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Issue: Vol. 5369 No. 3: 10 Nov. 2023
Type: Article
Published: 2023-11-10
DOI: 10.11646/zootaxa.5369.3.2
Page range: 336-368
Abstract views: 973
PDF downloaded: 19

Taxonomic diversity of racerunners with descriptions of two new Eremias species (Sauria: Lacertidae) from Central Iran

Zoological Museum of Moscow State University; 125009; B. Nikitskaya 2; Moscow; Russia; Zoological Institute of the Russian Academy of Sciences; Universitetskaya nab. 1; St. Petersburg; 199034; Russia; Laboratory of Comparative Ethology and Biocommunication; Severtsov Institute of Ecology and Evolution; Russian Academy of Sciences.; 119071 Leninsky Prospect 33; Moscow; Russia
Faculty of Sciences; Hakim Sabzevari University; Sabzevar; Iran
Department of Biodiversity; Institute of Science and High Technology and Environmental Sciences; Graduate University of Advanced Technology; Kerman; Iran
Department of Biology; Faculty of sciences; University of Qom; Qom; Iran
Faculty of Biology; Lomonosov Moscow State University; Moscow 119991; Russia
Zoological Institute of the Russian Academy of Sciences; Universitetskaya nab. 1; St. Petersburg; 199034; Russia
Zoological Museum of Moscow State University; 125009; B. Nikitskaya 2; Moscow; Russia; Laboratory of Comparative Ethology and Biocommunication; Severtsov Institute of Ecology and Evolution; Russian Academy of Sciences.; 119071 Leninsky Prospect 33; Moscow; Russia
Reptilia cryptic diversity Eremias Iranian Plateau taxonomy cytochrome oxidase I DNA Barcoding

Abstract

We provide a diversity assessment of Iranian species of the genus Eremias based on the cytochrome oxidase I mtDNA gene fragment. We analyzed 93 genetic samples from the entire distribution of the Eremias fasciata species complex in Iran and surrounding regions, along with morphological data to support the description of two new species from Central Iran. We hypothesize that the diversification of the Eremias fasciata species complex was largely influenced by the fragmentation of sand massifs in the region. This same hypothesis has been used to explain the high level of endemism among the sand-dwelling species of reptiles along the Iranian Plateau in the same area. The two new species described herein can be distinguished from other congeneric species by their phylogenetic position and a combination of morphological characters. We use these data to discuss the taxonomy of Eremias based on morphology, habitat choice, and genetic data.

 

References

  1. Anderson, J. (1872) On some Persian, Himalayan, and other Reptiles. Proceedings of the Zoological Society of London, 1872, 371–404.
  2. Anderson, S.C. (1999) The Lizards of Iran. Society for the Study of Amphibians and Reptiles, Ithaca, New York, 442 pp.
  3. Arnold, E.N. (1986) The hemipenis of lacertid lizards (Reptilia: Lacertidae): structure, variation and systematic implications. Journal of Natural History, 20, 1221–1257. https://doi.org/10.1080/00222938600770811
  4. Arnold, E.N. (1989) Towards a phylogeny and biogeography of the Lacertidae: relationships within an Old-World family of lizards derived from morphology. Bulletin of the British Museum Natural History, Zoology, 55 (2), 209–257.
  5. Arnold, E.N., Arribas, O. & Carranza, S. (2007) Systematics of the Palearctic and Oriental lizard tribe Lacertini (Squamata: Lacertidae: Lacertinae), with descriptions of eight new genera. Zootaxa, 1430 (1), 3–86. https://doi.org/10.11646/zootaxa.1430.1.1
  6. Baig, K.J. & Masroor, R. (2006) A new species of Eremias (Sauria: Lacertidae) from Cholistan Desert, Pakistan. Russian Journal of Herpetology, 13 (3), 167–174.
