Abstract
An integrative taxonomic analysis of the 10 species of the Cyrtodactylus sinyineensis group based on squamation, color pattern, and the mitochondrial gene NADH dehydrogenase subunit 2 (ND2) and its flanking tRNA regions, recovered the newly discovered populations from Datt Kyaik and Taung Wine Hills in Kayin State as the new species Cyrtodactylus dattkyaikensis sp. nov. and C. taungwineensis sp. nov. The Maximum Likelihood and Bayesian phylogenetic estimates supported C. dattkyaikensis sp. nov. as the sister species of C. bayinnyiensis and C. taungwineensis sp. nov. as the sister species of C. sinyineensis. Each new species is differentially diagnosable from all other C. sinyineensis group species based on their morphological placement in multivariate space and several statistically significant mean differences is meristic squamation and color pattern data. The C. sinyineensis group ranges across an archipelago of karstic habitat-islands in the Salween Basin of southern Myanmar. The discovery of these new species continues to underscore the unprecedented high degree of diversity and site-specific endemism in this relatively small region and the urgent need for the conservation of its karstic terranes.
References
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
Barraclough, T.G., Birky, C.W. Jr. & Burt, A. (2003) Diversification in sexual and asexual organisms. Evolution, 57, 2166–2172.
https://doi.org/10.1111/j.0014-3820.2003.tb00394.xBauer, A.M. (2003) Descriptions of seven new Cyrtodactylus (Squamata: Gekkonidae) with a key to the species of Myanmar (Burma). Proceedings of the California Academy of Sciences, 54, 463–498.
Connette, G.M., Oswald, P., Thura, M.K., Connette, K.J.L., Grindley, M.E., Songer, M., Zug, G.R., Mulchay, D.G. (2017) Rapid forest clearing in a Myanmar proposed national park threatens two newly discovered species of geckos (Gekkonidae: Cyrtodactylus). PLoS One, 12, e0174432.
https://doi.org/10.1371/journal.pone.0174432
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
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. & Barrenclough, T.G. (2007) Independently evolving species in asexual bdelloid rotifers. PLoS Biology, 5, e87.
https://doi.org/10.1371/journal.pbio.0050087
Grismer, L.L. (2017a) Herpetofaunal Survey of the Pyinyaung Limestone Mining Area. Unpublished survey report for Fauna & Flora International, Yangon, 12 pp.
Grismer, L.L. (2017b) Herpetofaunal Survey of Limestone Habitats in the Salween Basin. Unpublished survey report for Fauna & Flora International, Yangon, 17 pp.
Grismer, L.L. (2018a) Herpetofaunal Survey of Karstic Habitats in the Salween Basin. Unpublished survey report for Fauna & Flora International, Yangon, 13 pp.
Grismer, L.L. (2018b) Gecko Surveys of the Panlaung-Pyadalin Cave Wildlife Sanctuary and Adjacent Areas in Shan State and Mandalay Division. Unpublished survey report for Fauna & Flora International, Yangon, 12 pp.
Grismer, L.L., Wood Jr., P.L., Anuar, S., Davis, H.R, Cobos, A.J. & Murdoch, M.L. (2016) A new species of karst forest Bent-toed Gecko (genus Cyrtodactylus Gray) not yet threatened by foreign cement companies and a summary of Peninsular Malaysia’s endemic karst forest herpetofauna and the need for its conservation. Zootaxa, 4061 (1), 1–17.
https://doi.org/10.11646/zootaxa.4061.1.1
Grismer, L.L., Wood Jr., P.L., Chan, K.O., Anuar & Muin, M.A. (2014) Cyrts in the city: A new Bent-toed Gecko (Genus Cyrtodactylus) is the only endemic species of vertebrate from Batu Caves, Selangor, Peninsular Malaysia. Zootaxa, 3774 (4), 381–394.
https://doi.org/10.11646/zootaxa.3774.4.6
Grismer, L.L., Wood Jr.,P.L., Thura, M.K., Win, N.M. & Quah, E.S.H. (2018g)Two more new species of the Cyrtodactylus peguensis group (Squamata: Gekkonidae) from the fringes of the Ayeyarwady Basin, Myanmar. Zootaxa, 4577 (2), 274–294.
https://doi.org/10.11646/zootaxa.4577.2.3
Grismer, L.L., Wood Jr., P.L., Thura, M.K., Quah, E.S.H., Grismer, M.S., Murdoch, M.L., Espinoza, R.E. & Lin, A. (2018c) A new Cyrtodactylus Gray (Squamata, Gekkonidae) from the Shan Hills and the biogeography of Bent-toed Geckos from eastern Myanmar. Zootaxa, 4446 (4), 477–500.
