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Type: Articles
Published: 2012-09-07
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Review of the systematics, morphology and distribution of Asian Clawed Salamanders, genus Onychodactylus (Amphibia, Caudata: Hynobiidae), with the description of four new species

Department of Vertebrate Zoology, Biological faculty, Lomonosov Moscow State University, Leninskiye Gory, Moscow, GSP-1, 119991, Russia
State Key Laboratory of Genetic Resources and Evolution, and Yunnan Laboratory of Molecular Biology of Domestic Animals, Kunming Institute of Zoology, Chinese Academy of Sciences, Kunming 650223, China
Conservation Genome Resource Bank for Korean Wildlife (CGRB), Research Institute for veterinary science, College of Veterinary Medicine, Seoul National University, 151-742 Seoul, South Korea
Faculty of Agriculture and Life Science, Hirosaki University, 3 Bunkyo-cho, Hirosaki, Aomori, 036-8561, Japan
State Key Laboratory of Genetic Resources and Evolution, and Yunnan Laboratory of Molecular Biology of Domestic Animals, Kunming Institute of Zoology, Chinese Academy of Sciences, Kunming 650223, China
Jilin Huangnihe National Nature Reserve, Dunhua, 133704, China
Takashima 2nd Junior High School, 2-24-1 Takashimadaira, Itabashi-ku, Tokyo, 175-0082, Japan
Museo Nacional de Ciencias Naturales, Consejo Superior de Investigaciones Científicas (CSIC), C/José Gutiérrez Abascal 2, 28006, Madrid, Spain REFER Biodiversity Chair, University of Porto, CIBIO, Campus Agrário de Vairão, R. Padre Armando Quintas, 4485-661 Vairão, Portugal
Amphibia Caudata Hynobiidae Onychodactylus integrative taxonomy Asia salamander Onychodactylus koreanus Onychodactylus nipponoborealis Onychodactylus zhaoermii Onychodactylus zhangyapingi new species

Abstract

We describe four new species of Asian Clawed salamanders of the genus Onychodactylus (Caudata: Hynobiidae), basedon fresh material collected during fieldwork in Japan, Korea, the Russian Far East and northeastern China between 2003and 2010, as well as older voucher specimens deposited in several museums. Our analyses comprise all species currentlyrecognized within this genus across its entire distribution range. We follow an integrative taxonomic approach bycombining detailed morphological comparative analyses with molecular phylogenetic analyses. We find significantdifferences among species in this genus, based on morphological and molecular data, which resulted in the recognitionand description of four new species within this genus. The new species have uncorrected molecular divergences of over4.5–7.4% and 1.9–4.1% to their closest relatives in the mitochondrial COI and 16S rRNA genes respectively. In themolecular analyses, we found two very divergent lineages in Korea and Japan that need further investigation, as detailedmorphological data are not available for them. We also discuss our approach to delimit species on salamanders. For thenew species described in this group we evaluate their threat status according to IUCN criteria: O. koreanus sp. nov. Min,Poyarkov & Vieites and O. nipponoborealis sp. nov. Kuro-o, Poyarkov & Vieites are classified as Least Concern, whileO. zhaoermii sp. nov. Che, Poyarkov & Yan and O. zhangyapingi sp. nov. Che, Poyarkov, Li & Yan are classified as Vulnerable (Vu2a).

References

  1. Abé, Y. (1922) [On Ambystomidae from Japan]. Doubutsugaku zasshi [Zoological Magazine, Tokyo], 34, 328–332. (In Japanese)

    Adler, K. & Zhao, E.-M. (1990) Studies of hynobiid salamanders, with description of a new genus. Asiatic Herpetological Research, 3, 37–45.

    Akita, Y. & Miyazaki, K. (1991) Migration and cycle of reproduction of Onychodactylus japonicus in Mt. Hodatsu. Japanese Journal of Herpetology, 14, 29–38. (In Japanese)

    Akita, Y. (1982) Notes on the egg-laying site of Onychodactylus japonicus on Mt. Hodatsu. Japanese Journal of Herpetology, 9, 111–117. (In Japanese)

    Akita, Y. (1983) Winter reproduction of Onychodactylus japonicus. Ryoseihachurui Kenkyukaishi, 26, 1–6. (In Japanese)

    Akita, Y. (1985) Onychodactylus japonicus in Mt. Hodatsu during the reproductive period. Ryoseihachurui Kenkyukaishi, 31, 1–6. (In Japanese)

    Akita, Y. (1996) Onychodactylus japonicus. In: Sengoku, S., Hikida, T., Matsui, M., Nakaya, K. (Eds.), The Encyclopedia of Animals in Japan 5, Amphibians, Reptiles, Chondrichthyes. Heibonsha, Tokyo, p. 23. (In Japanese)

    AmphibiaWeb (2012): Information on amphibian biology and conservation. [web application]. 2008. Berkeley, California: AmphibiaWeb. Available: http://amphibiaweb.org/. (Accessed: Jan. 01, 2012)

    Antipenkova, T.P. (1982) [Tail ribs of the amphibians.] In: Morfofunktsionalnye Preobrazovaniya Pozvonochnykh v Protsesse Osvoeniya Sushi, Moscow, pp. 76–87. (In Russian)

    Barbour, T. & Loveridge, A. (1946) First supplement to typical reptiles and amphibians. Bulletin of the Museum of Comparative Zoology. Cambridge, Massachusetts, 96, 59–214.

    Boettger, O. (1892) Katalog der Batrachier-Sammlung im Museum der Senckenbergischen Naturforschenden Gesellshaft in Frankfurt am Main. Frankfurt a. M., Gebrüder Knauer, 73 pp.

    Bonaparte, C.L.J.L. (1839) Iconografia della fauna italica per le quattro classi degli animali vertebrati / di Carlo L. Principe Bonaparte, Tomo II. Amphibi; Roma: Dalla Tipografia Salviucci, (Fascicle 26). (In Italian)

    Boring, A.M. (1930) Distribution problems of North China Amphibia. Peking Natural History Bulletin, 1930–1931, V, 2, 39–47.

    Borkin, L.J. & Korotkov, Y, M. (1989) Amphibians and reptiles. In litt.: Redkiye pozvonochnye zhivotnye Sovetskogo Dalnego Vostoka i ikh okhrana [Rare vertebrate animals of the Soviet Far East and their conservation], Leningrad, pp. 21–22. (In Russian)

    Boulenger, G.A. (1886) First report on additions to the Batrachian collection in the Natural History Museum. Proceedings of the Zoological Society of London, June 29, 411–416.

    Boulenger, G.A. (1890) A list of the reptiles and batrachians of Amoorland. Annals and Magazine of Natural History, Series 6, 5, 137–144.