  7. Barabanov, A.V. (2009) Aspidorhinus Eichwald, 1841 as a valid subgeneric name for Eremias velox species group (Sauria, Lacertidae). Current Studies in Herpetology, 9 (1–2), 59–61. [in Russian with English abstract]
  8. Bedriaga, J.V. (1912) Amphibians and Reptiles. Part 4 (Zoological Section III (1)). In: Nauchnye rezultaty puteshestviy N. M. Przewalskogo po Tsentralnoy Azii [Scientific results of N.M. Przewalskii’s Travels in Central Asia]. Imperial Academy, St. Petersburg, pp. 503–769, pls. VIII–X. [in Russian and German]
  9. Berberian, M. & King, G.C.P. (1981) Towards a paleogeography and tectonic evolution of Iran. Canadian Journal of Earth Sciences, 18 (2), 210–265. https://doi.org/10.1139/e81-019
  10. Böhme, W. & Szczerbak, N. (1991) Ein neuer Wüstenrenner aus dem Hochland Afghanistans, Eremias (Eremias) afghanistanica sp. n. (Reptilia: Sauria: Lacertidae). Bonner zoologische Beiträge, 42, 137–141.
  11. Boulenger, G.A. (1918) A synopsis of the lizards of the genus Eremias. Journal of Zoological Research, 3, 1–12.
  12. Boulenger, G.A. (1920) Monograph of the Lacertidae. Vol. 1. Trustees of the British Museum of Natural History, London, 352 pp. https://doi.org/10.5962/bhl.title.8312
  13. Boulenger, G.A. (1921) Monograph of the Lacertidae. Vol. 2. Trustees of the British Museum of Natural History, London, 352 pp. https://doi.org/10.5962/bhl.title.8312
  14. Blanford, W.T. (1874) Descriptions of new lizards from Persia and Baluchistàn. Annals and Magazine of Natural History, Series 4, 13 (4), 453–455. https://doi.org/10.1080/00222937408680903
  15. Blanford, W.T. (1875) List of Reptilia and Amphibia collected by the late Dr. Stoliczka in Kashmir, Ladak, eastern Turkestan, and Wakhan, with descriptions of new species. Journal of the Asiatic Society of Bengal, 44 (2), 191–196 + 201, 2 pls.
  16. Blanford, W.T. (1876) Eastern Persia. In: An Account of the Journeys of the Persian Boundary Commission, 1870-1872. Vol. 2. Zoology and Geology. Macmillan and Co., London, 516 pp.
  17. Brushko, Z.K. (1995) Lizards of Desert Regions of Kazakhstan. Almaty, Konzhik, 231 pp. [in Russian]
  18. Darevsky, I.S. & Shcherbak, N.N. (1978) Eremias andersoni, a new lizard (Reptilia, Lacertilia, Lacertidae) from Iran. Journal of Herpetology, 12 (1), 13–15. https://doi.org/10.2307/1563496.
  19. Eichwald, E. (1841) Fauna Caspio-Caucasia Nonnulis Observationibus Novis. Litteris Typographiae Diarii Gall. Politic. Petropol., Petropoli, 233 pp., 40 col. Pls. https://doi.org/10.5962/bhl.title.126375
  20. Eremchenko, V.K. (1999) Nomenclature of Asian racerunners Eremias Wiegmann, 1884 in relation with following designation of monotypy (Sauria: Lacertidae). Izvestiya Natsionalnoy Akademii Nauk Kyrgyzstana, 1, 72–73. [in Russian]
  21. Guo, X., Chen, D.X., Papenfuss, T., Ananjeva, N.B., Melnikov, D.A. & Wang, Y. (2011) Phylogeny and divergence times of some racerunner lizards (Lacertidae: Eremias) inferred from mitochondrial 16S rRNA gene segments. Molecular Phylogenetics and Evolution, 61 (2), 400–412. https://doi.org/10.1016/j.ympev.2011.06.022
  22. Fu, J.Z. (1998) Towards the phylogeny of the family Lacertidae: implications from mitochondrial DNA 12S and 16S gene sequences (Reptilia: Squamata). Molecular Phylogenetics and Evolution, 9 (1), 118–130. https://doi.org/10.1006/mpev.1997.0456
  23. Fu, J.Z. (2000) Towards the phylogeny of the family Lacertidae—why 4708 base pairs of mtDNA cannot draw the picture. Biological Journal of the Linnaean Society, 71, 203–217. https://doi.org/10.1111/j.1095-8312.2000.tb01254.x
  24. Ivanova, N.V., Dewaard, J.R. & Hebert, P.D.N. (2006) An inexpensive, automation-friendly protocol for recovering high-quality DNA. Molecular Ecology Notes, 6 (4), 998–1002. https://doi.org/10.1111/j.1471-8286.2006.01428.x
  25. Hall, T. (1999) BioEdit: A user-friendly biological sequence alignment editor and analysis program for Windows 95/98/NT. Nucleic Acids Symposium Series, 41, 95–98.
