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Type: Monograph
Published: 2018-09-03
Page range: 1–81
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A molecular phylogenetic hypothesis for the Asian agamid lizard genus Phrynocephalus reveals discrete biogeographic clades implicated by plate tectonics

Genomics, Department of Biosciences, Merritt College, 12500 Campus Drive, Oakland, CA 94619, USA Museum of Vertebrate Zoology, University of California, Berkeley, CA 94720, USA
Department of Biology, 700 College Street, Center for the Sciences 338, Beloit College, Beloit, WI 53511, USA
Zoological Institute, Russian Academy of Sciences, St. Petersburg 199034, Russia
Genomics, Department of Biosciences, Merritt College, 12500 Campus Drive, Oakland, CA 94619, USA
Chengdu Institute of Biology, PO Box 416, Chengdu, Sichuan, China
Zoological Institute, Russian Academy of Sciences, St. Petersburg 199034, Russia
Department of Biology, Faculty of Science, Shahid Bahonar University of Kerman, PO Box 76169-133, Kerman, Iran
Department of Biology, Sultan Qaboos University, PO Box 36, Al Khoud, PC 123, Muscat, Oman
Institute of Zoology, Ministry of Education and Sciences, al-Farabi Av., 93, Academgoradok, Almaty, 050060, Kazakhstan
Genomics, Department of Biosciences, Merritt College, 12500 Campus Drive, Oakland, CA 94619, USA
Genomics, Department of Biosciences, Merritt College, 12500 Campus Drive, Oakland, CA 94619, USA
Genomics, Department of Biosciences, Merritt College, 12500 Campus Drive, Oakland, CA 94619, USA
Museum of Vertebrate Zoology, University of California, Berkeley, CA 94720, USA
Reptilia Squamata Agamidae Phrynocephalus Asia biogeography evolution phylogenetics tectonics mitochondrial DNA RAG-1 allozyme

Abstract

Phylogenetic relationships of the agamid lizard genus Phrynocephalus are described in the context of plate tectonics. A near comprehensive taxon sampling reports three data sets: (1) mitochondrial DNA from ND1 to COI (3’ end of ND1, tRNAGln, tRNAIle, tRNAMet, ND2, tRNATrp, tRNAAla, tRNAAsn, tRNACys, tRNATyr, and the 5’ end of COI) with 1761 aligned positional sites (1595 included, 839 informative), (2) nuclear RAG-1 DNA with 2760 aligned positional sites (342 informative), and (3) 25 informative allozyme loci with 213 alleles (107 informative when coded as presence/absence). It is hypothesized that Phrynocephalus phyletic patterns and speciation reflect fault lines of ancient plates now in Asia rejuvenated by the more recent Indian and Arabian plate collisions. Molecular estimates of lineage splits are highly congruent with geologic dates from the literature.  A southern origin for the genus in Southwest Asia is resolved in phylogenetic estimates and a northern origin is statistically rejected. On the basis of monophyly and molecular evidence several taxa previously recognized as subspecies are recognized as species: P. hongyuanensis, P. sogdianus, and P. strauchi as “Current Status”; Phrynocephalus bannikovi, Phrynocephalus longicaudatus, Phrynocephalus turcomanus, and Phrynocephalus vindumi are formally “New Status”. Phylogenetic evaluation indicates a soft substrate habitat of sand for the shared ancestor of modern Phrynocephalus. Size diversity maximally overlaps in the Caspian Basin and northwestern Iranian Plateau. The greatest species numbers of six in sympatry and regional allopatry are found in the southern Caspian Basin and southern Helmand Basin, both from numerous phylogenetic lineages in close proximity attributed to tectonic induced events.

 

References

  1. Abdrakhmatov, K.Ye., Aldazhanov, S.A., Hager, B.H., Hamburger, M.W., Herring, T.A., Kalabaev, K.B., Makakarov, V.I., Molnar, P., Pananasyuk, S.V., Prilepin, M.T., Reilinger, R.E., Sadybakasov, I.S., Souter, B.J., Trapeznikov, Y.A., Tsurkov, V.Ye. & Zubovich, A.V. (1996) Relatively recent construction of the Tien Shan inferred from GPS measurements of present-day crustal deformation rates. Nature, 384, 450–453.

    https://doi.org/10.1038/384450a0

    Anderson, J. (1872) On some Persian, Himalayan, and other Reptiles. Proceedings of the Zoological Society of London, 1872 (2), 371–404.

    Anderson, S.C. (1999) The Lizards of Iran. Contribution to Herpetology 15, Society for the Study of Amphibians and Reptiles, Oxford, Ohio, 442 pp.

