goriely, A. (2007) Component retention in principal component analysis with application of cDNA microarray data. Biology Direct, 2 (2), 1–21.
https://doi.org/10.1186/1745-6150-2-2
Cobos, A.L., Grismer, L.L., Wood Jr., P.L., Quah, E.S.H., Anuar, S. & Muin, M.A. (2016) Phylogenetic relationships of geckos of the Hemiphyllodactylus harterti group, a new species from Penang Island, Peninsular Malaysia, and a likely case of true cryptic speciation. Zootaxa, 4107 (3), 367–380.
https://doi.org/10.11646/zootaxa.4107.3.5
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/10.1098/rstb.1998.0210
De Queiroz, K. (2007) Species concepts and species delimitation. Systemtic 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
Eliades, S.J. Phimmachak, S., Sivongxay, N., Siler, C.D. & Stuart, B.L. (2019) Two new species of Hemiphyllodactylus (Reptilia: Gekkonidae) from Laos. Zootaxa, 4577 (1), 131–147.
https://doi.org/10.11646/zootaxa.4577.1.8
Grismer, L.L., Wood Jr., P.L., Anuar, S., Muin, M.A., Quah, E.S.H., McGuire, J.A., Brown, R.A., Ngo, V.T. & Pham, H.T. (2013) Integrative taxonomy uncovers high levels of cryptic species diversity in Hemiphyllodactylus Bleeker, 1860 (Squamata: Gekkonidae) and the description of a new species from Peninsular Malaysia. Zoological Journal of the Linnean Society, 169, 849–880.
https://doi.org/10.1111/zoj.12064
Grismer, L.L., Riyanto, A., Djoko, I.T. & McGuire, J.A. (2014a) A new species of Hemiphyllodactylus Bleeker, 1860 (Squamata: Gekkonidae) from Pulau Enggano, southwestern Sumatra, Indonesia. Zootaxa, 3821 (4), 485–495.
https://doi.org/10.11646/zootaxa.3821.4.7
Grismer, L.L., Wood Jr., PL. & Cota, M. (2014b) A new species of Hemiphyllodactylus Bleeker, 1860 (Squamata: Gekkonidae) from northwestern Thailand. Zootaxa, 3760 (1), 67–68.
https://doi.org/10.11646/zootaxa.3760.1.4
Grismer, L.L. Wood Jr., P.L., Zug, G.R., Thura, M.K., Grismer, M.S., Murdoch, M.L., Quah, E.S.H. & Lin, A. (2018) Two more new species of Hemiphyllodactylus Bleeker (Squamata: Gekkonidae) from the Shan Hills of eastern Myanmar (Burma). Zootaxa, 4483 (2), 295–316.
https://doi.org/10.11646/zootaxa.4483.2.4
Grismer, L.L., Wood Jr., P.L., Thura, M.K., Zin, T., Quah, E.S.H., Murdoch, M.L., Grismer, M.S., Li, A., Kyaw, H. & Ngwe, L. (2017b) Phylogenetic taxonomy of Hemiphyllodactylus Bleeker, 1860 (Squamata: Gekkonidae) with descriptions of three new species from Myanmar. Journal of Natural History, 52 (13–16), 881–915.
https://doi.org/10.1080/00222933.2017.1367045
Guo, W., Kaiya, Z., Jie, Y. & Peng, L. (2015) A new species of Hemiphyllodactylus Bleeker, 1860 (Squamata: Gekkonidae) from western Yunnan, China. Zootaxa, 3974 (3), 377–390.
https://doi.org/10.11646/zootaxa.3974.3.5
Heinicke, M.P., Greenbaum, E., Jackman, T.R. & Bauer, A.M. (2011) Phylogeny of a trans-Wallacean radiation (Squamata, Gekkonidae, Gehyra) supports a single early colonization of Australia. Zoologica Scripta, 40, 584–602.
https://doi.org/10.1111/j.1463-6409.2011.00495.x
Hillis, D.M. (2019) Species delimitation in herpetology. Journal of Herpetololgy, 53, 3–12.
https://doi.org/10.1670/18-123
Hoang, D.T., Chernomor, O., von Haeseler, A., Minh, B.Q. & Le, S.V. (2017) UFBoot2: Improving the Ultrafast Bootstrap Approximation. Moloecular Biology and Evolution, 35, 518–522.
https://doi.org/10.1093/molbev/msx281
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
Jombart, T., Collins, C. 2015. A tutorial for discriminant analysis of principal components (DAPC) using adegenet 2.0.0. Available from: http://adegenet.r-forge.r-project.org/files/tutorial-dapc-pdf (accessed 11 February 2020)
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
Knowles, L.L. & Carstens, B.C. (2007) Delimiting species without monophyletic gene trees. Systematic Biology, 56, 887–895.
https://doi.org/10.1080/10635150701701091
Kumar, S., Stecher, G. & Tamura, K. (2016) MEGA7: Molecular evolutionary genetics analaysis 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.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
Lleonart, J., Salat, J. & Torres, G.J. (2000) Removing allometric effects of body size in morphological analysis. Journal of Theoretical Biology, 205, 85–93.