  26. Harris, D.J., Arnold, E.N. & Thomas, R.N. (1998) Relationships of lacertid lizards (Reptilia: Lacertidae) estimated from mitochondrial DNA sequences and morphology. Proceedings of the Royal Society of London B, 265, 1939–1948. https://doi.org/10.1098/rspb.1998.0524
  27. Hillis, D.M., Moritz, C. & Mable, B.K. (1996) Molecular Systematics. 2nd Edition. Sinauer Associates, Inc, Sunderland, Massachusetts, xvi + 655 pp. https://doi.org/10.2307/1447682
  28. Heidari, N., Rastegar Pouyani, N., Rastegar Pouyani, E. & Radjabizadeh, M. (2013) A new species of Acanthodactylus Fitzinger 1834 (Sauria: Lacertidae) from southern Iran. Zootaxa, 3722 (3), 333–346. https://doi.org/10.11646/zootaxa.3722.3.3
  29. Hebert, P.D.N. & Gregory, T.R. (2005) The promise of DNA barcoding for taxonomy. Systematic Biology, 54, 852–859. https://doi.org/10.1080/10635150500354886
  30. Huelsenbeck, J.P. & Hillis, D.M. (1993) Success of phylogenetic methods in the four-taxon case. Systematic Biology, 42, 247–264. https://doi.org/10.1093/sysbio/42.3.247
  31. Huelsenbeck, J.P., Ronquist, F. (2001) MrBayes: Bayesian inference of phylogeny. Bioinformatics, 17, 754–755. https://doi.org/10.1093/bioinformatics/17.8.754
  32. Kamali, K. & Anderson, S. (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
  33. Kazemi, S.M., Qomi, M.F., Kami, H.G. & Anderson, S.C. (2011) A new species of Ophiomorus (Squamata: Scincidae) from Maranjab Desert, Isfahan Province, Iran, with a revised key to the genus. Amphibian & Reptile Conservation, 5 (1), 23–33.
  34. Khan, M.A., Jablonski, D., Nadeem, M.S., Masroor, R., Kehlmaier, C., Spitzweg, C. & Fritz, U. (2021) Molecular phylogeny of Eremias spp. from Pakistan contributes to a better understanding of the diversity of racerunners. Journal of Zoological Systematics and Evolutionary Research, 59, 466–483. https://doi.org/10.1111/jzs.12426
  35. Lantz, L.A. (1928) Les Eremias de l’Asie occidentale. Bulletin du Musée de Géorgie, 4–5, 1–136 + (1), pls. I–III.
  36. Leviton, A.E., Anderson, S.C., Adler, K. & Minton, S.A. (1992) Handbook to Middle East Amphibians and Reptiles. Society for the Study of Amphibians and Reptiles, Oxford, Ohio, 252 pp.
  37. Masroor, R., Khisroon, M., Khan, M.A. & Jablonski, D. (2020) A new species of Eremias (Squamata: Lacertidae) from the arid mountains of Pakistan. Zootaxa, 4786 (1), 101–121. https://doi.org/10.11646/zootaxa.4786.1.8
  38. Masroor, R., Khan, M.A., Nadeem, M.S., Amir, S.A., Khisroon, M. & Jablonski, D. (2022) Appearances often deceive in racerunners: integrative approach reveals two new species of Eremias (Squamata: Lacertidae) from Pakistan. Zootaxa, 5175 (1), 55–87. https://doi.org/10.11646/zootaxa.5175.1.3
  39. Mayer, W. & Pavlicev, M. (2007) The phylogeny of the family Lacertidae (Reptilia) based on nuclear DNA sequences: convergent adaptations to arid habitats within the subfamily Eremiainae. Molecular Phylogenetics and Evolution, 44, 1155–1163. https://doi.org/10.1016/j.ympev.2007.05.015
  40. Mattei, M., Cifelli, F., Muttoni, G. & Rashid, H. (2015) Post-Cimmerian (Jurassic–Cenozoic) paleogeography and vertical axis tectonic rotations of Central Iran and the Alborz Mountains. Journal of Asian Earth Sciences, 102, 92–101. https://doi.org/10.1016/j.jseaes.2014.09.038
  41. Moravec, J. (1994) A new lizard from Iran, Eremias (Eremias) lalezharica sp. n. (Reptilia: Lacertilia: Lacertidae). Bonner zoologische Beiträge, 45 (1), 61–66.
