Skip to main content Skip to main navigation menu Skip to site footer
Type: Articles
Published: 2013-01-08
Page range: 301–324
Abstract views: 33
PDF downloaded: 2

A preliminary phylogeny of the Palearctic naked-toed geckos (Reptilia: Squamata: Gekkonidae) with taxonomic implications

Department of Biology, Villanova University, 800 Lancaster Avenue, Villanova, Pennsylvania 19085, USA
Zoological Sciences Division, Pakistan Museum of Natural History, Garden Avenue, Shakarparian, Islamabad 44000, Pakistan
Department of Biology, Villanova University, 800 Lancaster Avenue, Villanova, Pennsylvania 19085, USA
Department of Natural Sciences, University of Michigan-Dearborn,125 Science Building, 4901 Evergreen Road, Dearborn, Michigan 48128, USA Department of Biology, Villanova University, 800 Lancaster Avenue, Villanova, Pennsylvania 19085, USA
Department of Biology, Villanova University, 800 Lancaster Avenue, Villanova, Pennsylvania 19085, USA
Department of Biology, Villanova University, 800 Lancaster Avenue, Villanova, Pennsylvania 19085, USA
Reptilia Cyrtopodion Tenuidactylus Mediodactylus Indogekko Altigekko Altiphylax Siwaligekko Asiocolotes Bunopus Agamura Crossobamon Tropiocolotes Stenodactylus Pseudoceramodactylus Cyrtodactylus Microgecko Alsophylax biogeography

Abstract

Palearctic naked-toed geckos are a group of gekkonid geckos that range from North Africa to northern India and western China, with their greatest diversity in Iran and Pakistan. Relationships among the constituent genera remain incompletely resolved and the monophyly of key genera remains unverified. Further, competing classifications are in current use and many species have been allocated to different genera by different authors. We used both mitochondrial (ND2) and nuclear genes (RAG1, PDC) to explore relationships among representatives of all but one genus in the group (Rhinogecko), including four genera not previously included in phylogenetic analyses (Asiocolotes, Altigekko, Indogekko, and Siwaligekko). Siwaligekko (and presumably other Tibeto-Himalayan species often referred to Cyrtopodion) are more closely related to tropical Asian Cyrtodactylus than to Palearctic naked-toed geckos. Sampled species of Asiocolotes and Altigekko are sister taxa, but both genera are here considered junior subjective synonyms of Altiphylax. Cyrtopodion sensu lato is non-monophyletic; Mediodactylus and Tenuidactylus, which have variably been considered as subgenera or synonyms of Cyrtopodion are both valid genera. Indogekko is embedded within Cyrtopodion and is here treated as a subgenus. Bunopus and Crossobamon are closely related to one-another, and with Agamura are interdigitated among taxa previously assigned to Cyrtopodion. Our data confirm the previous identification of a Saharo-Arabian Stenodactylus/Tropiocolotes/Pseudoceramodactylus clade and verify that Microgecko and Alsophylax are not members of the main clade of Palearctic naked-toed geckos. Osteological differences between Tropiocolotes and Microgecko, formerly treated as congeneric, are discussed and illustrated. The divergence between Cyrtodactylus and the Palearctic naked-toed clade predates the initial collision of the Indian and Eurasian plates, but deeper divergences within both groups are consistent with mountain building in the Himalayas and adjacent ranges as promoting cladogenic events. Miocene divergences within Tenuidactylus are consistent with vicariant speciation caused by uplift events in the Iranian and Transcaspian regions. Taxonomic implications of our phylogenetic results are discussed and a preliminary allocation of all species of padless Palearctic gekkonids to genus is provided.

References

  1. Agarwal, I. (2009) First record of Tropiocolotes persicus euphorbiacola Minton, Anderson and Anderson 1970 (Sauria: Gekkonidae) from the Republic of India, with notes on its habitat and natural history. Russian Journal of Herpetology, 16, 83–87.

    Ahmadzadeh, F., Flecks, M., Torki, F. & Böhme, W. (2011) A new species of angular-toed gecko, genus Cyrtopodion (Squamata: Gekkonidae) from southern Iran. Zootaxa, 2924, 22–32.

