Abstract
The Phong Nha-Ke Bang (PNKB National Park in the Central Highlands of Vietnam is a bastion for the protection and conservation of Vietnam’s natural heritage. Thus, the discovery of yet another new species of Cyrtodactylus, C. hangvaensis sp. nov., within the PNKB continues to underscore the parks ever-growing importance. Cyrtodactylus hangvaensis sp. nov. is a karst dwelling species of the angularis group that based on a molecular phylogenetic analysis, is most closely related to C. roesleri from the PNKB and C. sommerladi from karstic areas in nearby Laos. Cyrtodactylus hangvaensis sp. nov. occupies a unique position in morphospace where it is well separated from closely related species and other species within the PNKB. It is potentially diagnosable from all other members of the angularis by having a unique combination of morphological and color pattern characters and bearing an uncorrected pairwise sequence divergence of 9.5%–25.5%.
References
- 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.x
- Bauer, A.M., Sumontha, M. & Pauwels, O.S.G. (2003) Two new species of Cyrtodactylus (Reptilia: Squamata: Gekkonidae) from Thailand. Zootaxa, 376 (1), 1–18. https://doi.org/10.11646/zootaxa.376.1.1
- Bouckaert, R.R. & Drummond, A.J. (2017) bModelTest: Bayesian phylogenetic site model averaging and model comparison. BMC Evolutionary Biology, 17 (42), 1–11. https://doi.org/10.1186/s12862-017-0890-6
- Bui, M.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
- De Queiroz, K. (2007) Species concepts and species delimitation. Systematic Biology, 56, 879–886. https://doi.org/10.1080/10635150701701083
- Drummond, A.J., Suchard, M.A., Xie, D. & Rambaut, A. (2012) Bayesian phylogenetics with BEAUti and the BEAST 1.7. Molecular Biology and Evolution, 29, 1969–1973. https://doi.org/10.1093/molbev/mss075
- Frost, D.R. & Hillis, D.M. (1990) Species in concept and practice: herpetological application. Herpetologica, 46, 87–104.
- Frost, D.R. & Kluge, A.G. (1994) A consideration of epistemology in systematic biology, with special reference to species. Cladistics, 10, 259–294. https://doi.org/10.1111/j.1096-0031.1994.tb00178.x
- Gray, J.E. (1831) A synopsis of the species of Class Reptilia. In: Griffith, E. & Pidgeon, E., The animal kingdom arranged in conformity with its organisation by the Baron Cuvier with additional descriptions of all the species hither named, and of many not before noticed. Whittaker, Treacher and Co., London, pp. 1–481 + 1–110.
- Grismer, L.L. & Grismer, J.L. (2017) A re-evaluation of the phylogenetic relationships of the Cyrtodactylus condorensis group (Squamata; Gekkonidae) and a suggested 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., Wood., P.L. Jr., Poyarkov, N.A., Le, M.D., Karunarathna, S., Chomdej, S., Suwannapoom, C., Qi, S., Liu, S., Che, J., Quah, E.S.H., Kraus, F., Oliver, P.M., Riyanto, A., Pauwels, O.S.G. & Grismer, J.L. (2021a) Karstic landscapes are foci of species diversity in the world’s third-largest vertebrate genus Cyrtodactylus Gray, 1827 (Reptilia: Squamata; Gekkonidae). Diversity, 13 (5), 183. https://doi.org/10.3390/d13050183
- Grismer, L.L., Poyarkov, N.A., Quah, E.S.H., Grismer, J.L. & Wood, P.L. Jr. (2022) The biogeography of bent-toed geckos, Cyrtodactylus (Squamata: Gekkonidae). PeerJ, 10, e13153 https://doi.org/10.7717/peerj.13153
- Grismer, L.L., Wood, P.L. Jr., Thura, M.K., 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 underscore the urgent need for the conservation of karst habitats. Journal of Natural History, 52 (19–20), 1243–1294. https://doi.org/10.1080/00222933.2018.1449911
- Grismer, L.L., Wood, P.L. Jr., Quah, E.S.H., Grismer, M.S., Thura, M.K., Oaks J.R. & Lin, A. (2020) Two new species of Cyrtodactylus Gray, 1827 (Squamata: Gekkonidae) from a karstic archipelago in the Salween Basin of southern Myanmar (Burma). Zootaxa, 4718 (2), 151–183. https://doi.org/10.11646/zootaxa.4718.2.1
- Grismer, L.L., Wood, P.L. Jr., Thura, M.K., Zin T., Quah, E.S.H., Murdoch, M.L., Grismer, M.S., Lin, A., Kyaw, H. & Lwin, N. (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. https://doi.org/10.1093/zoolinnean/zlx057