https://doi.org/10.11646/zootaxa.4446.4.4
Grismer, L.L., Wood. Jr., P.L., Thura, M.K., Zin, T., Quah, E.S.H., Murdoch, M.L., Grismer, M.S., Lin, A., Kyaw, H. & Ngwe, L. (2018a) 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.
http://doi.org/10.1093/zoolinnean/zlx057
Grismer, L.L., Wood Jr., P.L., Thura, M.K., Win, N.M., Grismer, M.S., Trueblood, L.A. & Quah, E.H.S. (2018c) A re-description of Cyrtodactylus chrysopylos Bauer (Squamata: Gekkonidae) with comments on the adaptive significance of orange coloration in hatchlings and descriptions of two new species from eastern Myanmar (Burma). Zootaxa, 4527 (2), 151–185.
https://doi.org/10.11646/zootaxa.4527.2.1
Grismer, L.L., Wood Jr., P.L., Thura, M.K., Zin, T., Quah, E.S.H., Murdoch, M.L., Grismer, M.S., Herr, M.W., Lin, A. & Kyaw, H. (2018b) Three more new species of Cyrtodactylus (Squamata: Gekkonidae) from the Salween Basin of eastern Myanmar underscores the urgent need for the conservation of karst habitats. Journal of Natural History, 52, 1243–1294.
http://doi.org/10.1080/00222933.2018.1449911
Grismer, L.L., Wood Jr., P.L., Quah, E.S.H., Thura, M.K., Herr, M.X. & Lin, A.K. (2019b) A new species of forest-dwelling Cyrtdactylus (Squamata: Gekkonidae) from Indawgyi Wildlife Sanctuary, Kachin State, Myanmar. Zootaxa, 4623 (1), 1–25.
https://doi.org/10.11646/zootaxa.4623.1.1
Grismer, L.L., Wood Jr., P.L., Quah, E.S.H., Thura, M.K., Murdoch, M.L., Grismer, M.S., Herr, M.W., Espinoza, R.E., Brown, R.M. & Lin, A. (2018e)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., Thura, M.K., Zin, T., Quah, E.S.H., Murdoch, M.L., Grismer, M.S., Lin, A., Kyaw, H. & Ngwe, L. (2017) Phylogenetic taxonomy of Hemiphyllodactylus Bleeker, 1860 (Squamata: Gekkonidae) with descriptions of three new species from Myanmar. Journal of Natural History, 52, 881–195.
https://doi.org/10.1080/00222933.2017.1367045
Grismer, L.L., Wood Jr., P.L., Quah, E.S.H., Thura, M.K., Oaks, J.R. & Lin, A. (2019a) A new species of Bent-toed Gecko (Squamata, Gekkonidae, Cyrtodactylus) from the Shan Plateau in eastern Myanmar (Burma). Zootaxa, 4624 (3), 301–321.
https://doi.org/10.11646/zootaxa.4624.3.1
Grismer, L.L., Wood Jr., P.L., Zug, G.R., Thura, M.K., Grismer, M.S., Quah, E.S.H., Murdoch, M.L. & Lin, A. (2018d) Two more new species of Hemiphyllodaxctylus 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
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, 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
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
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
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 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.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
Macey, J.R., 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 (1), 91–104.
https://doi.org/10.1006/mpev.1997.0478
Maddison, W.P. & Maddison, D.R. (2015) Mesquite: a modular system for evolutionary analysis. Version 3.04. Available from: http://mesquiteproject.org (accessed 01 August 2018)
Miller, M.A., Pfeiffer, W. & Schwartz, T. (2010) Creating the CIPRES Science Gateway for inference of large phylogenetic trees. 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
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
R Core Team (2018) R: A language and environment for statistical computing. R Foundation for Statistical Computing. Vienna. Available from: http://www.R-project.org (accessed 1 August 2018)
Rambaut, A., Suchard, M.A., Xie, D. & Drummond, A.J. (2014) Tracer. Version 1.6. Available from: http://tree.bio.ed.ac.uk/software/tracer/ (accessec 13 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 (16 August 2016). American Society of Ichthyologists and Herpetologists, Washington, D.C. Available from: http://www.asih.org/ (accessed 01 October 2019)
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
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.
http://doi.org/10.1016/S1055-7903(02)00244-0
Wood Jr., P.L., 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.
https://doi.org/10.1016/j.ympev.2012.08.025