    Anderson, S.C. (1999) The Lizards of Iran. Society for the Study of Amphibians and Reptiles, Ithaca, New York, vii + 442 pp.

    Annandale, N. (1913). The Indian geckos of the genus Gymnodactylus. Records of the Indian Museum, 9, 309–326.

    Arévalo, E., Davis, S.K. & Sites, J.W., Jr. (1994) Mitochondrial DNA sequence divergence and phylogenetic relationships among eight chromosome races of the Sceloporus grammicus complex (Phrynosomatidae) in central Mexico. Systematic Biology, 43, 387–418.

    Arnold, E.N. (1980) The reptiles and amphibians of Dhofar, Southern Arabia. Journal of Oman Studies, Special Report 2, 273–332.

    Auffenberg, K., Krysko, K.L. & Auffenberg, W. (2004) Studies on Pakistan lizards: Cyrtopodion stoliczkai (Steindachner, 1867) (Gekkonidae: Gekkoninae). Asiatic Herpetological Research, 10, 151–160.

    Auffenberg, K., Krysko, K.L. & Rehman, H. (2010) Studies on Pakistan lizards: Cyrtopodion baturense (Khan and Baig 1992) and Cyrtopodion walli (Ingoldby 1922) (Sauria: Gekkonidae). Zootaxa, 2636, 1–20.

    Baig, K.J. (1998) A new species of Tenuidactylus (Sauria: Gekkonidae) from Balochistan, Pakistan. Hamadryad, 23, 127–132.

    Bauer, A.M., de Silva, A., Greenbaum, E. & Jackman, T.R. (2007) A new species of day gecko from high elevation in Sri Lanka, with a preliminary phylogeny of Sri Lankan Cnemaspis (Reptilia: Squamata: Gekkonidae). Mitteilungen aus dem Museum für Naturkunde in Berlin. Zoologische Reihe, 83, 22–32. http://dx.doi.org/10.1002/mmnz.200600022

    Blanford, W.T. (1874) Descriptions of new lizards from Persia and Baluchistan. Annals and Magazine of Natural History, series 4, 13, 453–455.

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

    Böhme, W. (1985) Zur Nomenklatur der paläarktischen Bogenfingergeckos, Gattung Tenuidactylus Ščerbak & Golubew, 1984 (Reptilia: Gekkonidae). Bonner zoologische Beiträge, 36, 95–98.

    Britton, T., Anderson, C. L., Jacquet, D., Lundqvist, S. & Bremer, K. (2007). Estimating divergence times in large phylogenetic trees. Systematic Biology, 56, 741–752. http://dx.doi.org/10.1080/10635150701613783

    Červenka, J. & Kratochvíl, L. (2010) Generic reassignment and validity of the recently described species Cyrtopodion dehkaroense. Herpetology Notes, 3, 135–137.

    Červenka, J., Frynta, D. & Kratochvíl, L. (2010) Phylogeny and taxonomy of the Middle Eastern geckos of the genus Cyrtopodion and their selected relatives. Zootaxa, 1931, 25–36.

    Červenka, J., Kratochvíl, L. & Frynta, D. (2008) Phylogenetic relationships of the gecko genus Carinatogecko (Reptilia: Gekkonidae). Zootaxa, 2636, 59–64.

    Clark, J., Clark, E.D., Anderson, S.C. & Leviton, A.E. (1969) Report on a collection of amphibians and reptiles from Afghanistan. Proceedings of the California Academy of Sciences, 4th series, 36, 279–315

    Das, I., Dattagupta, B. & Gayen, N.C. (1998) History and catalogue of reptile types in the collection of the Zoological Survey of India. Journal of South Asian Natural History, 3, 121–172.

    Daza, J.D. (2008) Cladistic analysis of the Gekkota (Reptilia) by means of craniological data. Unpublished Ph.D. dissertation, University of Puerto Rico, San Juan, Puerto Rico, USA, xvi + 227 pp.