    Brame, A.H., Jr. (1972) A Checklist of the Living and Fossil Salamanders (Order Caudata) of the World, Pasadena, California, pp. 1–387.

    Bruford, M., Hanotte, O., Brookfield, J. & Burke, T. (1992) Single locus and multilocus DNA fingerprint. In: Hoelzel, A. (Ed.), Molecular Genetic Analysis in Conservation. IRL Press, Oxford, pp. 225–270.

    Chang, M.L.Y. (1936) Contribution a l'étude Morphologique, Biologique, et Systematique des Amphibiens Urodèles de la Chine. Paris, Picart, 157 pp. (Reprinted, Society for the Study of Amphibians and Reptiles, 1968.)

    Chang, M-H., Kim, B.-S., Park, S.-G., Kim, T.-W. & Oh, H.-S. (2009) Herpetofauna in Mountain Unmun. Korean Journal of Herpetology, 1, 1, 53–60.

    Che, J., Chen, H.-M., Yang, J.-X., Jin, J.-Q., Jiang, K., Yuan, Z.-Y., Murphy, R.W. & Zhang, Y.-P. (2012) Universal COI primers for DNA barcoding amphibians. Molecular Ecology Resources, 12, 2, 247–258.

    Clemen, G. & Greven, H. (2009) Sex dimorphic dentition and notes on the skull and hyobranchium in the hynobiid salamander Pachyhynobius shangchengensis Fei, Qu & Wu, 1983 (Urodela: Amphibia). Vertebrate Zoology, 59, 1, 61–69.

    Cloete, S.E. (1961) The cranial morphology of Rhyacotriton olympicus olympicus (Gaige). Annale Universiteit van Stellenbosch, 36, A, 113–145.

    Cope, E.D. (1889) Batrachia of North America. Bulletin of the United States National Museum, 34; Washington: Government Printing Office: pp. 5–525. (Reprinted, Eric Lundberg, 1963.)

    Dayrat, B. (2005) Toward integrative taxonomy. Biological Journal of the Linnean Society, 85, 407–415.

    Doi, K. (1920) [On Dorongyeong (Salamanders)] Chosen Iho, 62, 485–492. (In Korean and Japanese)

    Donndorff, J.A. (1798) Zoologische Beyträge zur XIII. Ausgabe des Linneischen Natursystems. Dritter Band [vol. III]. Amphibien und Fische. Leipzig: Weidmannschen Buchhandlung, 980 pp. (In German)

    Duméril, A.M.C. & Bibron, G. (1841) Erpétologie Générale ou Histoire Naturelle Complète des Reptiles. Vol. 8. Librarie Encyclopedique de Roret, Paris, France. i–vii + 1–792. (In French)

    Duméril, A.M.C., Bibron, G. & Duméril, A.H.A. (1854) Erpétologie Générale ou Histoire Naturelle Complète des Reptiles. Vol. 9. Librarie Encyclopedique de Roret, Paris, France. xx, 440 pp. (In French)

    Dunn, E.R. (1918) The collection of Amphibia Caudata of the Museum of Comparative Zoology. Bulletin of the Museum of Comparative Zoology, 62, 445–471.

    Dunn, E.R. (1923) The salamanders of the family Hynobiidae. Proceedings of the American Academy of Arts and Sciences, Boston, 58, 445–523.

    Emelianov, A.A. (1923) [Amphibians and Reptiles of Primorye]. – In: Primorye: ego priroda i khozyaistvo, Vladivostok: pp. 128–140. (In Russian)

    Emelianov, A.A. (1934) Amphibian and reptile fauna of the Sudzukhe River Valley. Vestnik Dalnevostochnogo filiala AN SSSR, 9, 125–135. (In Russian)

    Emelianov, A.A. (1940) Amfibii i Reptilii Sovetskogo Dalnego Vostoka [Amphibians and Reptiles of the Soviet Far East]. D. Sc. Diss., Vladivostok, 2 v. (In Russian)

    Emelianov, A.A. (1947) On the biology of Onychodactylus fischeri. Zoologicheskiy Zhurnal, 26, 65–70. (In Russian)

    Fei, L. (Ed.) (1999) [Atlas of Amphibians of China.] Henan Science and Technology Press, Zhengzhou, 432 pp. (In Chinese)

    Fei, L., Hu, Sh.-Q., Ye, Ch.-Y. & Huang, Y.-Zh., eds. (2006) Fauna Sinica. Amphibia. Vol. 1. General accounts of Amphibia. Gymnophiona and Urodela. Science Press, Beijing, 472 pp. + XVI pl. (In Chinese)

    Fouquet, A., Vences, M., Salducci. M.-D., Meyer, A., Marty, C., Blanc, M. & Gilles, A. (2007) Revealing cryptic diversity using molecular phylogenetics and phylogeography in frogs of the Scinax ruber and Rhinella margaritifer species groups. Molecular Phylogenetics and Evolution, 43, 567–582.

    Fritze, A. (1891) Die Fauna von Yezo im Vergleich zur Fauna des übrigen Japan. Mittheilungen der Deutsche Gesellschaft für Natur- und Völkerkunde Ostasiens, (Tokyo), V, 235–248.

    Frost, D.R. (2009) Amphibian Species of the World: an Online Reference. V. 5.3 (12 February, 2009). Available: http://research.amnh.org/herpetology/amphibia/; American Museum of Natural History, New York, USA. (Accessed: Jan. 01, 2011).

    Frost, D.R. (2011) Amphibian Species of the World: an Online Reference. V. 5.5 (31 January, 2011). Available: http://research.amnh.org/vz/herpetology/amphibia/ American Museum of Natural History, New York, USA. (Accessed: Jan. 01, 2012)

    Fu, J.-Zh. & Zeng, X.-M. (2008) How many species are in the genus Batrachuperus? A phylogeographical analysis of the stream salamanders (family Hynobiidae) from southwestern China. Molecular Ecology, 17, 1469–1488.

    García-París, M. & Wake, D.B. (2000) Molecular phylogenetic analysis of relationships of the tropical salamander genera Oedipina and Nototriton, with descriptions of a new genus and three new species. Copeia, 2000, 1, 42–70.

    Gassó Miracle, M.E., van den Hoek Ostende, L.W. & Arntzen, J.W. (2007) Type specimens of amphibians in the National Museum of Natural History, Leiden, The Netherlands. Zootaxa, 1482, 25–68.

    Gee, N.G. & Boring, A.M.. (1929) Check list of Chinese Amphibia with notes on geographical distribution. Peking Natural History Bulletin, 4, 15–51.

    Geerts, A.J.C. (1881) Notice sur la grande salamandre du Japon, Cryptobranchus japonicus v. d. Hoevan (Salamandra maxima, Schl., Sieboldia maxima, Bonap.). Nouvelles Archives du Muséum d'Histoire Naturelle, Serie 2, Paris, 5, 273–290.