    Baig, K.J., Wagner, P., Ananjeva, N.B. & Böhme, W. (2012) A morphology-based taxonomic revision of Laudakia Gray, 1845 (Squamata: Agamidae). Vertebrate Zoology, 62 (2), 213–260.

    Bannikov, A.G., Darevsky, I.S., Ischchenko, V.G., Rustamov, A.K. & Szczerbak, N.N. (1977) Opredelitel’ Zemnovodnikh i Presmikayushchikhsya Fauni SSSR [Guide to the Reptile and Amphibian Fauna of the USSR]. Prosveshchenie, Moscow, 415 pp., figs. 1–108, 135 maps, 32 col. pls. [in Russian]

    Barabanov, A.V. & Ananjeva, N.B. (2007) Catalogue of the available scientific species-group names for lizards of the genus Phrynocephalus Kaup, 1825 (Reptilia, Sauria, Agamidae). Zootaxa, 1399, 1–56.

    https://doi.org/10.11646/zootaxa.1399.1.1

    Bedriaga, J. (“1905” 1906) 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.

    Bedriaga, J. (1907) Amphibien und Reptilien (Reptilia Przewalskiana). In: Wissenschaftliche Resultate der von N.M. Przewalski nach Central-Asien unternommenen Reisen. Zoologischer Theil, Bd. 3, Abth. 1, Lfrg. 2, St. Petersburg, pp. 71–278.

    Bedriaga, J. (1909) Amphibien und Reptilien (Reptilia Przewalskiana). In: Wissenschaftliche Resultate der von N.M. Przewalski nach Central-Asien unternommenen Reisen. Zoologischer Theil, Bd. 3, Abth. 1, Lfrg. 3, St. Petersburg, pp. 279–502.

    Bermingham, E., McCafferty, S.S. & Martin, A.P. (1997) Fish biogeography and molecular clocks: Perspectives from the Panamanian Isthmus. In: Kocher, T.D & Stepien C.A, (Eds.), Molecular Systematics of Fishes. Academic Press, San Diego, pp. 113–128.

    https://doi.org/10.1016/B978-012417540-2/50009-9

    Blanford, W.T. (1876) List of Reptilia and Amphibia collected by the late Dr. Stoliczka in Kashmir, Ladák, Eastern Turkestan and Wakhán, with descriptions of new species. Journal of the Asiatic Society of Bengal, 44 (3), 191–196.

    Bogdanov, O.P., Atayev, Ch. & Shammakov, S. (1974) The finding of Phrynocephalus maculatus in the USSR. Zoologicheskij Zhurnal, Moscow, 53 (2), 304–305. [in Russian]

    Boettger, O. (1888) Über die Reptilien und Batrachier Transcaspiens. Zoologischer Anzeiger, Leipzig, 11 (279), 259–263.

    Boulenger, G.A. (1887) Catalogue of the Lizards in the British Museum (Natural History), Second Edition. Vol. 3. Lacertidae, Gerrhosauridae, Scincidae, Anelytropidae, Dibamidae, Chamaeleontidae. Trustees of the British Museum, London, 575 pp.

    Buth, D.G. (1984) The application of electrophoretic data in systematic studies. Annual Review of Ecology and Systematics, 15, 501–522.

    https://doi.org/10.1146/annurev.es.15.110184.002441

    Chernov, S.A. (1948) Reptiles —Reptilia. In: Pavlovsky, E.N. & Vinogradova, B.S. (Eds.), The Animals of the USSR. Vol. 2. The desert zone. USSR Academy of Sciences, Moscow-Leningrad, pp. 127–161. [in Russian]

    Clayton, J.W. & Tretiak, D.N. (1972) Amine-citrate buffers for pH control in starch gel electrophoresis. Journal of the Fisheries Research Board Canada, 29, 1169–1172.

    https://doi.org/10.1139/f72-172

    Darevsky, I.S., Rustamov, A.K. & Shammakov, S. (1976) Status and distribution of Phrynocephalus reticulatus Eichwald (Sauria, Agamidae) in Central Asia. Teoreticheskije i prikladnyije aspekty okhrany prirody i okhotovedenija, Moscow, 84, 113–119. [in Russian]

    Dercourte, J., Zonenshain, L.P., Ricou, L.-E., Kazmin, V.G., Le Pichon, X., Knipper, A.L., Grandjacquet, C., Sbortshikov, I.M., Geyssant, J., Lepvrier, C., Pechersky, D.H., Boulin, J., Sibuet, J.-C., Savostin, L.A., Sorokhtin, O., Westphal, M., Bazhenov, M.L., Lauer, J.P. & Biju-Duval, B. (1986) Geological evolution of the Tethys belt from the Atlantic to the Pamirs since the Lias. Tectonophysics, 123, 241–315. [with maps, plates I–X]