    Drummond, A.J. & Rambaut, A. (2007) BEAST: Bayesian evolutionary analysis by sampling trees. BMC Evolutionary Biology 7, 214. http://dx.doi.org/10.1186/1471-2148-7-214

    Feng, J., Han, D., Bauer, A.M. & Zhou, K. (2007) Interrelationships among gekkonid geckos inferred from mitochondrial and nuclear gene sequences. Zoological Science, 24, 656–665. http://dx.doi.org/10.2108/zsj.24.656

    Fitzinger, L.J. (1826) Neue Classification der Reptilien nach ihren natürlichen Verwandschaften, nebst einer Verwandschafts-tafel und einem Verzeichnisse der Reptilien-Sammlung des K. K. Zoologischen Museum’s zu Wien. J. G. Heubner, Wien, [1], [1],[4], pp. 66, 1 folding table.

    Fitzinger, L.J. (1843) Systema Reptilium (Amblyglossae). Braumüller et Seidel, Vindobonae [Vienna], 106 pp. + vi, 3 pp. index (unnumbered).

    Fujita, M.K. & Papenfuss, T.J. (2011). Molecular systematics of Stenodactylus (Gekkonidae), an Afro-Arabian gecko species complex. Molecular Phylogenetics and Evolution, 58, 71–75. http://dx.doi.org/10.1016/j.ympev.2010.10.014

    Gamble, T. Bauer, A.M., Colli, G.R., Greenbaum, E., Jackman, T.R., Vitt, L.J. & Simons, A.M. (2011) Coming to America: multiple origins of New World geckos. Journal of Evolutionary Biology, 24, 231–244. http://dx.doi.org/10.1111/j.1420-9101.2010.02184.x

    Gamble, T., Greenbaum, E., Russell, A.P., Jackman, T.R. & Bauer, A.M. (2012) Repeated origin and loss of toepads in geckos. PLoS ONE, 7(6), e39429. Doi:10.1371/journal.pone.0039429. http://dx.doi.org/10.1371/journal.pone.0039429

    Golubev, M. L. (1984) The structure of the genus Tropiocolotes (Reptilia, Gekkonidae) [in Russian]. Vestnik Zoologii, 1984(6), 12.

    Golubev, M.L. & Szczerbak, N.N. (1981) Carinatogecko gen. n. (Reptilia, Gekkonidae) — a new genus from south-west Asia [in Russian]. Vestnik Zoologii, 1981(5), 34–41.

    Golubev, M.L., Khan, M.S. & Anderson, S.C. (1995) On the systematics of some Palearctic geckos. Abstracts, Second Asian Herpetological Meeting, 6–10 September, 1995, Ashgabat, Turkmenistan, 23–24.

    Gray, J.E. (1827) A synopsis of the genera of saurian reptiles, in which some new genera are indicated, and the others reviewed by actual examination. Philosophical Magazine, 2, 54–58.

    Greenbaum, E., Bauer, A.M., Jackman, T.R., Vences, M. & Glaw, F. (2007) A phylogeny of the enigmatic Madagascan geckos of the genus Uroplatus. Zootaxa, 1493, 41–51.

    Groth, J.G. & Barrowclough, G.F. (1999). Basal divergences in birds and the phylogenetic utility of the nuclear RAG-1 gene. Molecular Phylogenetics and Evolution, 12, 115–123. http://dx.doi.org/10.1006/mpev.1998.0603

    Guindon, S. & Gascuel, O. (2003). A simple, fast, and accurate algorithm to estimate large phylogenies by maximum likelihood. Systematic Biology, 52, 696–704. http://dx.doi.org/10.1080/10635150390235520

    Haas, G. (1957). Some amphibians and reptiles from Arabia. Proceedings of the California Academy of Sciences, 4th series, 29, 47–86.