    Glaw, F., Köhler, J., De la Riva, I., Vieites, D.R. & Vences, M. (2010) Integrative taxonomy of Malagasy treefrogs: combination of molecular genetics, bioacoustics and comparative morphology reveals twelve additional species of Boophis. Zootaxa, 283, 1–82.

    Good, D.A. & Wake, D.B. (1992) Geographic Variation and Speciation in the Torrent Salamanders of the Genus Rhyacotriton (Caudata: Rhyacotritonidae). University of California Publications in Zoology, 126, i-xi, 1–91.

    Göppert, E. (1896) Zur Phylogenese der Wirbeltierkralle. Morphologische Jahrbücher, 25, 1, 1–30 (1 tab., 19 ff.).

    Goris, R.C. & Maeda, N. (2004). Guide to the Amphibians and Reptiles of Japan. Krieger Publishing Company, Malabar, FL, 285 pp.

    Gray, J.E. (1850) Catalogue of the Specimens of Amphibia in the Collection of the British Museum. Part II. Batrachia Gradientia etc. Order of the Trustees, London, 72 pp.

    Griffin, P.C. & Solkin, V.A. (1995) Ecology and conservation of Onychodactylus fischeri (Caudata, Hynobiidae) in the Russian Far East. Abstracts of the Second Asian Herpetological Meeting, Ashgabat, p. 24.

    Guo, L.-j., Zhao, Y.-y., Zhou, Y.-y., Liang, Ch.-ch. & Wang, L.-w. (2009) Postembryonic Development of Respiratory Organs in Onychodactylus fischeri. Chinese Journal of Zoology, 44, 6, 103–108. (In Chinese)

    Hashimoto, J. (1910) [Ambystomatidae living on the north of Japan] Doubutsugaku zasshi [Zoological Magazine, Tokyo], 265, 497–499. (In Japanese)

    Hayase, N. & Yamane, S. (1982) Life history during the aquatic life period of a salamander, Onychodactylus japonicus (Houttuyn), at Mt. Tsukuba, Ibaraki, Japan. Japanese Journal of Ecology, 32, 395–403. (In Japanese)

    Highton, R. (1957) Correlating costal grooves with trunk vertebrae in salamanders. Copeia, 1957, 2, 107–109.

    Hilgendorf, F. (1880) Bemerkungen über die von ihm in Japan gesammelten Amphibien nebst Beschreibung zweier neuer Schlangenarten. Sitzungsberichte der Gesellschaft Naturforschender Freunde zu Berlin, 8, 111–121.

    Hillis, D.M., Moritz, C. & Mable, B.K. (1996) Molecular Systematics. Sinauer Assoc., 656 pp.

    Hoogmoed, M.S. (1978) An annotated review of the salamander types described in the Fauna Japonica. Zoologische Mededelingen (Leiden), 53, 91–105.

    Hou, M., Zhou, Z.-C., Li, P.-P. & Lue, S.-Q. (2009) Rediscovery of Pingia granulosus (Chang, 1933), and description of the neotype specimens. Sichuan Journal of Zoology, 28, 15–18. (In Chinese with English abstract)

    Houttuyn, M. (1782) Het onderscheid der salamanderen van de haagdissen in’t algemeen, en van de gekkos in’t byzonder. Verhandelingen / uitgegeven door het Zeeuwsch Genootschap der Wetenschappen te Vlissingen, 9, 305–336. (In Dutch)

    Houttuyn, M. (1787) Catalogus van eene uitmuntende Verzameling van allerley Soort van Dieren en dierlyke Zaaken, tot opheldering der Natuurlyke Historie …. Amsterdam, by J. van der Burgh en Zoon en J. C. Sepp, boekverkoopers, VIII + 178 + 1 p. (In Dutch)

    Ichimura, T. (1897) [Onychodactylus and Hynobius salamanders from Kaga]. Doubutsugaku zasshi [Zoological Magazine, Tokyo], 9, 107, 360–361. (In Japanese)

    Iizuka, K. & Yazawa, Sh. (1991) Onychodactylus fischeri from Korea. Heredity, 45, 10, 110–111. (In Japanese)

    Iizuka, K. & Yazawa, Sh. (1994) The karyotype, C bands and AgNO3 bands of a lungless salamander from Korea: Onychodactylus fischeri (Boulenger) (Amphibia, Urodela). Experientia, 50, 171–175.

    Iizuka, K., Poyarkov, N.A, & Koishi, H. (2011) Lectotype specimen of Japanese clawed salamander, Onychodactylus japonicus (Houttuyn 1782) collected by P. F. von Siebold. Bulletin of the Herpetological Society of Japan, 66, 2, 122–127. (In Japanese with English abstract)

    Iizuka, K., Poyarkov, N.A., Koishi, H., Shiraiwa, Y., Sessions, S.K. & Vassilieva, A.B. (2010) Limb development of Japanese clawed salamander, Onychodactylus japonicus (Houttuyn 1782). Bulletin of the Biogeographic Society of Japan, 65, 13–19. (In Japanese with English abstract).

    Ikebe, C., Amano, M., Koizumi, T., Kuro-o, M. & Kohno, S. (1995) Cytogenettic studies of Hynobiidae (Urodela) XIII. Morphological variation of chromosome No. 4 and the C-banded karyotype in Onychodactylus japonicus (Houttuyn). Japanese Journal of Herpetology, 16, 2, 35–41.

    Ikebe, C., Yamamoto, T. & Kohno, S. (1981) A preliminary note on the chromosomes of Onychodactylus japonicus (Houttuyn) (Amphibia: Urodela). Chromosome Information Service, 30, 17–18.

    Inukai, T. (1932) Urodelanarten aus Nordjapan mit Besonderer Berücksichtigung der Morphologie des Schädels. Journal of the Faculty of Science, Hokkaido Imperial University, Ser. 6 (Zoology), 1, 191–217. (In German)

    Inukai, T. (1933) On Onychodactylus from Korea. Annotationes Zoologicae Japonenses / Nihon Doubutsugaku ih, Tokyo, 14, 193–195, f. 1.

    Ivanova, N.V., deWaard, J. & Hebert, P.D.N. (2006) An inexpensive, automation-friendly proto-col for recovering high-quality DNA. Molecular Ecology Notes, 6, 998–1002.

