Abstract
An adult sand snake specimen was collected during a herpetofaunal survey conducted in the Turpan Basin in northwest China. Phylogenetic analyses revealed that this specimen, along with other snake sloughs and skins collected from different localities in the Turpan Basin formed a clade that is sister to Psammophis lineolatus. This taxon exhibited substantial divergence from its congeners (P. lineolatus and P. condanarus) with uncorrelated p-distances ranging from 11.9 ± 0.9% to 15.8 ± 1.6% for the ND4 gene and from 10.2 ± 0.8% to 13.8 ± 1.1% for the Cytb gene. Given the genetic differences along with morphological differences, we describe the specimen from the Turpan Basin as Psammophis turpanensis sp. nov. We provide detailed morphological descriptions, and compare this specimen with five Asian sand snakes and the Afro-Asian Sand Snake, P. schokari. In addition, we provide brief comments on the biogeography of Psammophis in China.
References
Aini, G., Aji, P. & Ubul, G. (2011) Analysis of the effect of global climate change on East Xinjiang Region (Turpan Basin as an example). Journal of Jinggangshan University (Natural Science), 32 (3), 70–75. [in Chinese with English abstract]
Amr, Z.S. & Disi, A.M. (2011) Systematics, distribution and ecology of the snakes of Jordan. Vertebrate Zoology, 61 (2), 179–266.
Ananjeva, N.B., Munkhbayar, K., Orlov, N.L., Orlova, V.F., Semenov, D.V., Terbish, K. (1997) Amphibians and Reptiles of Mongolia. Reptiles of Mongolia. In: Sokolov, V.E. (Ed.), Vertebrates of Mongolia. KMK Limited, Moscow, pp. 1–416. [in Russian]
Bannikov, A.G., Darevsky, I.S., Ischchenko, V.G., Rustamov, A.K. & Szczerbak, N.N. (1977) Guide to Amphibians and Reptiles of the USSR. Moscow: Prosveschenie Press, 414 pp. [in Russian]
Bedriaga, J.V. (1907) Verzeichnis der von der Central-Asiatischen expedition unter stabs-kapitän W. Roborowski in den Jahren 1893–1895 gesammelten reptilien. Annuaire du Musée Zoologique de l’Académie Impériale des Sciences de St.-Pétersbourg, 10 (3–4), 159–200. [in German]
Boulenger, G.A. (1896) Catalogue of the Snakes in the British Museum (Natural History). Vol. III. Taylor and Francis, London, xiv + 727 pp.
Branch, W.R., Baptista, N., Keates, C. & Edwards, S. (2019) Rediscovery, taxonomic status, and phylogenetic relationships of two rare and endemic snakes (Serpentes: Psammophiinae) from the southwestern Angolan plateau. Zootaxa, 4590 (3), 4342–4366.
https://doi.org/10.11646/zootaxa.4590.3.2
Broadley, D.G. (1966) A review of the African Stripe-bellied Sand-snakes of the genus Psammophis. Arnoldia, Rhodesia, 2, 1–9.
Broadley, D.G. (1977) A review of the genus Psammophis in southern Africa (Serpentes: Colubridae). Arnoldia, Rhodesia, 8, 1–29.
Cheng, J., Lv, X., Xia, L., Ge, D., Zhang, Q., Lu, L. & Yang, Q. (2019) Impact of orogeny and environmental change on genetic divergence and demographic history of Dipus sagitta (Dipodoidea, Dipodinae) since the Pliocene in inland East Asia. Journal of Mammalian Evolution, 26, 253–266.
https://doi.org/10.1007/s10914-017-9397-6
Chirikova, M.A., Dujsebayeva, T.N., Liu, J. & Guo, X. (2019) Geographical distribution and morphological variability of the Rapid Racerunner, Eremias velox (Pallas, 1771) (Reptilia, Lacertidae) in the Eastern Periphery of its range. Asian Herpetological Research, 10 (4), 230–245.
https://doi.org/10.16373/j.cnki.ahr.190009
Cottone, A.M. & Bauer, A.M. (2009a) Sexual size dimorphism, diet, and reproductive biology of the Afro-Asian Sand Snake, Psammophis schokari (Psammophiidae). Amphibia-Reptilia, 30 (3), 331–340.