    Haas, G. & Battersby, J.C. (1959). Amphibians and reptiles from Arabia. Copeia, 1959, 196–202. http://dx.doi.org/10.2307/1440386

    Han, D., Zhou, K. & Bauer, A.M. (2004) Phylogenetic relationships among the higher taxonomic categories of gekkotan lizards inferred from C-mos nuclear DNA sequences. Biological Journal of the Linnean Society, 83, 353–368. http://dx.doi.org/10.1111/j.1095-8312.2004.00393.x

    Hearn, T.M. & Ni, J.F. (1994) Pn velocities beneath continental collision zones: the Turkish-Iranian Plateau. Geophysical Journal International, 117, 273–283. http://dx.doi.org/10.1111/j.1365-246X.1994.tb03931.x

    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. http://dx.doi.org/10.1111/j.1463-6409.2011.00495.x

    Huang, B., Piper, J.D.A., Peng, S., Liu, T., Li, Z., Wang, Q. & Zhu, R. (2006) Magnetostratigraphic study of the Kuche Depression, Tarim Basin, and Cenozoic uplift of the Tien Shan Range, western China. Earth and Planetary Science Letters, 251, 346–364. http://dx.doi.org/10.1016/j.epsl.2006.09.020

    Jeremčenko, V.K. & Szczerbak, N.N. (1984) A new gecko lizard species (Reptilia, Gekkonidae) from Tien-Shan [in Russian]. Vestnik Zoologii, 1984(2), 46–50.

    Khan, M.S. (1988) A new cyrtodactylid gecko from northwestern Punjab, Pakistan. Journal of Herpetology, 22, 241–243. http://dx.doi.org/10.2307/1564007

    Khan, M.S. (1989) Rediscovery and redescription of the highland ground gecko Tenuidactylus montiumsalsorum (Annandale, 1913). Herpetologica, 45, 46–54.

    Khan, M.S. (1991) A new Tenuidactylus gecko from the Sulaiman Range, Punjab, Pakistan. Journal of Herpetology, 25, 199–204. http://dx.doi.org/10.2307/1564648

    Khan, M.S. (1993a) A new angular-toed gecko from Pakistan, with remarks on the taxonomy and a key to the species belonging to genus Cyrtodactylus (Reptilia: Sauria: Gekkonidae). Pakistan Journal of Zoology, 25, 67–73.

    Khan, M.S. (1993b). A new sandstone gecko from Fort Munro, Dera Ghazi Khan District, Punjab, Pakistan. Pakistan Journal of Zoology, 25, 217–221.

    Khan, M.S. (1993c) A checklist and key to the gekkonid lizards of Pakistan. Hamadryad, 18, 35–41.

    Khan, M.S. (1994) Validity and redescription of Tenuidactylus yarkandensis (J. Anderson, 1872). Pakistan Journal of Zoology, 26, 139–143.

    Khan, M.S. (1997) Biodiversity of geckkonid [sic] fauna of Pakistan. In: Mufti, S.A., Woods, C.A. & Hasan, S.A. (eds.), Biodiversity of Pakistan. Florida Museum of Natural History, Gainesville, pp. 383–389.

    Khan, M.S. (2000) Redescription and generic redesignation of Gymnodactylus stoliczkai Steindachner, 1869. Pakistan Journal of Zoology, 32, 157–163.

    Khan, M.S. (2001) Taxonomic notes on angular-toed Gekkota of Pakistan, with description of a new species of genus Cyrtopodion. Pakistan Journal of Zoology, 33, 13–24.

    Khan, M.S. (2003a) Anmerkungen zur Morphologie, Verbreitung und den Habitatpräferenzen einiger pakistanischer Geckos. Sauria, 25, 35–47.

    Khan, M.S. (2003b) Notes on circum Indus geckos of genus Cyrtopodion (Squamata: Gekkonidae). Gekkota 4, 43–59.

    Khan, M.S. (2003c) Questions of generic designation of angular-toed geckos of Pakistan with descriptions of three new genera (Reptilia: Gekkonidae). Journal of Natural History and Wildlife, 2, 1–9.

    Khan, M.S. (2004) Notes on high altitude geckos of the genus Altigekko in northeastern Pakistan. Bulletin of the Chicago Herpetological Society, 39, 221–224.

    Khan, M.S. (2005) Notes on geckos of the genus Siwaligekko Khan, 2003 (Reptilia: Gekkonidae) in northeastern Pakistan. Bulletin of the Chicago Herpetological Society, 40, 54–57.

    Khan, M.S. (2006) Amphibians and Reptiles of Pakistan. Krieger Publishing Company, Malabar, Florida, xvi + 311pp.