  42. Mozaffari, O. & Parham, J.F. (2007) A new species of racerunner lizard (Lacertidae: Eremias) from Iran. Proceedings of the California Academy of Sciences, 58 (28), 569–574.
  43. Mozaffari, O., Ahmazadeh, F. & Parham, J.F. (2011) Eremias papenfussi sp. nov., a new lacertid lizard (Sauria: Lacertidae) from Tehran Province, Iran. Zootaxa, 3114 (1), 57–62. https://doi.org/10.11646/zootaxa.3114.1.6
  44. Mozaffari, O., Ahmazadeh, F. & Saberi-Pirooz, R. (2020) Fahimi’s racerunner, a new species of the genus Eremias Fitzinger, 1834 (Sauria: Lacertidae) from Iran. Zootaxa, 4768 (4), 565–578. https://doi.org/10.11646/zootaxa.4768.4.7
  45. Nazarov, R.A., Radjabizadeh, M., Poyarkov, N.A., Ananjeva, N.B., Melnikov, D.A. & Rastegar-Pouyani, E. (2017) A New Species of Frog-Eyed Gecko, genus Teratoscincus Strauch, 1863 (Squamata: Sauria: Sphaerodactylidae), from Central Iran. Russian Journal of Herpetology, 24 (4), 291–310. https://doi.org/10.30906/1026-2296-2019-24-4-291-310
  46. Nazarov, R.A., Melnikov, D.A., Radjabizadeh, M. & Poyarkov, N.A. (2018) A new species of short-fingered geckos Stenodactylus (Squamata, Geckonidae) from South Iran with taxonomic notes on validity of the genus Trigonodactylus Hass, 1957. Zootaxa, 4457 (1), 93–113. https://doi.org/10.11646/zootaxa.4457.1.4
  47. Nikolsky, A.M. (1897) Les reptiles, amphibiens et poissons recueillis (part.) Mr. N. Zaroudny dans la Perse orientale. Annuaire Musée Zoologique de l’Académie Impériale des Sciences de St.-Pétersbourg, 2, 306–348. [in Russian with French title]
  48. Nikolsky, A.M. (“1899” 1900) Reptiles, amphibiens et poissons, recueillis pendant le voyage de Mr. N. A. Zaroudny en 1898dans la Perse. Annuaire du Musée Zoologique de l’Académie Impériale des Sciences de St. Pétersbourg, 4 (4), 375–417. [in Russian with French title]
  49. Nikolsky, A.M. (1905) Herpetologia Rossica. Mémoires de l’Académie Imperiale des Sciences, Phys.-Math. Classe, St.-Pétersbourg, XVII (1), 1–518
  50. Nikolsky, A.M. (1915) The Fauna of Russia and Adjacent Countries. Reptiles (Reptilia). Vol 1. Chelonia and Sauria. Academy of Science, Petrograd, 534 pp. [in Russian]
  51. Orlova, V.F. & Terbish, Kh. (1997) Family Lacertidae Cope, 1864. In: Ananjeva, N.B., Munkhbayar, Kh., Orlov, N.L., Orlova, V.F., Semenov, D.V. & Terbish, Kh. Amphibians and Reptiles of Mongolia. Reptiles. KMK Ltd., Moscow, pp. 133–266. [in Russian]
  52. Orlova, V.F. & Nazarov, R.A. (2017) Diversity of racerunners (Sauria: Lacertidae, Eremias) of Iran: Review of species and taxonomy issues. In: Spatial-temporal Dynamics of Biota and Ecosystems in the Aral-Caspian Basin. Materials of the II International Conference, Dedicated to the Memory of the Outstanding Naturalist and Traveler Nikolai Alekseevich Zarudny. IPK “University”, Orenburg, pp. 298–302. [in Russian]
  53. Orlova, V.F., Poyarkov, N.A., Chirikova, M.A., Nazarov, R.A., Munkhbaatar, M., Munkhbayar, K. & Terbish, K. (2017) MtDNA Differentiation and Taxonomy of Central Asian Racerunners of Eremias multiocellata–E. przewalskii species complex (Squamata, Lacertidae). Zootaxa, 4282 (1), 1–42. https://doi.org/10.11646/zootaxa.4282.1.1
  54. Orlova, V.F., Solovyeva, E.N., Dunayev, E.A. & Ananjeva, N.B. (2022) Integrative taxonomy within Eremias multiocellata complex (Sauria, Lacertidae) from the western part of range: evidence from historical DNA. Genes, 13, 941. https://doi.org/10.3390/genes13060941
  55. Popov, S.V., Antipov, M.P., Zastrozhnov, A.S., Kurina, E.E. & Pinchuk, T.N. (2010) Sea‐level fluctuations on the northern shelf of the Eastern Paratethys in the Oligocene‐Neogene. Stratigraphy and Geological Correlation, 18 (2), 200–224.