- Grismer, L.L., Aowphol, A., Grismer, J.L., Aksonneam, A., Quah. E.S.H., Murdoch, M.L., Nguyen, E., Kaatz, A., Bringsøe, H. & Rujirwana, A. (2024) A new species of the Cyrtodactylus chauquangensis group (Squamata: Gekkonidae) from northern Thailand. ZooKeys. [submitted]
- Heidrich, A., Rösler, H., Vu, T.N., Böhme, W. & Ziegler, T. (2007) Another new Cyrtodactylus (Squamata: Gekkonidae) from Phong Nha-Ke Bang National Park, central Truong Son, Vietnam. Zootaxa, 1445 (1), 35–48. https://doi.org/10.11646/zootaxa.1445.1.3
- 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., Bui, M.Q. & Le, V.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
- Husson, F., Josse, J., Le, S. & Mazet, J. (2017) FactoMine R: exploratory data analysis and data mining. R package, version 1.36. [program]
- Loos, J., von Wehrden, H., Dang, K.N. & Ziegler, T. (2011) Niche Segregation In Microhabitat Use Of Three Sympatric Cyrtodactylus In The Phong Nha-Ke Bang National Park, Central Vietnam. Herpetological Conservation and Biology, 7 (1), 101–108. [https://herpconbio.org/Volume_7/Issue_1/Loos_etal_2012.pdf]
- Kassambara, A. & Mundt, F. (2017) Factoextra: extract and visualize the result of multivariate data analyses. R package, version 1.0.5.999. [program]
- 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
- Luu, V.Q., Bonkowski, M., Nguyen, T.Q., Le, M.D., Schneider, N., Ngo, H.T. & Ziegler, T. (2016a) Evolution in karst massifs: Cryptic diversity among bent-toed geckos along the Truong Son Range with descriptions of three new species and one new country record from Laos. Zootaxa, 4107 (2), 101–140. https://doi.org/10.11646/zootaxa.4107.2.1
- Luu, V.Q., Calame, T., Bonkowski, M., Nguyen, Q. T. & Ziegler, T. (2014) A new species of Cyrtodactylus (Squamata: Gek- konidae) from Khammouane Province, Laos. Zootaxa, 3760 (1), 54−66. https://doi.org/10.11646/zootaxa.3760.1.3
- Luu, V.Q., Calame, T., Nguyen, T.Q., Bonkowski, M. & Ziegler, T. (2015) A new species of Cyrtodactylus (Squamata: Gek- konidae) from limestone forest, Khammouane Province, central Laos. Zootaxa, 4058 (3), 388–402. https://doi.org/10.11646/zootaxa.4058.3.6
- Luu, V.Q., Calame, T., Nguyen, T.Q., Le, M.D., Bonkowski, M. & Ziegler, T. (2016b) Cyrtodactylus rufford, a new cave-dwelling bent-toed gecko (Squamata: Gekkonidae) from Khammouane Province, central Laos. Zootaxa, 4067 (2), 185–199. https://doi.org/10.11646/zootaxa.4067.2.4
- Luu, V.Q., Dung, T.V., Nguyen, T.Q., Le, M.D. & Ziegler, T. (2017) A new species of the Cyrtodactylus irregularis complex (Squamata: Gekkonidae) from Gia Lai Province, Central Highlands. Zootaxa, 4362 (3), 385–404. https://doi.org/10.11646/zootaxa.4362.3.4
- Luu, V.Q., Nguyen, T.Q., Do, H.Q. & Ziegler, T. (2011) A new Cyrtodactylus (Squamata: Gekkonidae) from Huong Son lime- stone forest, Hanoi, northern Vietnam. Zootaxa, 3129 (1), 39–50. https://doi.org/10.11646/zootaxa.3129.1.3
- Luu, V.Q., Nguyen, T.Q., Le, M.D., Bonkowski, M. & Ziegler, T. (2016c) A new species of karst-dwelling bent-toed gecko (Squamata: Gekkonidae) from Khammouane Province, central Laos. Zootaxa, 4079 (1), 87–102. https://doi.org/10.11646/zootaxa.4079.1.6
- Miller, M.A., Pfeiffe, W. & Schwartz, T. (2010) Creating the CIPRES Science Gateway for inference of large phylogenetic trees. Gateway Computing Environments Workshop (GCE), New Orleans (USA), November 2010, IEEE, 1–8. https://doi.org/10.1109/GCE.2010.5676129
- Maddison, W.P. & Maddison, D.R. (1992) MacClade version 3: Analysis of phylogeny and character evolution. Sinauer Associates, Sunderland, Massachusetts. [program]
- Murdoch, M.L., Grismer, L.L., Wood, P.L. Jr., Neang, T., Poyarkov, N.A., Ngo, V.T., Nazarov, R.A., Aowphol, A., Pauwels, O.S.G., Nguyen, H.N. & Grismer, J.L. (2019) Six new species of the Cyrtodactylus intermedius complex (Squamata: Gekkonidae) from the Cardamom Mountains and associated highlands of Southeast Asia. Zootaxa, 4554 (1), 1–62. https://doi.org/10.11646/zootaxa.4554.1.1
- Nazarov, R.A., Pauwels, O.S.G., Konstantinov, E.L., Chulisov, A.S., Orlov, N.L. & Poyarkov, N.A. Jr. (2018) A new karst-dwelling bent-toed gecko (Squamata: Gekkonidae: Cyrtodactylus) from Xiangkhoang Province, northeastern Laos. Zoological Research, 39 (3), 202–219. https://doi.org/10.24272/j.issn.2095-8137.2018.010