    Khan, M.S. (2008a) Biology and distribution of geckos of genus Indogekko Khan, 2003 (Sauria: Gekkonidae). Russian Journal of Herpetology, 15, 87–92.

    Khan, M.S. (2008b). Review of the morphology, ecology, and distribution of geckos of the genus Cyrtopodion, with a note on generic placement of Cyrtopodion brachykolon Krysko et. al., 2007. Caspian Journal of Environmental Sciences, 6, 79–86.

    Khan, M.S. (2009) Intergeneric relations of the angular-toed geckos of Circum Western Himalayas (Sauria: Gekkonidae). Pakistan Journal of Zoology, 41, 29–34.

    Khan, M.S. & Baig, K.J. (1992) A new Tenuidactylus gecko from northeastern Gilgit Agency, north Pakistan. Pakistan Journal of Zoology, 24, 273–277.

    Khan, M.S. & Rösler, H. (1999) Redescription and generic redesignation of the Ladakhian gecko Gymnodactylus stoliczkai Steindachner, 1969 [sic]. Asiatic Herpetological Research, 8, 60–68.

    Khan, M.S. & Tasnim, R. (1990) A new gecko of the genus Tenuidactylus from northeastern Punjab, Pakistan, and southwestern Azad Kashmir. Herpetologica, 46, 142–148.

    Kluge, A.G. (1967) Higher taxonomic categories of gekkonid lizards and their evolution. Bulletin of the American Museum of Natural History, 135, 1–60, pls. 1–5.

    Kluge, A.G. (1983) Cladistic relationships among gekkonid lizards. Copeia, 1983, 465–475. http://dx.doi.org/10.2307/1444392

    Kluge, A.G. (1985) Notes on gecko nomenclature (Sauria: Gekkonidae). Zoologische Mededelingen, 59, 95–100.

    Kluge, A.G. (1987) Cladistic relationships in the Gekkonoidea (Squamata, Sauria). Miscelaneous Publications of the Museum of Zoology, University of Michigan, (173), i–iv,1–54.

    Kluge, A.G. (1991) Checklist of gekkonid lizards. Smithsonian Herpetological Information Service, (85), 1–35.

    Kluge, A. G. (1993) Gekkonoid Lizard Taxonomy. International Gecko Society, San Diego, 245 pp.

    Kluge, A.G. (2001) Gekkotan lizard taxonomy. Hamadryad, 26, 1–209.

    Krysko, K.L., Rehman, H. & Auffenberg, K. (2007) A new species of Cyrtopodion (Gekkonidae: Gekkoninae) from Pakistan. Herpetologica, 63, 100–113. http://dx.doi.org/10.1655/0018-0831(2007)63[100:ANSOCG]2.0.CO;2

    Kuhl, H. & van Hasselt, J.C. (1822). Uittreksels uit brieven van de Heeren Kuhl en van Hasselt aan de Heeren C. J. Temminck, Th. van Swinderen en W. de Haan. Algemeene Konst- en Letter-Bode, 1822(1), 99–103.

    Leviton, A.E. & Anderson, S.C. (1963) Third contribution to the herpetology of Afghanistan. Proceedings of the California Academy of Sciences, 4th series, 31, 329–339.

    Leviton, A.E. & Anderson, S.C. (1972) Description of a new species of Tropiocolotes (Reptilia: Gekkonidae) with a revised key to the genus. Occasional Papers of the California Academy of Sciences (96), 1–7.

    Leviton, A.E. & Anderson, S.C. (1984) Description of a new species of Cyrtodactylus from Afghanistan with remarks on the status of Gymnodactylus longipes and Cyrtodactylus fedtschenkoi. Journal of Herpetology, 18, 270–276. http://dx.doi.org/10.2307/1564080

    Li, P., Zhao, E. & Dong, B. (2010) Amphibians and Reptiles of Tibet [in Chinese]. Science Press, Beijing, xii + 251 pp.