    Dewey, J.F., Shackleton, R.M., Chang, C. & Sun, Y. (1988) The tectonic evolution of the Tibetan Plateau. In: The Geological Evolution of Tibet. Report of the 1985 Royal Society – Academia Sinica Geotraverse of the Qinghai – Xizang Plateau Led by Chang Chengfu, Robert M. Shackleton, F.R.S, John F. Dewey, F.R.S., and Yin Jixiang. Philosophical Transactions of the Royal Society of London, London, A 327 (1594), pp. 379–313.

    https://doi.org/10.1098/rsta.1988.0135

    Dewey, J.F., Cande, S. & Pitman, W.C., III. (1989) Tectonic evolution of the India/Eurasia collision zone. Eclogae Geologicae Helvetiae, 82 (3), 717–734.

    https://doi.org/10.5169/seals-166399

    Dunayev, E.A. (1995) Reviewed description of the types of Phrynocephalus strauchi Nikolsky, 1899 (Squamata, Agamidae) and materials on the history of its study, distribution, and variability. Russian Journal of Herpetology, 2 (2), 87–94.

    Eichwald, E. (1831) Zoologia specialis quam expositis animalibus tum vivis, tum fossilibus potissimum Rossiae in universum, et Poloniae in specie, in usum lectionum publicarum in Universitate Caesarea Vilnensi habendarum. Pars posterior. Josephi Zawadzki, Vilnae, 396 pp.

    Felsenstein, J. (1985) Confidence limits on phylogenies with a molecular clock. Systematic Zoology, 34 (4), 152–161.

    https://doi.org/10.1111/j.1558-5646.1985.tb00420.x

    Feng, Y., Coleman, R.G., Tilton, G. & Xiao, X. (1989) Tectonic evolution of the west Junggar region, Xinjiang, China. Tectonics, 8 (4), 729–752.

    https://doi.org/10.1029/TC008i004p00729

    Golubev, M.L. (1991) About the name Agama ocellata Lichtenstein in Eversmann, 1823 (Reptilia, Agamidae) with redescription of the types. Herpetological Researches, Leningrad, 1, 12–17. [in Russian with English summary]

    Golubev, M.L. (1998) A new subspecies of Phrynocephalus ornatus Boulenger (Reptilia: Agamidae) from Eastern Iran, with a key to South-Western and Middle Asian microphrynocephalids. Hamadryad, 23 (2), 162–168.

    Gozdik, A. & Fu, J. (2009) Are Toad-Headed lizards Phrynocephalus przewalskii and P. frontalis (family Agamidae) the same species? Defining species boundaries with morphological and molecular data. Russian Journal of Herpetology, 16 (2), 107–118.

    Haas, G. (1957) Some amphibians and reptiles from Arabia. Proceedings of the California Academy of Sciences, 29 (3), 47–86.

    Hall, R. (1996) Reconstructing Cenozoic SE Asia. In: Hall, R. & Blundell, D. (Eds.), Reconstructing Cenozoic SE Asia. Geological Society of London Special Publication No. 106, London, pp. 153–184.

    https://doi.org/10.1144/GSL.SP.1996.106.01.11

    Harris, H. & Hopkinson, D.A. (1976 et seq.) Handbook of Enzyme Electrophoresis in Human Genetics. Oxford: North Holland Publishing Co., Amsterdam. [loose leaf with supplements in 1977 and 1978]

    Jackman, T.R. & Wake, D.B. (1994) Evolutionary and historical analysis of protein variation in the blotched forms of salamanders of the Ensatina complex (Amphibia: Plethodontidae). Evolution, 48 (3), 876–897.

    https://doi.org/10.1111/j.1558-5646.1994.tb01369.x

    Jiang, Y.-M., Huang, Q.-Y. & Zhao, E.-M. (1980) A new subspecies of Phrynocephalus vlangalii (Strauch) and preliminary observation on its ecology. Acta Zoologica Sinica, Beijing, 26 (2), 178–183. [in Chinese with English summary]

    Jin, Y.-T., Brown, R.P. & Liu, N.-F. (2008) Cladogenesis and phylogeography of the lizard Phrynocephalus vlangalii (Agamidae) on the Tibetan plateau. Molecular Ecology, 17 (8), 1971–1982.

    https://doi.org/10.1111/j.1365-294X.2008.03721.x

    Jin, Y.-T. & Brown, R.P. (2013) Species history and divergence times of viviparous and oviparous Chinese toad-headed sand lizards (Phrynocephalus) on the Qinghai-Tibetan Plateau. Molecular Phylogenetics and Evolution, 68 (2), 259–268.

    https://doi.org/10.1016/j.ympev.2013.03.022

    Kamali, K. & Anderson, S.C. (2015) A new Iranian Phrynocephalus (Reptilia: Squamata: Agamidae) from the hottest place on earth and a key to the genus Phrynocephalus in southwestern Asia and Arabia. Zootaxa, 3904 (2), 249–260.