  56. https://doi.org/10.1134/s0869593810020073
  57. Posada, D. & Crandall, K.A. (1998) MODELTEST: testing the model of DNA substitution. Bioinformatics, 14, 817–818. https://doi.org/10.1093/bioinformatics/14.9.817
  58. Poyarkov, N.A., Jr., Orlova, V.F. & Chirikova, M.A. (2014) The mitochondrial phylogeography and intraspecific taxonomy of the Steppe Racerunner, Eremias arguta (Pallas) (Lacertidae: Sauria, Reptilia), reflects biogeographic patterns in MiddleAsia. Zootaxa, 3895 (2), 208–224. https://doi.org/10.11646/zootaxa.3895.2.4
  59. Rastegar-Pouyani, N. & Rastegar-Pouyani, E. (2001) A new species of Eremias (Sauria: Lacertidae) from Highlands of Kermanshah Province, Western Iran. Asiatic Herpetological Research, 9, 107–112. https://doi.org/10.5962/bhl.part.15563
  60. Rastegar-Pouyani, N. & Rastegar-Pouyani, E. (2006) A new form of Eremias (Sauria: Lacertidae) from the Alvand Mountains, Hamedan Province, Western Iran. Iranian Journal of Animal Biosystematics (UAB), 1 (1), 14–20.
  61. Rastegar-Pouyani, E., Hosseinian, S., Rafiee, S., Kami, H.G., Rajabizadeh, M. & Wink, M. (2016) A new species of the genus Eremias Fitzinger, 1834 (Squamata: Lacertidae) from Central Iran, supported by mtDNA sequences and morphology. Zootaxa, 4132 (2), 207–220. https://doi.org/10.11646/zootaxa.4132.2.2
  62. Ronquist, F.& Huelsenbeck, J.P. (2003) MrBayes 3: Bayesian phylogenetic inference under mixed models. Bioinformatics, 19, 1572–1574. https://doi.org/10.1093/bioinformatics/btg180
  63. Safaei-Mahroo, B., Ghaffari, H., Fahami, H., Broomad, S., Yazdanian, M., Najafi Majd, E., Hosseinian Yousefkhani, S., Rezzadeh, E., Hosseinzadeh, M., Nasrabadi, R., Rajabizadeh, M., Mashayekhi, M., Motesharei, A., Naderi, A. & Kazemi, S. (2015) The herpetofauna of Iran: Checklist of taxonomy, distribution and conservation Status. Asian Herpetological Research, 6 (4), 257–290. https://doi.org/10.16373/j.cnki.ahr.140062
  64. Szczerbak, N.N. (1971) Taxonomy of the genus Eremias (Sauria, Reptilia) in connection with the focuses of the desert-steppe fauna development in Paleoarctic. Vestnik Zoologii, 2, 47–55. [in Russian]
  65. Szczerbak, N.N. (1974) Racerunners of the Palaearctic. Naukova Dumka, Kiev, 293 pp. [in Russian]
  66. Smith, A.M., Poyarkov, N.A. & Hebert, P.D.N. (2008) CO1 DNA barcoding amphibians: take the chance, meet the challenge. Molecular Ecology Notes, 8 (2), 235–246. https://doi.org/10.1111/j.1471-8286.2007.01964.x
  67. Solovyeva, E.N., Dunayev, E.N., Nazarov, R.A., Radjabizadeh, M. & Poyarkov, Jr. N.A. (2018) Molecular and morphological differentiation of Secret Toad-headed agama, Phrynocephalus mystaceus, with the description of a new subspecies from Iran (Reptilia, Agamidae). ZooKeys, 748, 97–129. https://doi.org/10.3897/zookeys.748.20507
  68. Strauch, A. (1863) Characteristik zweier neuen Eidechsen aus Persien. Bulletin de l’Académie Impériale des Sciences de St. Pétersbourg, 6, 477–480.