    Iwasawa, H. & Kera, Y. (1978) Oviposition in the lungless salamander Onychodactylus japonicus induced by injection of frog pituitary and commercial gonadotropins. Japanese Journal of Herpetology, 7, 3, 51–55. (In Japanese)

    Iwasawa, H. & Kera, Y. (1980) Stages of normal development of Onychodactylus japonicus. Japanese Journal of Herpetology, 8, 73–89. (In Japanese)

    Iwasawa, H. (1976) Methods of collecting and smoking of Onychodactylus in Hinoemata. Hachu Ryosei Zatsu, 6, 105–109. (In Japanese)

    Iwasawa, H., Nomura, T. & Usuda, H. (1992) Spawning site of Onychodactylus japonicus found at the northern foot of Mt. Namba, Niigata Prefecture. Japanese Journal of Herpetology, 14, 212–213. (In Japanese)

    Ji, D., Liu, M., Liu, Z., Zhou, Y., Huang, K., Wen, S. & Zou, B. (1987) Fauna Liaoningica. Amphibia, Reptilia. Liaoning Sci. and Technol. Press: Shenyang, 170 pp. (In Chinese)

    Jiang, Y.-M. (1998) Lamei ying liangqi paxing dongwu ci hui [Latin-Chinese-English Names of Amphibians and Reptiles], Science Press, Beijing: 329 pp. (In Chinese and English)

    Kaneko, Y. & Matsui, M. (2004) Onychodactylus japonicus. In: IUCN 2011. IUCN Red List of Threatened Species. Version 2011.2. Accesible at: www.iucnredlist.org. (Accessed: Apr. 24, 2012)

    Kang, Y.-S. & Yoon, I.-B. (1975) Amphibia, Reptilia. In: Yoo, K.-C. (Ed.), Illustrated Encyclopedia of Fauna and Flora of Korea, 17, Korean Ministry of Education, Seoul, pp. 55–69. (In Korean)

    Kim, J. –B. (2009) Taxonomic List and Distribution of Korean Amphibians. Korean Journal of Herpetology, 1, 1–13.

    Kim, J.-B., Min, M.-S. & Matsui, M. (2003) A new species of lentic breeding Korean salamander of the genus Hynobius (Amphibia, Urodela). Zoological Science, 20, 1163–1169.

    Kobayashi, H. (1931) [Two salamanders from Shikoku.] Doubutsugaku zasshi [Zoological Magazine, Tokyo], 43, 709–710. (In Japanese)

    Kozik, S.S. & Truberg, A.G. (1985) Onychodactylus fischeri. Priroda (Moscow), 9, 55–57. (In Russian)

    Kozik, S.S. (1991) Record of the Onychodactylus fischeri egg-laying site. Zoologicheskiy Zhurnal, 70, 159–160. (In Russian)

    Kudo, T. (1934a) [Onychodactylus japonicus: notes on its breeding habits.] Acta Anatomica (Inst. Niigata), 4, 1–16, pls. I–III. (In Japanese)

    Kudo, T. (1934b) [Über die Entwicklung der lungelosen Salamander] Mitteilungen aus dem Anatomische Institut Niigata, 4: 1–18, pl. I. (In Japanese)

    Kudo, T. (1935) On breeding habits of Onychodactylus japonicus. Mitteilungen aus dem Anatomische Institut Niigata, 5, 1–16.

    Kuzmin, S.L. & Maslova, I.V. (2004) Amphibians of the Russian Far East. Advances in Amphibian Research in the former Soviet Union, Vol. 8, Pensoft Publishers, Sofia-Moscow, 462 pp.

    Kuzmin, S.L. & Maslova, I.V. (2005) Zemnovodnye rossiyskogo Dalnego Vostoka [Amphibians of the Russian Far East.] Moscow, KMK Scientific Press, 434 pp. (In Russian)

    Kuzmin, S.L. & Solkin, V.A. (1993) Foraging and seasonal changes in food composition of Onychodactylus fischeri (Boulenger) (Caudata: Hynobiidae). Herpetozoa, 6, 3/4, 83–87.

    Kuzmin, S.L. (1995) The Clawed Salamanders of Asia. Genus Onychodactylus. Biology, distribution, and conservation. Die Neue Brehm-Buecherei, 622, 108 pp.

    Kuzmin, S.L. (1999) The Amphibians of the Former Soviet Union. Pensoft Series Faunistica No. 12. Sofia, Bulgaria, and Moscow, Russia: Pensoft, 538 pp.

    Kuzmin, S.L., Matsui, M., Zhao W.-G., Maslova, I.V., & Tuniyev, B.S. (2004) Onychodactylus fischeri. In: IUCN 2011. IUCN Red List of Threatened Species. Version 2011.2. Accesible at: www.iucnredlist.org. (Accessed: Apr. 24, 2012)

    Lai, J.-S., & Lue, K.-Y. (2008) Two new Hynobius (Caudata: Hynobiidae) salamanders from Taiwan. Herpetologica, 64, 63–80.

    Larson, A., Weisrock, D.W. & Kozak, K.H. (2003) Phylogenetic Systematics of Salamanders (Amphibia: Urodela), A Review. In: Sever, D.M. (Ed.) Reproductive Biology and Phylogeny of Urodela. Reproductive Biology and Phylogeny, 1, pt. A, pp. 31–108.

    Lebedkina, N.S. (1981) [Some peculiarities of bony skull development in the Ussurian salamander, Onychodactylus fischeri Blgr. (Hynobiidae).] In: Gerpetologicheskiye Issledovaniya v Sibiri i na Dalnem Vostoke [Herpetological Investigations in Siberia and the Far East], Leningrad, pp. 61–65. (In Russian).

    Lee, J.-H., Park, D.-S., Ra, N.-Y. & Kim, J.-K. (2009) Herpetofauna in Odaesan National Park. Korean Journal of Herpetology, 1, 1, 45–52.

    Lee, J.-H., Ra, N.-Y., Eom, J.H. & Park, D.S. (2008) Population dynamics of the long-tailed clawed salamander larva, Onychodactylus fischeri, and its age structure in Korea. Journal of Ecological Field Biology, 31, 31–36.

    Li, J.-L. (2004) On the biology of Onychodactylus fischeri found in Xiuyan Co., and protection of its habitat. Sichuan Journal of Zoology, 23, 3, 169–171. (In Chinese)

    Li, S., Tian, Y.-Zh., & Gu, X.-M. (2010) A new species of the genus Pseudohynobius (Caudata, Hynobiidae) from Guizhou, China. Acta Zootaxonomica Sinica, 35, 2, 407–412. (In Chinese with English abstract)

    Litvinchuk, S.N. & Borkin, L.J. (2003) Variation in the number of trunk vertebrae and in count of costal grooves in salamanders of the family Hynobiidae. Contributions to Zoology, 72, 4, 195–209.

    Litvinchuk, S.N. & Borkin, L.J. (2009) [Evolution, taxonomy and distribution of Crested Newts (Triturus cristatus complex) on territories of Russia and adjacent countries]. St.-Petersburg, Evropeyskiy Dom, 590 pp. + 24 pl. (In Russian)

    Litvinchuk, S.N., Borkin, L.J. & Rosanov, J.M. (2004) Intraspecific and interspecific genome size variation in Hynobiid salamanders of Russia and Kazakhstan: determination by flow cytometry. Asiatic Herpetological Research, 10, 282–294.