Macey, J.J., Larson, A., Ananjeva, N.B., Fang, Z. & Papenfuss, T.J. (1997) Two
https://doi.org/10.1006/jtbi.2000.2043novel 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
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://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
Ngo, V.T., Grismer, L.L., Thai, P.H. & Wood Jr., P.L. (2014) A new species of Hemiphyllodactylus Bleeker, 1860 (Squamata: Gekkonidae) from Ba Na-Nui Nature reserve, central Vietnam, Zootaxa, 3760 (4), 539–552.
https://doi.org/10.11646/zootaxa.3760.4.3
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, T.Q., Botoc, A., Le, M.D., Nophaseud, L., Zug, G., Bonkowski, M. & Ziegler, T. (2014) A new species of Hemiphyllodactylus (Reptilia: Gekkonidae) from northern Laos. Zootaxa, 3827 (1), 45–56.
https://doi.org/10.11646/zootaxa.3827.1.4
Nguyen, T.Q., Tanja, L., Minh, D.L., Ha, T.D., Michael, B. & Ziegler, T. (2013) A new species of Hemiphyllodactylus (Reptilia: Gekkonidae) from northern Vietnam. Zootaxa, 3736 (1), 89–98.
https://doi.org/10.11646/zootaxa.3736.1.5
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 19 November 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
Sabaj, M.H. (2016) Standard symbolic codes for institutional resource collections in herpetology and ichthyology: an Online Reference. Version 6.5. Available from: http://www.asih.org/ (accessed 23 November 2019)
Siler, C.D., Oaks, 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.
https://doi.org/10.1016/j.ympev.2010.01.027
Smith, M.A. (1935) Fauna of British India, including Ceylon and Burma. Reptilia and Amphibia. Vol. II Sauria. Taylor & Francis Ltd, London, 440 pp.
https://doi.org/10.2307/1436417
Sukprasert, A., Sutthiwises S., Lauhachinda V. & Taksintum W. (2018) Two new species of Hemiphyllodactylus (Squamata: Gekkonidae) from Thailand. Zootaxa, 4369 (3), 363–376.
https://doi.org/10.11646/zootaxa.4369.3.4
Sung, Y-H., Wing-Ho, L., Ng, N.-H., Zhang, Y. & Yang, J-H (2018) A new species of Hemiphyllodactylus (Squamata: Gekkonidae) from Hong Kong. Zootaxa, 4392 (2), 361–373.
https://doi.org/10.11646/zootaxa.4392.2.8
Taylor, E.H. (1963) The lizards of Thailand. The University of Kansas Science Bulletin, 44, 687–1077.
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 analyzing 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
Trifinopoulos, J., Nguyen, L.-T., von Haeseler, A. & Minh, B.Q. (2016) W-IQ-TREE: a fast online phylogenetic tool for maximum likelihood analysis. Nucleic Acids Research, 44, W232–W235.
https://doi.org/10.1093/nar/gkw256
Turan, C. (1999) A note on the examination of morphometric differentiation among fish populations: The Truss System. Turkish Journal of Zoology, 23, 259–263.
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
Yan, J., Yubo, L., Guo, W., Peng, Li. & Kaiya, Z. (2016) A new species of Hemiphyllodactylus Bleeker, 1860 (Squamata: Gekkonidae) from Guizhou, China. Zootaxa, 4117 (4), 543–554.
https://doi.org/10.11646/zootaxa.4117.4.6
Zhou, K.-Y., Liu, Y.-Z. &Yang, G.-P. (1981) Three new subspecies of Hemiphyllodactylus yunnanensis (Boulenger) from China. Acta Zootaxonomica Sinica, 6, 202–209. [in Chinese, English translation by H. Ota (1996) in Smithsonian Herpetological Information Service, 110, 1–8, 1 pl.]
Zug, G.R. (2010) Speciation and dispersal in a low diversity taxon: the Slender geckos Hemiphyllodactylus (Reptilia, Gekkonidae). Smithsonian Contributions to Zoology, 631, 1–70.
https://doi.org/10.5479/si.00810282.631