https://doi.org/10.1163/156853809788795182
Cottone, A.M. & Bauer, A.M. (2009b) Notes on sexual size dimorphism and reproduction in the Asian Sand Snake, Psammophis condanarus (Psammophiidae). Hamadryad, 34 (1), 181–184.
https://doi.org/10.1163/156853809788795182
Dayrat, B. (2005) Towards integrative taxonomy. Biological Journal of the Linnean Society, 85 (3), 407−417.
https://doi.org/10.1111/j.1095-8312.2005.00503.x
de Queiroz, K. (1998) The general lineage concept of species, species criteria, and the process of speciation: a conceptual unification and terminological recommendations. In: Howard, D.J. & Berlocher, S.H. (Eds.), Endless Forms: Species and Speciation. Oxford University Press, New York, New York, pp. 57–75.
de Queiroz, K. (2007) Species concepts and species delimitation. Systematic Biology, 56 (6), 879–886.
https://doi.org/10.1080/10635150701701083
Doronin, I.V., Dujsebayeva, T.N., Akhmedenov, K.M., Bakiev, A.G. & Plakhov, K.N. (2020) On the type locality of the Steppe Ribbon Racer, Psammophis lineolatus (Brandt, 1838) (Serpentes: Lamprophiidae). Proceedings of the Zoological Institute RAS, 324 (2), 262–272. [in Russian with English abstract]
https://doi.org/10.31610/trudyzin/2020.324.2.262
Dowling, H.G. (1951) A proposed standard system of counting ventral in snakes. British Journal of Herpetology, 1 (5), 97–99.
Edgar, R.C. (2004) MUSCLE: multiple sequence alignment with high accuracy and high throughput. Nucleic Acids Research, 32 (5), 1792–1797.
https://doi.org/10.1093/nar/gkh340
Erixon, P., Svennblad, B., Britton, T. & Oxelman, B. (2003) Reliability of Bayesian posterior probabilities and bootstrap frequencies in phylogenetics. Systematic Biology, 52 (5), 665–673.
https://doi.org/10.1080/10635150390235485
Felsenstein, J. (1985) Confidence limits on phylogenies: an approach using the bootstrap. Evolution, 39 (4), 783–791.
https://doi.org/10.2307/2408678
Ganesh, S.R., Sharma, V. & Guptha, M.B. (2017) Records of the Indian Sand Snake Psammophis condanarus (Merrem, 1820) (Reptilia: Lamprophiidae) in southern India. Journal of Threatened Taxa, 9 (7), 10453–10458.
https://doi.org/10.11609/jott.3468.9.7.10453-10458
Gonçalves D.V., Martínez-Freiría, F., Crochet, P.A., Geniez, P., Carranza, S. & Brito, J.C. (2018) The role of climatic cycles and trans-Saharan migration corridors in species diversification: biogeography of Psammophis schokari group in North Africa. Molecular Phylogenetics and Evolution, 118, 64–74.
https://doi.org/10.1016/j.ympev.2017.09.009
Gouy, M., Guindon, S. & Gascuel, O. (2010) SeaView version 4: A multiplatform graphical user interface for sequence alignment and phylogenetic tree building. Molecular Biology and Evolution, 27 (2), 221–224.
https://doi.org/10.1093/molbev/msp259
Greene, T.J., Carroll, A.R., Wartes, M., Graham, S.A. & Wooden, J.L. (2005) Integrated provenance analysis of a complex orogenic terrane: mesozoic uplift of the Bogda Shan and inception of the Turpan-Hami Basin, NW China. Journal of Sedimentary Research, 75 (2), 251–267.
https://doi.org/10.2110/jsr.2005.019
Guo, X. & Wang, Y. (2007) Partitioned Bayesian analyses, dispersal-vicariance analysis, and the biogeography of Chinese toad-headed lizards (Agamidae: Phrynocephalus): A reevaluation. Molecular Phylogenetics and Evolution, 45 (2), 643–662.
Haidar, I.K.A., Romon, B.B. & Chowdhury, M.A.W. (2020) First record of the Condanarous sand snake (Psammophis condanarus) (Reptilia: Squamata: Lamprophiidae) from Bangladesh. Journal of Asia-Pacific Biodiversity, 13 (4), 716–719.
https://doi.org/10.1016/j.japb.2020.08.013
Hartmann, T., Handschuh, M. & Böhme, W. (2011) First record of Psammophis indochinensis Smith, 1943 from Cambodia, within the context of a distributional species account. Cambodian Journal of Natural History, 1, 7–10.