    Lu, Y.-Y., Wang, J., Liu, X.-H., Zhou, Zh.-Y., Chen, X.-Zh., Wang, X.-A., Zhang, P. & Li, P.-P. (2004) Microstructure of Skin of FischerpsFischer’s Clawed Salamander (Onychodactylus fischeri). Sichuan Journal of Zoology, 123, 13, 178–182, pl. VI.

    Maslova, I.V. (1998) The current status of populations of the Long-Tailed Clawed Salamander (Onychodactylus fischeri) in the Ussuriiskii Nature Reserve. Advances in Amphibian Research in the former Soviet Union, Vol. 3, Pensoft Publishers, Sofia-Moscow, pp. 193–198.

    Maslova, I.V. (1999) [New findings of Onychodactylus fischeri.] In: Tezisy Dokladov Tretyey Dalnevostochnoy Konferentsii po Zapovednomu Delu, Vladivostok, pp. 70–71. (In Russian)

    Matsui, M., Kokuryo, Y., Misawa, Y. & Nishikawa, K. (2004) A new species of salamander of the genus Hynobius from central Honshu, Japan (Amphibia, Urodela). Zoological Science, 21, 661–669.

    Matsui, M., Nishikawa, K., Utsunomiya, T. & Tanabe, S. (2006) Geographic allozyme variation in the Japanese clouded salamander Hynobius nebulosus (Amphibia: Urodela). Biological Journal of the Linnean Society, 89, 311–330.

    Matsui, M., Yoshikawa, N., Tominaga, A., Sato, T., Takenaka, S., Tanabe, S., Nishikawa, K. & Nakabayashi, S. (2008) Phylogenetic relationships of two Salamandrella species as revealed by mitochondrial DNA and allozyme variation (Amphibia: Caudata: Hynobiidae). Molecular phylogenetics and evolution, 48, 84–93.

    Mazin, A.L. (1978) Genome size in some Caudata and Anura of the Far East. In: Gerpetofauna Dalnego Vostoka i Sibiri, Vladivostok, pp. 20-21. (In Russian).

    Medvedeva, I.M. (1975) [Distribution of seismosensory organs of the head in the most early free living larvae of Onychodactylus fischeri (Amphibia, Urodela).] Doklady Acad. Nauk SSSR, 246, 724–727. (In Russian)

    Merrem, B. (1820) Versuch eines Systems der Amphibien / Tentamen Systematis Amphibiorum. J.C. Krieger, Marburg, Germany. xv, xv, 185–191 pp. (In German)

    Meyer, F.A.A. (1795) Synopsis Reptilium, novam ipsorum sistens generum methodum, nec no Gottingensium huius ordinis animalium enumerationem. Gottingae, apud Vandenhoek & Ruprecht, 32 pp.

    Min, M.-S., Yang, S.-Y., Bonett, R.-M., Vieites, D.-R., Brandon, R.-A. & Wake, D.-B. (2005) Discovery of the first Asian plethodontid salamander. Nature, 435, 87–90.

    Misawa, Y. (1989) The method of counting costal grooves. In: Matsui, M. et al. (Eds.), Current Herpetology in East Asia, Kyoto, Herpetological Society of Japan, pp. 129–134.

    Miyadi, D. & Koba, K. (1940) [On Manchurian Urodeles]. Kanto-shu oyobi Manshu-koku riku-sui seibutsu chosa-sho, 20–21, f. 3. (In Japanese)

    Morescalchi, A., Odierna, G. & Olmo, E. (1979) Karyology of the primitive salamanders, family Hynobiidae. Experientia, 35, 1434–1436.

    Mori, T. (1928) “1927” A Hand-list of the Manchurian and Eastern Mongolian Vertebrata. Published by the author, Tokyo, 1, 1, 10, 2, 186, 23, 1 pp.

    Mott, T. & Vieites, D.-R. (2009) Molecular phylogenetics reveals extreme morphological homoplasy in Brazilian worm lizards challenging current taxonomy. Molecular Phylogenetics and Evolution, 51, 190–200.

    Murphy, R.W., Lowcock, L.A., Smith, C., Darevsky, I.S., Orlov, N., MacCulloch, R.D. & Upton, D.E. (1997) Flow cytometry in biodiversity surveys: methods, utility and constraints. Amphibia-Reptilia, 18, 1–13.

    Nakamura, D. (1941) [On the ecology of Onychodactylus japonicus]. Botany and Zoology (Tokyo), 9, 422–426, f. 2; 515–521, f. 8. (In Japanese)

    Nakamura, K., & Ueno, S. (1963) Genshoku Nihon ryousei hachuuri zukan [Japanese Reptiles and Amphibians in Colour]. Hoikusha, Osaka, 215 pp. (In Japanese)

    Nambu, H. (1996) Notes on the breeding site of Onychodactylus japonicus from Ooyama-machi, Toyama Prefecture, central Japan. Bulletin of the Toyama Science Museum, 19, 41–43. (In Japanese)

    Namiye, M. (1908) [Onychodactylus japonicus from Mt. Tsukuba.] Doubutsugaku zasshi [Zoological Magazine, Tokyo], 20, 399. (In Japanese)

    Nikitin, V.B. (1986) On the nasal muscles in Anura and Urodela. In: Studies in Herpetology. Prague, pp. 251–254.

    Nikolskii, A.M. (1906) "1905". [Les reptiles et les amphibiens de l'Empire de Russie]. Mémoires de l'Académie Impériale des Sciences de St. Pétersbourg, Série 8, 17, 1–518. (In Russian)

    Nikolsky [Nikolskii], A.M. (1896). Geomolge fischeri Blgr., nazemnyi triton iz Ussuriyskogo Kraya. [Geomolge fischeri Blgr., a terrestrial newt from the Ussuri Territory.] Annuaire du Musée Zoologique de l'Academie Impériale des Sciences de St. Pétersbourg, 1, 77-80. (In Russian)

    Nikolsky [Nikolskii], A.M. (1912) 1914. Novyi vid tritona iz Primorskoy Oblasti. [A new species of newt from Maritime Province.] Trudy Trotskosavsko-Kyakhtinskogo otdeleniya Priamurskogo Otdela Imperatorskogo RGO, 15, 3, 31–41. (In Russian)

    Nikolsky [Nikolskii], A.M. (1914) "1913". Onychodactylus rossicus sp. n. (fam. Salamandridae). Annuaire du Musée Zoologique de l'Academie Impériale des Sciences de St. Pétersbourg, 18, 260–262.