    https://doi.org/10.11646/zootaxa.3904.2.4

    Kaup, J.J. (1825) Einige Bemerkungen zu Merrems Handbuch [Some Remarks on Merrem’s Handbook]. Isis von Oken, 16, cols., 589–593. [in German]

    Kuchta, S.R., Parks, D.S., Mueller, R.L. & Wake, D.B. (2009) Closing the ring: Historical biogeography of the salamander ring species Ensatina eschscholtzii. Journal of Biogeography, 36 (5), 982–995.

    https://doi.org/10.1111/j.1365-2699.2008.02052.x

    Kumazawa, Y. & Nishida, M. (1993) Sequence evolution of mitochondrial transfer RNA genes and deep-branch animal phylogenetics. Journal of Molecular Evolution, 37 (4), 380–398.

    https://doi.org/10.1007/BF00178868

    Kwon, S.-T., Tilton, R.G., Coleman, R.G. & Feng, Y. (1989) Isotopic studies bearing on the tectonics of the west Junggar region, Xinjiang, China. Tectonics, 8 (4), 719–727.

    https://doi.org/10.1029/TC008i004p00719

    Lanfear, R., Calcott, B., Ho, S.Y.W. & Guindon, S. (2012) PartitionFinder: Combined selection of partitioning schemes and substitution models for phylogenetic analyses. Molecular Biology and Evolution, 29 (6), 1695–1701.

    https://doi.org/10.1093/molbev/mss020

    Larson, A. (1994) The comparison of morphological and molecular data in phylogenetic systematics. In: Schierwater, B., Streit, B., Wagner, G.P. & DeSalle, R. (Eds.), Molecular Ecology and Evolution: Approaches and Applications. Birkhäuser Verlag, Basel, pp. 371–390.

    https://doi.org/10.1007/978-3-0348-7527-1_22

    Leviton, A.E. & Anderson, S.C. (2010) The herpetological literature for Southwestern Asia: An indexed bibliography. Occasional Papers of the California Academy of Sciences, No. 157, i–vii, 1–622.

    Leviton, A.E. & Anderson, S.C. (2013) The herpetological literature for Southwestern Asia: An indexed bibliography Second Edition. Occasional Papers of the California Academy of Sciences, No. 161, i–x, 1–944.

    Lichtenstein, H. (1856) Nomenclator Reptilium et Amphibiorum Musei Zoologici Berolinensis. Namenverzeichniss der in der zoologischen Sammlung der Königlichen Universitat zu Berlin aufgestellten Arten von Reptilien und Amphibien nach ihren Ordnungen, Familien und Gattungen. Königlich Preussischen Akademie der Wissenschaften, Berlin, 48 pp. [in German]

    Mabee, P.M. & Humphries, J. (1993) Coding polymorphic data: Examples from allozymes and ontogeny. Systematic Biology, 42 (2), 166–181.

    https://doi.org/10.1093/sysbio/42.2.166

    Macey, J.R. (2005) Plethodontid salamander mitochondrial genomics: A parsimony evaluation of character conflict and implications for historical biogeography. Cladistics, 21 (2), 194–202.

    https://doi.org/10.1111/j.1096-0031.2005.00054.x

    Macey, J.R., Larson, A., Ananjeva, N.B., Fang, Z.L. & Papenfuss, T. J. (1997a) 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/10.1093/oxfordjournals.molbev.a025706

    Macey, J.R., Larson, A., Ananjeva, N.B. & Papenfuss, T J. (1997b) Replication slippage may cause parallel evolution in the secondary structures of mitochondrial transfer RNAs. Molecular Biology and Evolution, 14 (1), 30–39.

    https://doi.org/10.1093/oxfordjournals.molbev.a025699

    Macey, J.R., Larson, A., Ananjeva, N.B. & Papenfuss, T. J. (1997c) Evolutionary shifts in three major structural features of the mitochondrial genome among iguanian lizards. Journal of Molecular Evolution, 44 (6), 660–674.

    https://doi.org/10.1007/PL00006190

    Macey, J.R. & Verma, A. (1997) Homology in phylogenetic analysis: Alignment of transfer RNA genes and the phylogenetic position of snakes. Molecular Phylogenetics and Evolution, 7 (2), 272–279.

    https://doi.org/10.1006/mpev.1997.0379

    Macey, J.R., Schulte, J.A. II, Ananjeva, N.B., Larson, A., Rastegar-Pouyani, N., Shammakov, S.M. & Papenfuss, T.J. (1998a) Phylogenetic relationships among agamid lizards of the Laudakia caucasia species group: Testing hypotheses of biogeographic fragmentation and an area cladogram for the Iranian Plateau. Molecular Phylogenetics and Evolution, 10 (1), 118–131.