Hillis, D.M. (2019) Species delimitation in herpetology. Journal of Herpetology, 53 (1), 3–12.
https://doi.org/10.1670/18-123
Hillis, D.M. & Bull, J.J. (1993) An empirical test of bootstrapping as a method for assessing confidence in phylogenetic analysis. Systematic Biology, 42 (2), 182–192.
https://doi.org/10.1093/sysbio/42.2.182
Huelsenbeck, J.P. & Rannala, B. (2004) Frequentist properties of Bayesian posterior probabilities of phylogenetic trees under simple and complex substitution models. Systematic Biology, 53 (6), 904–913.
https://doi.org/10.1080/10635150490522629
Hughes, B. (1999) Critical review of a revision of Psammophis (Linnaeus 1758) (Serpentes, Reptilia) by Frank Brandstätter. African Journal of Herpetology, 48 (1-2), 63–70.
https://doi.org/10.1080/21564574.1999.9651073
Kark, S., Warburg, I. & Werner, Y.L. (1993) Polymorphism of Psammophis schokari (Ophidia: Colubridae) in Israel and Sinai. Journal of Arid Environments, 37, 513–527.
https://doi.org/10.1006/jare.1997.0296
Kelly, C.M.R., Barker, N.P., Villet, M.H., Broadley, D.G. & Branch, W.R. (2008) The snake family Psammophiidae (Reptilia: Serpentes): phylogenetics and species delimitation in the African sand snakes (Psammophis Boie, 1825) and allied genera. Molecular Phylogenetics and Evolution, 47 (3), 1045–1060.
https://doi.org/10.1016/j.ympev.2008.03.025
Khan, M.S. (2002) A Guide to the Snakes of Pakistan. Edition Chimaira, Frankfurt-am-Main, 73 pp.
Kumar, S., Stecher, G. & Tamura, K. (2016) MEGA7: molecular evolutionary genetics analysis version 7.0 for bigger datasets. Molecular Biology and Evolution, 33 (7), 1870–1874.
https://doi.org/10.1093/molbev/msw054
Lanfear, R., Frandsen, P.B., Wright, A.M., Senfeld, T. & Calcott, B. (2017) PartitionFinder 2: new methods for selecting partitioned models of evolution for molecular and morphological phylogenetic analyses. Molecular Biology and Evolution, 34 (3), 772–773.
https://doi.org/10.1093/molbev/msw260
Li, G., Peng, Z., Zhang, R., Tang, Y., Tong, C., Feng, C., Zhang, C. & Zhao, K. (2016) Mito-nuclear phylogeography of the cyprinid fish Gymnodiptychus dybowskii in the arid Tien Shan region of Central Asia. Biological Journal of the Linnean Society, 118, 304–314.
https://doi.org/10.1111/bij.12724
Liu, J., Guo, X., Chen, D., Li, J., Yue, B. & Zeng, X. (2019) Diversification and historical demography of the rapid racerunner (Eremias velox) in relation to geological history and Pleistocene climatic oscillations in arid Central Asia. Molecular Phylogenetics and Evolution, 130, 244–258.
https://doi.org/10.1016/j.ympev.2018.10.029
Macey, J.R., Ananjeva, N.B., Wang, Y. & Papenfuss, T.J. (1997) A taxonomic reevaluation of the gekkonid lizard genus Teratoscincus in China. Russian Journal of Herpetology, 4 (1), 8–16.
https://doi.org/10.30906/1026-2296-1997-4-1-8-16
Macey, R.J., Fong, J.J., Kuehl, J.V., Shafiei, S., Ananjeva, N.B., Papenfuss, T.J. & Boore, J.L. (2005). The complete mitochondrial genome of a gecko and the phylogenetic position of the Middle Eastern Teratoscincus keyserlingii. Molecular Phylogenetics and Evolution, 36 (1), 188–93.
https://doi.org/10.1016/j.ympev.2005.03.025
Meng, W., Yang, T.Y., Guo, Y., Hai, S., Ma, Y.W, Ma, X.F. & Cai, L.G. (2015) Remarkable genetic divergence of Gymnodiptychus dybowskii between south and north of Tianshan Mountain in northwest China. Biochemical Systematics and Ecology, 58, 43–50.