- Ngo, T.V. & Pauwels, O.S.G. (2010) A new cave-dwelling species of Cyrtodactylus Gray, 1827 (Squamata: Gekkonidae) from Khammouane Province, southern Laos. Zootaxa, 2730 (1), 44–56. https://doi.org/10.11646/zootaxa.2730.1.3
- 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
- Nazarov, R.A., Poyarkov, N.A.Jr., Orlov, N.L., Nguyen, N.S., Milto, K.D., Martynov, A.A., Konstantinov, E.L. & Chulisov, A.S. (2015) A review of genus Cyrtodactylus (Reptilia: Sauria: Gekkonidae) in fauna of Laos with description of four new species. Труды Зоологического института РАН, 318 (4), 391–423. [Proceedings of the Zoological Institute RAS, 318 (4), 391–423 (2014)] https://doi.org/10.31610/trudyzin/2014.318.4.391
- Nguyen, D.X., Nguyen, H.T., Nguyen, D.Q. & Nguyen, N.X. (2016) Nature Conservation and Sustainable Natural Resource Management in Phong Nha-Ke Bang National Park Region Project, Quang Binh Province, Vietnam (Phong Nha-Ke Bang Region Project). Available from: https://hinnamno.org/wp-content/uploads/formidable/11/30.Nguyen_2016_BIODIVERSITY IN PHONG NHA-KE BANG AND HIN NAM NO.pdf (accessed 4 February 2024)
- Pagès, J. (2015) Multiple Factor Analysis by Example Using R. CRC Press, New York, New York, 272 pp. https://doi.org/10.1201/b17700
- Rambaut, A. & Drummond, A.J. (2013) TreeAnnotator v1.8.0 MCMC Output Analysis.
- Rambaut, A., Drummond, A.J., Xie, D., Baele, G. & Suchard, M.A. (2018) Posterior summarization in Bayesian phylogenetics using Tracer 1.7. Systemastic Biology, 67, 901–904. https://doi.org/10.1093/sysbio/syy032
- Reist, J.D. (1986) An empirical evaluation of coefficients used in residual and allometric adjustment of size covariation. Canadian Journal of Zoology, 64, 1363–1368. https://doi.org/10.1139/z86-203
- Ronquist, F., Teslenko, M., Van Der Mark, P., Ayres, D.L., Darling, A., Höhna, S., Larget, B., Liu, L., Suchard, M.A. & Huelsenbeck, J.P. (2012) MrBayes 3.2: Efficient Bayesian phylogenetic inference and model choice across a large model space. Systematic Biology, 61 (3), 539–542. https://doi.org/10.1093/sysbio/sys029
- Sitthivong, S., Luu, V.Q., Ha, N.V., Nguyen, T.Q., Le, M.D. & Ziegler, T. (2019) A New Species of Cyrtodactylus (Squamata: Gekkonidae) from Vientiane Province, northern Laos. Zootaxa, 4701 (3), 257–275. https://doi.org/10.11646/zootaxa.4701.3.3
- Smith, M.A. (1921) New or little-known reptiles and batrachians from southern Annam (Indochina). Proceedings of the Zoological Society of London, 1921, 423–440. https://doi.org/10.1111/j.1096-3642.1921.tb03271.x
- Tamura, K., Stecher, G. & Kumar, S. (2021) MEGA11: Molecular evolutionary genetics analysis version 11. Molecular Biology and Evolution, 38, 3022–3027. https://doi.org/10.1093/molbev/msab120
- Thorpe, R.S. (1975) Quantitative handling of characters useful in snake systematics with particular reference to intraspecific 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. Vol. 1. Springer, Berlin and Heidelberg, pp. 404–423. https://doi.org/10.1007/978-3-642-69024-2_43
- Trifinopoulos, J., Nguyen, L.T., von Haeseler, A. & Bui, M.Q. (2016) W-IQ-TREE: a fast online phylogenetic tool for maximum likelihood analysis. Nucleic Acids Research, 44, 232–235. 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., 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
- Ziegler, T., Phung, T.M., Le, M.D. & Nguyen, T.Q. (2013) A new Cyrtodactylus (Squamata: Gekkonidae) from Phu Yen Province, southern Vietnam. Zootaxa, 3686 (4), 432–446. https://doi.org/10.11646/zootaxa.3686.4.2
- Ziegler, T. & Vu, T.N. (2009) Ten years of herpetodiversity research in Phong Nha-Ke Bang National Park, Central Vietnam. In: Tri, N.T.D., Dang, K.N. & Yen, P.T.H. (Eds.), Phong Nha-Ke Bang National Park and Cologne Zoo, 10 years of Cooperation (1999–2009), Quang Binh, 2009, pp. 103–124.
- Ziegler, T., Nazarov, R., Orlov, N., Nguyen, T.Q., Vu, T.N., Dang, K.N., Dinh, T.H. & Schmitz, A. (2010) A third new Cyrtodactylus (Squamata: Gekkonidae) from Phong Nha-Ke Bang National Park, Truong Son Range, Vietnam. Zootaxa, 2413 (1), 20–36. https://doi.org/10.11646/zootaxa.2413.1.2