    https://doi.org/10.1006/mpev.1997.0478

    Macey, J.R., Schulte, J.A. II, Larson, A., Fang, Z., Wang, Y., Tuniyev, B.S. & Papenfuss, T.J. (1998b). Phylogenetic relationships of toads in the Bufo bufo species group from the eastern escarpment of the Tibetan Plateau: A case of vicariance and dispersal. Molecular Phylogenetics and Evolution, 9 (1), 80–87.

    https://doi.org/10.1006/mpev.1997.0440

    Macey, J.R., Wang, Y., Ananjeva, N.B., Larson, A. & Papenfuss, T.J. (1999) Vicariant patterns of fragmentation among gekkonid lizards of the genus Teratoscincus produced by the Indian Collision: A molecular phylogenetic perspective and an area cladogram for Central Asia. Molecular Phylogenetics and Evolution, 12 (3), 320-332.

    https://doi.org/10.1006/mpev.1999.0641

    Macey, J.R., Schulte, J.A. II & Larson, A. (2000a) Evolution and phylogenetic information content of mitochondrial genomic structural features illustrated with acrodont lizards. Systematic Biology, 49 (1), 257–277.

    https://doi.org/10.1093/sysbio/49.2.257

    Macey, J.R., Schulte, J.A. II, Larson, A., Ananjeva, N.B., Wang, Y., Pethiyagoda, R., Rastegar-Pouyani, N. & Papenfuss, T. J. (2000b) Evaluating trans-Tethys migration: An example using acrodont lizard phylogenetics. Systematic Biology, 49 (2), 233–256.

    https://doi.org/10.1093/sysbio/49.2.233

    Macey, J.R., Schulte, J.A. II, Fong, J.J., Das, I. & Papenfuss, T.J. (2006) The complete mitochondrial genome of an agamid lizard from the Afro-Asian subfamily Agaminae and the phylogenetic position of Bufoniceps and Xenagama. Molecular Phylogenetics and Evolution, 39 (3), 881–886.

    https://doi.org/10.1016/j.ympev.2005.08.020

    Maddison, W.P. & Maddison, D.R. (2000) MacClade4: Analysis of Phylogeny and Character Evolution. Sinauer Associates, Sunderland, MA.

    Maniatis, T.E., Fritsch, F. & Sambrook, J. (1982) Molecular Cloning: A Laboratory Manual. Cold Spring Harbor Laboratory, Cold Spring Harbor, NY, 545 pp.

    McIntrye, A., Moore, T.C., Andersen, B., Balsam, W., Bé, A., Brunner, C., Cooley, J., Crowley, T., Denton, G., Gardner, J., Geitzenauer, K., Hays, J.D., Hutson, W., Imbrie, J., Irwing, G., Kellogg, T., Kennett, J., Kipp, N., Kukla, G., Kukla, H., Lozano, J., Luz, B., Mangion, S., Matthews, R.K., Mayewski, P., Molfino, B., Ninkovich, D., Opdyke, N., Prell, W., Robertson, J., Ruddiman, W.F., Sachs, H., Saito, T., Shackleton, N., Thierstein, H. & Thompson, P. (1976) The surface of the ice-age earth. Science, 191 (4232), 1131–1137.

    https://doi.org/10.1126/science.191.4232.1131

    Melnikov, D., Melnikova, E., Nazarov, R., Al-Johany, A. & Ananjeva, N.B. (2015) A new species of Phrynocephalus (Agamidae, Sauria) from Al Sharqiyah sands, northeastern Oman, dedicated to the memory of Sako Tuniyev (1983 – 2015). Russian Journal of Herpetology, 22 (4), 301–309.

    Melville, J., Hale, J., Mantziou, G., Ananjeva, N.B., Milto, K. & Clemann, N. (2009) Historical biogeography, phylogenetic relationships and intraspecific diversity of agamid lizards in the Central Asian deserts of Kazakhstan and Uzbekistan. Molecular Phylogenetics and Evolution, 53 (1), 99–112.

    https://doi.org/10.1016/j.ympev.2009.05.011

    Mertens, R. & Müller, L. (1928) Liste der Amphibien und Reptilien Europas. Abhandlungen der Senckenbergischen Naturforschenden Gesellschaft, 41 (1), 1–62.

    Metcalfe, I. (1996) Pre-Cretaceous evolution of SE Asia terranes. In: Hall, R. & Blundell, D. (Eds.), Tectonic Evolution of Southeast Asia. Geological Society of London Special Publication No. 106, London, pp. 97–122.