https://doi.org/10.1016/j.bse.2014.10.005
Meng, H.H. & Zhang, M.L. (2013) Diversification of plant species in arid Northwest China: species-level phylogeographical history of Lagochilus Bunge ex Bentham (Lamiaceae). Molecular Phylogenetics and Evolution, 68 (3), 398–409.
https://doi.org/10.1016/j.ympev.2013.04.012
Miller, A.H., Davis, H.R., Luong, A.M., Do, Q.H., Pham, C.T., Ziegler, T., Lee, J.L., De Queiroz, K., Reynolds, R.G. & Nguyen, T.Q. (2020) Discovery of a new species of enigmatic Odd-Scaled Snake (Serpentes: Xenodermidae: Achalinus) from Ha Giang Province, Vietnam. Copeia, 108 (4), 768–808.
https://doi.org/10.1643/CH2020060
Neang, T., Hartmann, T., Hun, S., Souter, N.J. & Furey, N.M. (2014) A new species of wolf snake (Colubridae: Lycodon Fitzinger, 1826) from Phnom Samkos Wildlife Sanctuary, Cardamom Mountains, southwest Cambodia. Zootaxa, 3814 (1), 68–80. https://doi.org/10.11646/zootaxa.3814.1.3
Nei, M. (1987) Molecular Evolutionary Genetics. Columbia University Press, New York, 512 pp.
Padial, J.M., Miralles, A., De la Riva, I. & Vences, M. (2010) The integrative future of taxonomy. Frontiers in Zoology, 7, 16.
https://doi.org/10.1186/1742-9994-7-16
Przewalski, N.M. (1876) Mongolia and the Tangut Country. A Three-Year Travel in Eastern High Asia. Vols. 1-2. Imperial Russian Geographic Society, St. Petersburg, 55 pp. [In Russian]
Rambaut, A., Drummond, A.J, Xie, D., Baele, G. & Suchard, M.A. (2018) Posterior summarization in bayesian phylogenetics using Tracer 1.7. Systematic Biology, 67 (5), 901–904.
https://doi.org/10.1093/sysbio/syy032
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
Schwarz, G. (1978) Estimating the dimension of a model. The Annals of Statistics, 6 (2), 461–464.
https://doi.org/10.1214/aos/1176344136
Shammakov, S.M. (1981) Reptiles of the Plains of Turkmenistan. Ylym Press, Ashkhabad, 312 pp. [in Russian]
Shi, L. & Zhao, E.M. (2011) A new gecko in the genus Cyrtopodion Fitzinger, 1843 (Reptilia: Squamata: Gekkonidae) from western China. Herpetologica, 67 (2), 186–193.
https://doi.org/10.1655/09-036.1
Smith, M.A. (1943) The Fauna of British India, Ceylon and Burma including the Whole of the Indo-Chinese Sub-region. Reptilia and Amphibia. Vol. III. Serpentes. Taylor and Francis, London, 583 pp.
Stamatakis, A. (2006) RAxML-VI-HPC: maximum likelihood-based phylogenetic analyses with thousands of taxa and mixed models. Bioinformatics, 22 (21), 2688–2690.
https://doi.org/10.1093/bioinformatics/btl446
Sun, J.M., Zhu, R.X. & Bowler, J. (2004) Timing of the Tianshan Mountains uplift constrained by magnetostratigraphic analysis of molasse deposits. Earth and Planetary Science Letters, 219 (3–4), 239–253.
https://doi.org/10.1016/S0012-821X(04)00008-1
Taft, J.M. (2018) Phylogenetic Assessment and Historical Biogeography of the Psammophis leightoni Complex. Master Thesis. University of Western Cape, Bellville, Cape Town, 54 pp.
Tamar, K., Els, J., Kornilios, P., Soorae, P., Tarroso, P., Thanou, E., Pereira, J., Shah, J.N., Elhassan, E.E.M., Aguhob, J.C., Badaam, S.F., Eltayeb, M.M., Pusey, R., Papenfuss, T.J., Macey J.R. & Carranza, S. (2021) The demise of a wonder: evolutionary history and conservation assessments of the Wonder Gecko Teratoscincus keyserlingii (Gekkota, Sphaerodactylidae) in Arabia. PLoS ONE, 16 (1), e0244150.