    Nikolsky [Nikolskii], A.M. (1918) Fauna Rossii i sopredel’nykh stran. Zemnovodnye (Amphibia). [Fauna of Russia and Adjacent Countries. Amphibians (Amphibia).] Petrograd, Russia: Russian Academy of Sciences, 311 pp. (In Russian) (Reprinted by Israel Program for Scientific Translations, 1962.)

    Nishikawa, K., Jiang, J.-P. & Matsui, M. (2011b) Two New Species of Pachytriton from Anhui and Guangxi, China (Amphibia: Urodela: Salamandridae). Current Herpetology, 30, 1, 15–31.

    Nishikawa, K., Jiang, J.-P., Matsui, M. & Chen, C.-S. (2009) Morphological variation in Pachytriton labiatus and a re-assessment of the taxonomic status of P. granulosus (Amphibia: Urodela: Salamandridae). Current Herpetology, 28, 49–64.

    Nishikawa, K., Jiang, J.-P., Matsui, M. & Mo, Y.-M. (2011a) Unmasking Pachytriton labiatus (Amphibia: Urodela: Salamandridae), with description of a new species of Pachytriton from Guangxi, China. Zoological Science, 28, 453–461.

    Nishikawa, K., Matsui, M., Tanabe, S. & Sato, S. (2007) Morphological and allozymic variation in Hynobius boulengeri and H. stejnegeri (Amphibia: Urodela: Hynobiidae). Zoological Science, 24, 752–766.

    Nomura, H. (1905) [Collecting on Onychodactylus japonicus.] Doubutsugaku zasshi [Zoological Magazine, Tokyo], 17, 226–228. (In Japanese)

    Ohno, M. (1982) Onychodactylus japonicus (Houttuyn). In: [Distributions of Important Amphibians and Reptiles in Japan], Environment Agency of Japan, Tokyo, 2, pp. 134–141. (In Japanese)

    Okada, S. (1891) [Catalogue of the Vertebrated Animals of Japan]. Tokyo, Japan: Kinkodo, 128 pp. (In Japanese)

    Okada, Y. (1933) [The herpetological fauna in the vicinity of Nikko, Japan]. Tokyo Bunrika Daigaku, 1, B, 159–173. (In Japanese)

    Okada, Y. (1934) A contribution toward a check list of the Urodeles of Japan. Copeia, 12, 1, 16–19.

    Okada, Y. (1935) A general review of the classification and distribution of Japanese salamanders. Doubutsugaku zasshi [Zoological Magazine, Tokyo], 47, 575–589. (In Japanese)

    Okada, Y. (1937a) [Notes on the Amphibia of the Tohoku district, Northern Japan.] Saito Hoonkai Museum Research Bulletin, 12, 177–206, ff. 10–13. (In Japanese)

    Okada, Y. (1937b) [On Onychodactylus japonicus Houttuyn.] Doubutsugaku zasshi [Zoological Magazine, Tokyo], 35, 263–268. (In Japanese)

    Okajima, K. (1908) Die Osteologie des Onychodactylus japonicus. Zeitschrift für Wissenschaftliche Zoologie, Leipzig, 91, 351–381. (In German)

    Okajima, K. (1909a) 1910. Untersuchungen über die Sinnesorgane von Onychodactylus. Zeitschrift für Wissenschaftliche Zoologie, Leipzig, 94, 171–239. (In German)

    Okajima, K. (1909b) Über des Hyobranchialskelett von Onychodactylus. Anatomische Anzeiger, 34, 182. (In German)

    Okajima, K. (1922) Zur Anatomie des Hyobranchialskelettes bei den Onychodactyluslarven. Folia Anatomica Japonica, 1, 195–200. (In German)

    Okajima, K., & Tsusaki, T. (1921) Beiträge zur Morphologie des Skleralknorpels bei den Urodelen. Zeitschrift fur Anatomie und Entwicklungsgeschichte, 60, 631–651.

    Okhotina, M.V. (1959) On the specification of range borders of some amphibians and reptiles of Primorye. Soobscheniya Dalnevostochnogo filiala AN SSSR, 11, 139–143. (In Russian)

    Olmo, E. (1983) Nucleotype and cell size in vertebrates: a review. Basic Applications of Histochemistry, 27, 227–256.

    Padial, J.M., Castroviejo-Fisher, S., Köhler, J., Vilà, C., Chaparro, J.C. & De la Riva, I. (2009) Deciphering the products of evolution at the species level: the need for an integrative taxonomy. Zoologica Scripta, 38, 431-447.

    Padial, J.M., Miralles, A., De la Riva, I., & Vences, M. (2010) The integrative future of taxonomy. Frontiers in Zoology, 2010, 7, 16.

    Park, D.-S. (2005) The First observation of breeding of the Long-tailed Clawed Salamander, Onychodactylus fischeri, in the field. Current Herpetology, 24, 1, 7–12.

    Parker, W.K. (1879) On the morphology of the skull in the Amphibia Urodela. Transactions of the Linnean Society of London, part 2, 165–212; part 3, No. 5, 188, pl. 19, ff. 1–3.

    Peng, R., Zhang, P., Xiong, J.-L., Gu, H.-J., Zeng, X.-M. & Zou, F.-D. (2010) Rediscovery of Protohynobius puxiongensis (Caudata: Hynobiidae) and its phylogenetic position based on complete mitochondrial genomes. Molecular phylogenetics and evolution, 56, 1, 252–258.

    Poyarkov, N.A. & Kuzmin, S.L. (2008) Phylogeography of the Siberian Newt Salamandrella keyserlingii by Mitochondrial DNA Sequence Analysis. Russian Journal of Genetics, 44, 8, 948–958.

    Ratnikov, V.Y. & Litvinchuk, S.N. (2007) Comparative morphology of trunk and sacral vertebrae of tailed amphibians of Russia and adjacent countries. Russian Journal of Herpetology, 14, 3, 177–190.

    Ratnikov, V.Y. & Litvinchuk, S.N. (2009) Atlantal vertebra of tailed amphibians of Russia and adjacent countries. Russian Journal of Herpetology, 16, 1, 57–68.

    Regel, E.D. & Epstein [Epshtein], S.M. (1975). [Some peculiarities of biology of Onychodactylus fischeri.] Zoologicheskiy Zhurnal, 54, 1515–1524. (In Russian)

    Regel, E.D. & Epstein [Epshtein], S.M. (1977) [External morphology of the larvae of Onychodactylus fischeri Boul., Hynobiidae.] Zoologicheskiy Zhurnal, 56, 901–912. (In Russian)

    Regel, E.D. (1970) [Some eco-morphological peculiarities of Onychodactylus fischeri Blg.] In: Amfibii Dalnego Vostoka i Sibiri, Vladivostok: pp. 29–30. (In Russian)

    Regel, E.D. (1979) [Peculiarities of development and morphology of olfactory capsules in the Ussurian salamander (Onychodactylus fischeri Blgr., Hynobiidae, Urodela) in comparison with other amphibians.] In: Ecology and Systematics of Amphibians and Reptiles, Leningrad: pp. 71–97. (In Russian)

    Regel, E.D. (1981) [Some peculiarities of structure of the heart in the urodelan family Hynobiidae.] In: Gerpetologicheskiye Issledovaniya v Sibiri i na Dalnem Vostoke / Herpetological Investigations in Siberia and the Far East, Leningrad: pp. 91–103. (In Russian)

    Rein, J.J. (1884) Reptiles and Batrachians of Japan: Travels and Researches. Translated from German. New York, 1884, pp. 186–189.