  69. Strauch, A.A. (1876) Reptiles and amphibians. In: Przewalski, N.M. (Ed.), Mongolia, the Tangut Country, and the Solitudes of Northern Tibet: Being a Narrative of Three-Years in Journey in Eastern High Asia. Vol. 2. Russian Geographical Society, St. Petersburg, pp. 1–55. [in Russian]
  70. Swofford, D. (2002) PAUP*. Phylogenetic Analysis Using Parsimony (*and Other Methods). Sinauer Associates, Sunderland, Massachusetts. [program]
  71. Tamar, K., Carranza, S., Sindaco, R., Moravec, J., Trape, J.F. & Meiri, S. (2016) Out of Africa: Phylogeny and biogeography of the wide­spread genus Acanthodactylus (Reptilia: Lacertidae). Molecular Phylogenetics and Evolution, 103, 6–18. https://doi.org/10.1016/j.ympev.2016.07.003
  72. Tamura, K., Peterson, D., Peterson, N., Stecher, G., Nei, M. & Kumar, S. (2011) MEGA5: Molecular Evolutionary Genetics Analysis using maximum likelihood, evolutionary distance, and maximum parsimony methods. Molecular Biology and Evolution, 28, 2731–2739. https://doi.org/10.1093/molbev/msr121
  73. Uetz, P., Freed, P., Aguilar, R. & Hošek, J. (Eds.) (2022) The Reptile Database, http://www.reptile-database.org (accessed 28 July 2022)
  74. Wiegmann, A.F.A. (1834) Herpetologia Mexicana, seu Descriptio Amphibiorum Novae Hispaniae. Saurorum Species [Pars Prima Ed.]. Sumptibus C. G. Luderitz, Berlin, 54 pp., 10 pls. [reprint (1969) Society for the Study of Amphibian and Reptiles, Athens, Ohio, USA]
  75. Witte, G.-F. de (1973) Description d’un Gekkonidae nouveau de l’Iran (Reptilia, Sauria). Bulletin de l’Institut royal des sciences naturelles de Belgique, 49 (1), 1–6
  76. Zarudny, N.A. (1896) Itinéraire de l’expédition dans la Perse orientale en 1896. Annuaire du Musée Zooloogique de l’Academie Impériale des Sciences de St. Petersburg, I, XVIII–XXI. [in Russian]
  77. Zarudny, N.A. (1897) Une note sur les reptiles écailleux et nus du nord-est de la Perse. Annuaire du Musée Zoologique del’Academie Impériale des Sciences de St. Petersburg, II, 349–361. [in Russian]
  78. Zarudny, N.A. (1898) Itinéraire de l’expédition dans la Perse orientale en 1898. Annuaire du Musée Zoologique de l’Academie Impériale des Sciences de St. Petersburg, III, V–XII. [in Russian]
  79. Zarudny, N.A. (1902) Itinéraire de l’expédition de la Société Impériale Russe de Géographie dans la Perse orientale en 1900–1901. Annuaire du Musée Zoologique de l’Academie Impériale des Sciences de St. Petersburg, VII, I–IX. [in Russian]
  80. Zarudny, N.A. (1903) About reptiles and fishes of East Persia. Herpetological and ichthyological results of the excursion in Eastern Persia in 1898. Bulletin of the Imperial Russian Geographical Society, St. Petersburg, 36 (3), 1–42. [in Russian]
  81. Zarudny, N.A. (1904) Itinéraire de l’expédition dans la Perse occidentale en 1903–1904. Annuaire du Musée Zoologique del’Academie Impériale des Sciences de St. Petersburg, IX, I–VII. [in Russian]
  82. Zhao, E. & Adler, K. (1993) Herpetology of China. Society for the Study of Amphibians and Reptiles, Oxford, Ohio, 522 pp., 42 figs., 48 pls.