https://doi.org/10.1371/journal.pone.0244150
Trape, J.F., Crochet, P.A., Broadley, D.G., Sourouille, P., Mané, Y., Burger, M., Böhme, W., Saleh, M., Karan, A., Lanza, B. & Mediannikov, O. (2019) On the Psammophis sibilans group (Serpentes, Lamprophiidae, Psammophiinae) north of 12°S, with the description of a new species from West Africa. Bonn Zoological Bulletin, 68 (1), 61–91.
https://doi.org/10.20363/BZB-2019.68.1.061
Trifonov, V.G., Ivanova, T.P. & Bachmanov, D.M. (2012) Recent mountain building of the central Alpine-Himalayan Belt. Geotectonics, 46 (5), 315–332.
https://doi.org/10.1134/S0016852112050068
Uetz, P., Freed, P. & Hošek, J. (2020) The Reptile Database. Available from: http://www.reptile-database.org (accessed 20 December 2020)
Vyas, R. & Patel, H. (2013) Notes on distribution and natural history of Psammophis longifrons Boulenger 1896 (Serpentes: Psammophiidae: Psammophiinae) in Gujarat, India. Russian Journal of Herpetology, 20 (3), 217–222.
Wang, L., Ji, J., Sun, D., Xu, Q. & Han, B. (2015) Chronological constraints on multi-staged rapid cooling of the Tianshan Mountains inferred from apatite fission track analysis of modern river sands. Science China Earth Sciences, 58 (8), 1305–1319.
https://doi.org/10.1007/s11430-015-5072-z
Wang, K., Yu, Z.-B., Vogel, G. & Che, J. (2021) Contribution to the taxonomy of the genus Lycodon H. Boie in Fitzinger, 1827 (Reptilia: Squamata: Colubridae) in China, with description of two new species and resurrection and elevation of Dindon septentrionale chapanense Angel, Bourret, 1933. Zoological Research, 42 (1), 62–86.
https://doi.org/10.24272/j.issn.2095-8137.2020.286
Wang, Y., Zhao, L.M., Fang, F.J., Liao, J.C. & Liu, N.F. (2013) Intraspecific molecular phylogeny and phylogeography of the Meriones meridianus (Rodentia: Cricetidae) complex in northern China reflect the processes of desertification and the Tianshan Mountains uplift. Biological Journal of the Linnean Society, 110 (2), 362–383.
https://doi.org/10.1111/bij.12123
Wallach, V., Williams, K.L. & Boundy, J. (2014) Snakes of the World: A Catalogue of Living and Extinct Species. Taylor and Francis, CRC Press, Florida, 1237 pp.
https://doi.org/10.1201/b16901
Whitaker, R. & Captain, A. (2004) Snakes of India: the Field Guide. Draco Books, Chengelpat, Tamil Nadu, 479 pp.
Windley, B.F., Allen, M.B., Zhang, C., Zhao, Z.-Y. & Wang, G.R. (1990) Paleozoic accretion and Cenozoic redeformation of the Chinese Tien Shan Range, central Asia. Geology, 18, 128–131.
https://doi.org/10.1130/0091-7613(1990)018<0128:PAACRO>2.3.CO;2
Zaher, H., Murphy, R.W., Arredondo, J.C., Graboski, R., Machado-Filho, P.R., Mahlow, K., Montingelli, G.G., Quadros, A.B., Orlov, N.L., Wilkinson, M., Zhang, Y.P. & Grazziotin, F.G. (2019) Large-scale molecular phylogeny, morphology, divergence-time estimation, and the fossil record of advanced caenophidian snakes (Squamata: Serpentes). PLoS ONE, 14 (5), e0216148.
https://doi.org/10.1371/journal.pone.0216148
Zhang, H.X. & Zhang, M.L. (2012) Genetic structure of the Delphinium naviculare species group tracks Pleistocene climatic oscillations in the Tianshan Mountains, arid Central Asia. Palaeogeography Palaeoclimatology Palaeoecology, 353–355, 93–103.
https://doi.org/10.1016/j.palaeo.2012.07.013
Zhao, E.M. (2006) The Snakes of China. Vol. I. Anhui Sience & Technology Publishing House, Hefei, 372 pp. [in Chinese]
Zhao, E.M., Huang, M.H. & Zong, Y. (1998) Fauna Sinica: Reptilia. Vol. 3. Squamata, Serpentes. Science Press, Beijing, 522 pp. [in Chinese]