    Sambrook, J. & Russell, D.W. (2001) Molecular Cloning: A Laboratory Manual. Cold Spring Harbor Laboratory Press, Cold Spring Harbor, New York, 2344 pp.

    Sato, I. (1934a) [Taxonomical study of the salamanders of the nebulosus group, with special reference to the morphology of the skull.] Doubutsugaku zasshi [Zoological Magazine, Tokyo], 46, 214–224. (In Japanese)

    Sato, I. (1934b) [On the species of Hynobius of the naevius group with special reference to the morphology of the skull.] Botany and Zoology (Tokyo), 2, 1843–1852. (In Japanese)

    Sato, I. (1934c) [Studies on tailed amphibians from Shikoku. I. On the salamanders found at Mt. Ishizuchi and its vicinity.] Doubutsugaku zasshi [Zoological Magazine, Tokyo], 46, 464–472. (In Japanese)

    Sato, I. (1937a) A synopsis of the family Hynobiidae of Japan. Bulletin of the Biogeographical Society of Japan, 7, 31–45.

    Sato, I. (1937b) Fauna Nipponica. vol. XV (III) 1. Amphibia, Caudata. Tokyo, 74 pp.

    Sato, I. (1939). [Salamander studies in Tokyo]. In: Yoshida hakase shukuga kinen-shi [Celebrations for commemoration of Prof. Yoshida], Osaka, pp. 731-801, f. 52. (In Japanese)

    Sato, I. (1943) Nippon-san yubirui sosetsu [A Monograph of the Japanese Tailed Batrachians]. Nippon Shuppansha, Osaka, 520 pp. (In Japanese).

    Schlegel, H. (1858) Handleiding tot de beoefening der dierkunde. Breda, 628 pp. (In Dutch)

    Schneider, J.G. (1799) Historia Amphibiorum Naturalis et Literarariae. Fasciculus Primus. Continens Ranas, Calamitas, Bufones, Salamandras et Hydros in Genera et Species Descriptos Notisque suis Distinctos. Fasc. 1. Frederick Frommann, Jena. I: xiv, 1-266. (In Latin)

    Sclater, W.L. (1892) A List of the Batrachia of the [Calcutta] Indian Museum. London: Taylor and Francis, 43 pp.

    Serbinova, I.A., & Solkin, V.A. (1995) Reproductive behavior in the long-tailed salamander (Onychodactylus fischeri Boulenger). Asiatic Herpetological Research, 6, 114–119.

    Sessions, S.K., Leon, P.E. & Kezer, J. (1982) Cytogenetic studies of the Chinese giant salamander Andrias davidianus (Blanchard): the evolutionary significance of the cryptobranchoid karyotypes. Chromosoma, 86, 341–357.

    Shen, Y.-H., Deng, X.-J. & Wang, B. (2004) A new hynobiid species, Hynobius guabangshenensis, from Hunan Province, China (Amphiba: Hynobiidae). Acta Zoologica Sinica, 50, 209–215. (In Chinese with English abstract)

    Shen, Y.-H., Shen, D.-W. & Mo, X.-Y. (2008) A new species of salamander Pachytriton archospotus from Hunan Province, China (Amphibia, Salamandridae). Acta Zoologica Sinica, 54, 645–652. (In Chinese with English abstract)

    Shim, J.-H. (2001) [The frog, singing of life.] Seoul: DDworld, 217 pp. (In Korean)

    Siebold, P.F., von (1897) Nippon. Archiv zur Beschreibung von Japan und dessen Neben- und Schutzländern: Jezo mit den Südlichen Kurilen, Krafto, Koorai und den Liukiu-Inseln. 2. veränderte und ergänzte Auflage, hrsg. von seinen Söhnen, 2 vol., Wurzburg und Leipzig, 1350 pp.

    Smirina, E.M., Serbinova, I.A. & Makarov, A.N. (1994) [Some difficulties in age determination of amphibians by annual layers in their bones with special reference to the Long-tailed salamander, Onychodactylus fischeri (Amphibia, Hynobiidae).] Zoologicheskiy Zhurnal, 66, 735–745. (In Russian)

    Smirnov, S.V. & Vassilieva, A.B. (2002) Skeletal and dental ontogeny in the long-tailed clawed salamander, Onychodactylus fischeri (Urodela: Hynobiidae). Russian Journal of Herpetology, 9, 1, 21–32.

    Smirnov, S.V. & Vorobyeva, E.I. (1988) Morphological and evolutionary change in the amphibian sound-conducting apparatus. Anatomische Anzeiger, 166, 317–322.

    Smirnov, S.V. (1987) The sound-conducting apparatus of Urodela: structure, function and evolutionary pathways. Zoologicheskiy Zhurnal, 66, 735–745. (In Russian)

    Smith, M.A., 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.

    Solkin, V.A. (1993) On the eclogy of the salamander Onychodactylus fischeri (Caudata: Hynobiidae). Herpetozoa, 6, 1/2, 29–36.

    Song, J.-Y. & Lee, I.-S. (2005) Elevation distribution of Korean Amphibia. Yangseo Pachunglyu Haghoeji [Journal of Herpetology], 1, 1, 15–19. (In Korean)

    Sparreboom, M. (2012). Salamanders of the Old World: an Online Catalogue. Electronic database accessible at science.naturalis.nl/salamanders, Netherlands Centre for Biodiversity Naturalis. Leiden. The Netherlands. (Accessed: Jan. 01, 2012)

    Stejneger, L. (1907) Herpetology of Japan and adjacent territory. Bulletin of the United States National Museum, 58, 1–577. (Reprinted Society for the Study of Amphibians and Reptiles, 1996.)