    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), 14 Nov. 2010, 1–8.

    https://doi.org/10.1109/GCE.2010.5676129

    Molnar, P., Burchfiel, B.C., Liang, K. & Zhao, Z. (1987) Geomorphic evidence for active faulting in the Altyn Tagh and northern Tibet and qualitative estimates of its contribution to the convergence of India and Eurasia. Geology, 15 (3), 249–253.

    https://doi.org/10.1130/0091-7613(1987)15<249:GEFAFI>2.0.CO;2

    Moritz, C., Dowling, T.E. & Brown, W.M. (1987) Evolution of animal mitochondrial DNA: Relevance for population biology and systematics. Annual Review of Ecology and Systematics, 18, 269–292.

    https://doi.org/10.1146/annurev.es.18.110187.001413

    Mouthereau, F., Lacombe, O. & Vergés, J. (2012) Building the Zagros collisional orogen: Timing, strain distribution and the dynamics of Arabia/Eurasia plate convergence. Tectonophysics, 532–535, 27–60.

    https://doi.org/10.1016/j.tecto.2012.01.022

    Murphy, R.W., Sites, J.W. Jr., Buth, D.G. & Haufler, C.H. (1990) Proteins I: Isozyme electrophoresis. In: Hillis, D.M. & Moritz, C. (Eds.), Molecular Systematics. Sinaur Associates, Sunderland, MA, pp. 45–126.

    Nikolsky, A.M. (1899) Reptiles and amphibians of the Turkestan general-governorship (Herpetologia Turanica). In: Fedschenko, A.P. Travel to the Turkestan of the member-founder of the A.P. Fedchenko Society, accomplished on behalf of the Imperial Society of Amateurs of natural history, anthropology, and ethnography. Zoogeographical Investigations. University Publishing, Moscow, Issue 23, Vol. 2, part 7, pp. 21–84. [in Russian]

    Nikolsky, A.M. (1905) Reptiles and amphibians of the Russian Empire (Herpetologica Rossica). Mémoires de l'Académie Impériale des Sciences de St.-Pétersbourg, 17 (1), 518 pp. [in Russian].

    Nikolsky A. M. (1916) Faunae de la Russie et des pays limitrophes fondée principalement sur les collections du Musée Zoologique de l’Académie Impériale des Sciences de Petrograd. Reptiles (Reptilia). Vol. II. Ophidia. Imperial Academy of Sciences, Petrograd. 349 pp. [in Russian with French title].

    Noble, D.W.A., Qi, Y. & Fu, J. (2010) Species delineation using Bayesian model-based assignment tests: A case study using Chinese toad-headed agamas (genus Phrynocephalus). BMC Evolutionary Biology, 2010 (10), 197.

    https://doi.org/10.1186/1471-2148-10-197

    Pallas, P.S. (1771) Reise durch verschiedene Provinzen des Russischen Reichs. Bd.I. St.Petersburg: Kayserliche Acade-mie der Wissenschaften, 540 pp.

    Pang, J., Wang, Y., Zhong, Y., Hoelzel, A.R., Papenfuss, T.J., Zeng, X., Ananjeva, N.B. & Zhang, Y.-P. (2003) A phylogeny of Chinese species in the genus Phrynocephalus (Agamidae) inferred from mitochondrial DNA sequences. Molecular Phylogenetics and Evolution, 27 (3), 398–409.

    https://doi.org/10.1016/S1055-7903(03)00019-8

    Qi, Y., Yang, W., Lu, B. & Fu, J. (2013) Genetic evidence for male-biased dispersal in the Qinghai toad-headed agamid Phrynocephalus vlangalii and its potential link to individual social interactions. Ecology and Evolution, 3 (5), 1219–1230.

    https://doi.org/10.1002/ece3.532

    Richardson, B.J., Baverstock, P.R. & Adams, M. (1986) Allozyme Electrophoresis. A Handbook for Animal Systematics and Population Studies. Academic Press, Sydney, 410 pp.

    Richter, B. & Fuller, M. (1996) Palaeomagnetism of the Sibumasu and Indochina blocks: Implications for the extrusion tectonic model. In: Hall, R. & Blundell, D. (Eds.), Reconstructing Cenozoic SE Asia. Geological Society of London Special Publication No. 106, London, pp. 203–224.

    https://doi.org/10.1144/GSL.SP.1996.106.01.13

    Royden, L.H., Burchfiel, B.C. & van der Hilst, R.D. (2008) The Geological Evolution of the Tibetan Plateau. Science, 321 (5892), 1054–1058.

    https://doi.org/10.1126/science.1155371

    Sattarov, T.S. (1993) Reptiles of North Tajikistan. Donish, Dushanbe, 276 pp. [in Russian]

    Selander, R.K., Smith, H.M., Yang, S.Y., Johnson, W.E. & Gentry, J.R. (1971) Biochemical polymorphism and systematics in the genus Peromyscus. I. Variation in the old-field mouse (Peromyscus polionotus). Studies in Genetics VI, University of Texas Publication, 7103, 49–90.