    Macey, J.R., Larson, A., Ananjeva, N.B., Feng, 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, 91-–104. http://dx.doi.org/10.1093/oxfordjournals.molbev.a025706

    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, 320–332. http://dx.doi.org/10.1006/mpev.1999.0641

    Macey, J.R., Ananjeva, N.B. Wang, Y. & Papenfuss, T.J. (2000) Phylogenetic relationships among Asian gekkonid lizards formerly of the genus Cyrtodactylus based on cladistic analyses of allozymic data: monophyly of Cyrtopodion and Mediodactylus. Journal of Herpetology, 34, 258–265. http://dx.doi.org/10.2307/1565422

    Macey, J.R., Fong, J.J., Kuehl, J.V., ShaWei, S., Ananjeva, N.B., Papenfuss, T.J. & Boore, J.J. (2005) The complete mitochondrial genome of a gecko and the phylogenetic position of the Middle Eastern Teratoscincus keyserlingii. Molecular Phylogenetics and Evolution, 36, 188–193. http://dx.doi.org/10.1016/j.ympev.2005.03.025

    Maddison, D.R. & Maddison, W.P. (2005) MacClade 4: Analysis of phylogeny and character evolution. Version 4.08a. http://macclade.org.

    Masroor, R. (2009) A new arboreal species of Cyrtopodion (Squamata: Gekkonidae) from Deh Akro-II Wetlands Complex, Sindh, Pakistan. Zootaxa, 2243, 57–67 [erratum in Zootaxa, 2307, 68].

    Mees, G.F. (1987) On the generic names Gonydactylus Kuhl and van Hasselt, 1822, Goniodactylus Schlegel, 1826, and Gonydactylus Wagler, 1830 (Reptilia: Sauria: Gekkonidae). Zoologische Mededelingen, 61,137–140.

    Mertens, R. (1965) Bemerkungen über einige Eidechsen aus Afghanistan. Senckenbergiana Biologica, 46, 1–4.

    Minton, S.A., Jr. (1966) A contribution to the herpetology of West Pakistan. Bulletin of the American Museum of Natural History, 134, 27–184, pls. 9–36.

    Minton, S.A., Anderson, S.C. & Anderson, J.A. (1970) Remarks on some geckos from southwest Asia, with descriptions of three new forms and a key to the genus Tropiocolotes. Proceedings of the California Academy of Sciences, 4th series, 37, 333–362.

    Nazarov, R., Ananjeva, N. & Radjabizadeh, M. (2009) Two new species of angular-toed geckoes (Squamata: Gekkonidae) from South Iran. Russian Journal of Herpetology, 16, 311–324.

    Nikolsky, A. M. 1907. Reptiles et amphibies, recueillis par Mr. N. A. Zaroudny en Perse en 1903–1904. Annuaire du Musée Zoologique de l'Académie Impériale des Sciences de St.-Pétersbourg, 10, 260–301.

    Nylander, J., Wilgenbusch, J., Warren, D. & Swofford, D. (2008) AWTY A system for graphical exploration of MCMC convergence in Bayesian phylogenetic inference. http://ceb.csit.fsu.edu/awty.

    Perret, J.L. (1976) Garzoniella: Un nouveau genre de Gekkonidae saharien. Revue Suisse de Zoologie, 83,761–764.

    Peters, W. (1880) Über die von Hrn. Gerhard Rohlfs und Dr. A. Stecker auf der Reise nach der Oase Kufra gesammelten Amphibien. Monatsberichte der Königlich Preussischen Akademie der Wissenschaften zu Berlin, 1880, 305–309.

    Posada, D. (2008). jModelTest: Phylogenetic model averaging. Molecular Biology and Evolution, 25, 1253–1256. http://dx.doi.org/10.1093/molbev/msn083

    Rajabizadeh, M., Rastegar-Pouyani, N., Faizi, H. & Bostanchi, H. (2010) New insights inn the taxonomy of the Persian sand gecko, Tropiocolotes persicus (Nikolsky, 1903). Zoology in the Middle East, 49, 55–62.