    Sengör, A.M.C. (1984) The Cimmeride orogenic system and the tectonics of Eurasia. Geological Society of America, Special Paper No. 195, 1–82.

    https://doi.org/10.1130/SPE195-p1

    Sengör, A.M.C., Altiner, D., Cin, A.,Ustaomer, T. & Hsu, K.J. (1988) Origin and assembly of the Tethyside orogenic collage at the expense of Gondwana Land. In: Audley-Charles, M.G. & Hallam, A. (Eds.), Gondwana and the Tethys. Geological Society Special Publication No. 37, Oxford University Press, Oxford, pp. 119–181.

    Schulte, J.A. II, Macey, J.R., Larson, A. & Papenfuss, T.J. (1998) Molecular tests of phylogenetic taxonomies: A general procedure and example using four subfamilies of the lizard family Iguanidae. Molecular Phylogenetics and Evolution, 10 (3), 367–376.

    https://doi.org/10.1006/mpev.1998.0541

    Shammakov, S. (1981) Reptiles of the Turkmenian plains. Ilym, Ashkhabad, 312 pp. [in Russian]

    Sharma, R.C. (1978) A new species of Phrynocephalus Kaup (Reptilia: Agamidae) from the Rajasthan Desert, India with notes on its ecology. Bulletin of the Zoological Survey of India, 1 (3), 291–294.

    Shackleton, R.M. & Chang, C. (1988) Cenozoic uplift and deformation of the Tibetan Plateau: The geomorphological evidence. In: The Geological Evolution of Tibet. Report of the 1985 Royal Society – Academia Sinica Geotraverse of the Qinghai – Xizang Plateau Led by Chang Chengfu, Robert M. Shackleton, F.R.S, John F. Dewey, F.R.S., and Yin Jixiang. Philosophical Transactions of the Royal Society of London, London, A 327 (1594), pp. 365–377.

    https://doi.org/10.1098/rsta.1988.0134

    Shenbrot, G.I. & Semenov (as Semyonov), D.V. (1990) New species of the genus Phrynocephalus (Reptilia, Agamidae) from Turkmenia. Zoologichesky Zhurnal, Moscow, 69 (9), 154–156. [in Russian with English summary]

    Smit, J.H.W., Cloetingh, S.A.P.L., Burov, E., Tesauro, M., Sokoutis, D. & Kaban, M. (2013) Interference of lithospheric folding in western Central Asia by simultaneous Indian and Arabian plate indentation. Tectonophysics, 602, 176–193.

    https://doi.org/10.1016/j.tecto.2012.10.032

    Solovyeva, E.N., Poyarkov, N.A., Dunaev, E.A., Duysebayeva, T.N. & Bannikova, A.A. (2011) Molecular differentiation and taxonomy of the Sunwatcher Toad-Headed Agama species complex Phrynocephalus superspecies helioscopus (Pallas 1771) (Reptilia: Agamidae). Russian Journal of Genetics, 47 (7), 842–856.

    https://doi.org/10.1134/S1022795411070155

    Solovyeva, E.N., Dunayev, E.A. & Poyarkov, N.A. (2012) Interspecific taxonomy of Sunwatcher Toadhead Agama species complex (Phrynocephalus helioscopus, Squamata). Zoologicheski Zhurnal, Moscow, 91 (11), 1377–1396. [in Russian with English summary]

    Solovyeva, E.N., Poyarkov, N.A., Dunayev, E.A., Nazarov, R.A., Lebedev, V.S. & Bannikova, A.A. (2014) Phylogenetic relationships and subgeneric taxonomy of Toad Headed Agamas Phrynocephalus (Reptilia, Squamata, Agamidae) as determined by mitochondrial DNA sequencing. Doklady Biological Sciences, 455, 119–124.

    https://doi.org/10.7868/S0869565214100260

    Stamatakis, A., Hoover, P. & Rougemont, J. (2008) A rapid bootstrap algorithm for the RAxML web servers. Systematic Biology, 57 (5), 758–771.

    https://doi.org/10.1080/10635150802429642

    Staub, N.L., Brown, C.W. & Wake, D.B. (1995) Patterns of growth and movements in a population of Ensatina eschscholtzii platensis (Caudata: Plethodontidae) in the Sierra Nevada, California. Journal of Herpetology, 29 (4), 593–599.