    Rambaut, A. & Drummond, A.J. (2007) Tracer v1.4, available from http://beast.bio.edu.ac.uk/Tracer

    Ronquist, F. & Huelsenbeck, J.P. (2003) MrBayes 3: Bayesian phylogenetic inference under mixed models. Bioinformatics, 19, 1572–1574. http://dx.doi.org/10.1093/bioinformatics/btg180

    Rowley, D.B. (1996) Age of initiation of collision between India and Asia: A review of stratigraphic data. Earth and Planetary Science Letters, 145, 1–13. http://dx.doi.org/10.1016/S0012-821X(96)00201-4

    Shi, L. & Zhao, E. (2011) A new gecko in the genus Cyrtopodion Fitzinger, 1843 (Reptilia: Squamata: Gekkonidae) from Western China. Herpetologica, 67, 186–193. http://dx.doi.org/10.1655/09-036.1

    Shimodaira, H. & Hasegawa, M. (1999) Multiple comparisons of log-likelihoods with applications to phylogenetic inference. Molecular Biology and Evolution, 16,1114–1116. http://dx.doi.org/10.1093/oxfordjournals.molbev.a026201

    Sindaco, R. & Jeremčenko, V.K. (2008) The Reptiles of the Western Palearctic. Monografie della Societas Herpetologica Italica 1. Edizione Belvedere, Latina, Italy. 579 pp.

    Spix, J.B. (1825) Animalia Nova Sive Species Novae Lacertarum, quas in Itinere per Brasiliam Annis MDCCCXVII–MDCCCXX Jussu et Auspicius Maximiliani Josephi I. Bavariae Regis. Typis Franc. Seraph. Hübschmanni, Monachii [Munich], 26 + [4] pp., 30 pls. [I–XXVIII, XIIIa, XIVa].

    Stamatakis, A. (2006). RAxML-VI-HPC: Maximum Likelihood-based phylogenetic analyses with thousands of taxa and mixed models. Bioinformatics, 22, 2688–2690. http://dx.doi.org/10.1093/bioinformatics/btl446

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

    Szczerbak, N.N. & Golubev, M.L. (1977) Systematics of the Palearctic geckos (genera Gymnodactylus, Bunopus, Alsophylax) [in Russian]. Trudy Instituta Zoologii, Akademiya Nauk USSR, 74, 120–133.

    Szczerbak, N.N. & Golubev, M.L. (1984) On generic assignement of the Palearctic Cyrtodactylus lizard species (Reptilia, Gekkonidae) [in Russian]. Vestnik Zoologii, 1984(2), 50–56.

    Szczerbak, N.N. & Golubev, M.L. (1986) Gecko Fauna of the USSR and Adjacent Regions [in Russian]. Nauka Dymka, Kiev, ix + 232 pp, pls. 1–8.

    Szczerbak, N.N. & Golubev, M.L. (1996) Gecko Fauna of the USSR and Contiguous Regions. [English edition translated from the Russian by M.L. Golubev & S.A. Malinsky; Leviton, A.E. & Zug, G.R., eds.]. Society for the Study of Amphibians and Reptiles, Ithaca, New York, ix + 233 pp., 8 pls.

    Torki, F. (2008) Morphologische und ökologische Aspekte von Tropiocolotes helenae (Nikolsky, 1907) (Reptilia: Gekkonidae). Sauria, 30(1), 5–12.

    Torki, F. (2011) Description of a new species of Carinatogecko (Squamata: Gekkonidae) from Iran. Salamandra, 47, 103–111.

    Torki, F., Gharzi, A., Nazari-Serenjeh, F., Javanmardi, S. & Heidari, N. (2008) Geckos of the genera Tropiocolotes and Asaccus in the Zagros Mountains, Iran. Gekko, 5(2), 31–43.

    Underwood G. (1954) On the classification and evolution of geckos. Proceedings of the Zoological Society of London, 124, 469–492. http://dx.doi.org/10.1111/j.1469-7998.1954.tb07789.x

    Witte, G.F. de (1973) Description d'un Gekkonidae nouveau de l'Iran (Reptilia Sauria). Bulletin de Institut Royal des Sciences Naturelles de Belgique, Bruxelles, 49, 1–6.

    Wood, P.L., Jr., 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. http://dx.doi.org/10.1016/j.ympev.2012.08.025

    Zhao, E. & Li, S. (1987) A new lizard of Tenuidactylus and a new Tibetan snake record of Amphiesma [in Chinese]. Acta Herpetologica Sinica, 6, 48–51.