    https://doi.org/10.2307/1564743

    Steininger, F.F. & Rögl, F. (1984) Paleogeography and palinspastic reconstruction of the Neogene of the Mediterranean and Paratethys. In: Dixon, J.E. & Robertson, A.H.F. (Eds.), The Geological Evolution of the Eastern Mediterranean. Geological Society Special Publication No. 17, Blackwell Science, Oxford, pp. 659–668.

    https://doi.org/10.1144/GSL.SP.1984.017.01.52

    Strauch, A.A. (1876) Part III. Reptilia and Amphibia. In: Imperial Russian Geographic Society (Ed.), Przevalsky N. M., Mongolia and the Tangut Lands, a Three-Year Journey in Eastern High Asia. Vol. II. Reptiles and Amphibians. V.S. Balashov printing house, St. Petersburg, pp. 1–55. [in Russian]

    Swofford, D.L. (2002) PAUP*. Phylogenetic Analysis Using Parsimony (*and Other Methods). Sinauer Associates, Sunderland, Massachusetts.

    Tapponier, P., Mattauer, M., Proust, F. & Cassaigneau, C. (1981) Mesozoic ophiolites, sutures, and large-scale tectonic movements in Afghanistan. Earth and Planetary Science Letters, 52 (2), 355–371.

    https://doi.org/10.1016/0012-821X(81)90189-8

    Templeton, A.R. (1983) Phylogenetic inference from restriction endonuclease cleavage site maps with particular reference to the evolution of humans and the apes. Evolution, 37 (2), 221–244.

    https://doi.org/10.1111/j.1558-5646.1983.tb05533.x

    Townsend, T., Larson, A., Louis, E. & Macey, J.R. (2004) A new hypothesis of squamate evolutionary relationships from nuclear and mitochondrial DNA sequence data. Systematic Biology, 53 (5), 735–757.

    https://doi.org/10.1080/10635150490522340

    Tsarevsky (as Carevskij), S. (1926) Contributions sur la systématique et la distribution des lézards du genre Phrynocephalus (Reptilia). Comptes Rendus de l’Académie des Sciences de l’URSS, 1926, 211–214. [in Russian with French title]

    Urquhart, J., Wang, Y. & Fu, J. (2009) Historical vicariance and male-mediated gene flow in the toad-headed lizards Phrynocephalus przewalskii. Molecular Ecology, 18 (17), 3714–3729.

    https://doi.org/10.1111/j.1365-294X.2009.04310.x

    Wake, D.B. & Schneider, C.J. (1998) Taxonomy of the plethodontid salamander genus Ensatina. Herpetologica, 54 (2), 279–298.

    Wang, Y. & Fu, J. (2004) Cladogenesis and vicariance patterns in the toad-headed lizard Phrynocephalus versicolor species complex. Copeia, 2004 (2), 199–206.

    https://doi.org/10.1643/CG-03-082R1

    Wagner, P., Bauer, A.M., Leviton, A.E., Wilms, T.M. & Böhme, W. (2016) A checklist of the amphibians and reptiles of Afghanistan exploring herpetodiversity using biodiversity archives. Proceedings of the California Academy of Sciences, Series 4, 63, 457–565.

    Weisrock, D.W., Macey, J.R., Ugurtas, I.H., Larson, A. & Papenfuss, T.J. (2001) Molecular phylogenetics and historical biogeography among salamandrids of the “true” salamander clade: Rapid branching of numerous highly divergent lineages with the rise of Anatolia in Mertensiella luschani. Molecular Phylogenetics and Evolution, 18 (3), 434–448.

    https://doi.org/10.1006/mpev.2000.0905

    Weisrock, D.W., Macey, J.R., Matsui, M., Mulcahy, D.G. & Papenfuss, T.J. (2013) Molecular phylogenetic reconstruction of the endemic Asian salamander family Hynobiidae (Amphibia, Caudata). Zootaxa, 3626, 77–93.

    https://doi.org/10.11646/zootaxa.3626.1.3

    Windley, B.F. (1988) Tectonic framework of the Himalaya, Karakorum and Tibet, and problems of their evolution. In: Shackleton, R.M., Dewey, J.F. & Windley, B.F. (Eds.), Tectonic Evolution of the Himalayas and Tibet. Philosophical Transactions of the Royal Society of London, Series A, Vol. 326, No. 1589, Cambridge University Press, Cambridge, pp. 3–16.

    Wu, Y., Fu, J., Yue, B. & Yin, Q. (2015) An atypical reproductive cycle in a common viviparous Asia Agamid Phrynocephalus vlangalii. Ecology and Evolution, 5 (21), 5138–5147.

    https://doi.org/10.1002/ece3.1783

    Zhao, K. (1979) A survey of the classification and distribution of the toad-headed agamids (Phrynocephalus) in China. Acta Scientiarum Naturalium Universitatis Intramongolicae, Hohhot, 1979 (2), 111–121. [in Chinese with English summary]