    Strauch, A. (1870) Revision der Salamandriden-Gattungen nebst Beschreibung einiger oder weniger bekannten Arten dieser Familie. Mémoires de l'Académie Impériale des Sciences de St. Pétersbourg, Ser. 7, 16, 4, 1–110. (In German)

    Tagirova, V.T. (1977) Zoologicheskiye Ekskursii v Primaurye [Zoological Excursions in Priamurye]. Khabarovsk, 95 pp. (In Russian)

    Tago, K. (1903) [Notes on a salamander collection from Nikko] Doubutsugaku zasshi [Zoological Magazine, Tokyo], 15, 182, 460–464. (In Japanese)

    Tago, K. (1904) [Observations on Japanese Salamanders]. Doubutsugaku zasshi [Zoological Magazine, Tokyo], 16, 184, 41–46, color pl. 1. (In Japanese)

    Tago, K. (1907a) [A research on reproduction of Japanese salamanders] Doubutsugaku zasshi [Zoological Magazine, Tokyo], 19, 225, 191–205, fig. 1. (In Japanese)

    Tago, K. (1907b) [A research on reproduction of Japanese salamanders] Doubutsugaku zasshi [Zoological Magazine, Tokyo], 19, 226, 229–248, fig. 3; 19, 228, pl. 1. (In Japanese)

    Tago, K. (1931) Imori to Sanshouo. [Newts and Salamanders]. Geisoudo, Kyoto, 210 + 8 pp., pls. 28, fig. XXV. (In Japanese)

    Takashima, H. (1936) [A survey of Japanese Caudata.] Tokyo Hakubutsugaku Zasshi, 34, 17–25. (In Japanese)

    Temminck, C.J. & Schlegel, H. (1838) Fauna Japonica. Les Sauriens. Les Batraciens. Vol. 3. In: Siebold, P.F.v. (Ed.) Fauna Japonica sive Descriptio animalium, quae in itinere per Japoniam, jussu et auspiciis Superiorum, qui summum in India Batava imperium tenent, suscepto annis 1823-1830, collegit, notis, observationibus et adumbrationibus illustravit, Lugduni Batavorum, Amsterdam, 329 pp. (In French)

    Terentjev, P.V. & Chernov, S.A. (1936) Kratkiy Opredelitel’ Zemnovodnykh i Presmykayushchikhsya SSSR [Brief Guide to Amphibians and Reptiles of the USSR; 1st edition]. Moscow and Leningrad: Uchpedgiz, 96 pp. (In Russian)

    Terentjev, P.V. & Chernov, S.A. (1940) Opredelitel’ presmykayushchikhsya i zemnovodnykh SSSR [Guide to Reptiles and Amphibians of the USSR, 2d edition]. Moscow and Leningrad: Uchpedgiz, 184 pp. (In Russian)

    Terentjev, P.V. & Chernov, S.A. (1949) Opredelitel’ presmykayushchikhsya i zemnovodnykh [Guide to Reptiles and Amphibians], 3d edition. Moscow: Sovetskaya Nauka, 340 pp. (In Russian)

    Terentjev, P.W. [P.V.] (1938) Notes on salamanders of the family Hynobiidae. Copeia, 1938, 1, 17–18.

    Thiesmeier, B. (1996) A morphological and ecological approach to the classification of the larvae of the Semirechensk salamander (Ranodon sibiricus) and the long-tailed clawed salamander (Onychodactylus fischeri). Advances in Amphibian Research in the Former Soviet Union, 1, 131–138.

    Thireau, M. (1972) Catalogue des types D`Urodèles du Muséum National d`Histoire Naturelle. Revue Critique, Muséum National d`Histoire Naturelle, Laboratoire de zoologie (Reptiles et Amphibiens), 97 pp. (In French).

    Thireau, M. (1975a) L’allometrie pondérale encephalo-somatique chez les urodeles. I. Relations intraspecifiques. Bulletin du Muséum national d'Histoire naturelle, Paris, 3 ser., 297, Zool. 207, 467–482.

    Thireau, M. (1975b) L’allometrie pondérale encephalo-somatique chez les urodeles. I. Relations interspecifiques. Bulletin du Muséum national d'Histoire naturelle, Paris, 3 ser, 297, Zool. 207, 483–501.

    Thompson, J.D., Gibson, T.J., Plewniak, F., Jeanmougin, F. & Higgins, D.G. (1997) The CLUSTAL_X windows interface: flexible strategies for multiple sequence alignment aided by quality analysis tools. Nucleic Acids Research, 25, 24, 4876–4882.

    Thorn, R. (1968) Les Salamandres d’Europe d’Asie et d’Afrique du Nord. Paul Lechevalier: Paris. (4), iv, 376 pp. (In French)

    Thunberg, C.P. (1787a) Beskrifning på några sällsynte och okände Ödlor. Kongliga Svenska Vetenskaps-Akademiens Nya Handlingar, Stockholm, 8, 123–128. (In Swedish)

    Thunberg, C.P. (1787b) Museum Naturalium Academiae Upsaliensis, Upsala, Pars 2, 32 pp. (In Latin)

    Thunberg, C.P. (1791) Resa uti Europa, Afrika, Asia, Forrattad aren 1770-1779. Del 1-4, Upsala, pp. 1788–1793. (In Swedish)

    Tominaga, A., Matsui, M., Nishikawa, K., Tanabe, S. & Sato, S. (2005) Morphological discrimanation of two genetic groups of a Japanese salamander, Hynobius naevius (Amphibia, Caudata). Zoological Science, 22, 1229–1244.

    Troschel, F.H. (1877) Über Onychodactylus japonicus Bonap. Archiv für Naturgeschichte, 43, 1, 199–215, pl. XV.

    Tschudi, J.J. von (1838) Classification der Batrachier, mit Berucksichtigung der fossilen Thiere dieser Abtheilung der Reptilien. Mem. Soc. Sci. Neuchâtel: Petitpierre, 99 (2) pp. (In German) (Reprinted Society for the Study of Amphibians and Reptiles, 1967.)

    Uchiyama, R., Maeda, N., Numata, K. & Seki, S. (2002) [A Photographic Guide; Amphibians and Reptiles in Japan]. Heibonsha, Tokyo, 336 pp. (In Japanese)

    Vences, M., Thomas, M., Bonett, R.M. & Vieites, D.R. (2005a) Deciphering amphibian diversity through DNA barcoding: chances and challenges. Philosophical Transactions of the Royal Society London, Ser. B, 360, 1859–1868.

    Vences, M., Thomas, M., van der Meijden, A., Chiari, Y. & Vieites, D.R. (2005b) Comparative performance of the 16S rRNA gene in DNA barcoding of amphibians. Frontiers in Zoology, 2, 5, 1–12.

    Venczel, M. (1999a) Land salamanders of the family Hynobiidae from the Neogene and Quaternary of Europe. Amphibia-Reptilia, 20, 401-412.

    Venczel, M. (1999b) Fossil land salamanders (Caudata, Hynobiidae) from the Carpathian basin: relation between extinct and extant genera. Acta Paleontologica Romaniae, 2, 489–492.

    Vieites, D.R., Min, M.-S. & Wake, D.B. (2007) Rapid diversification and dispersal during periods of global warming by plethodontid salamanders. Proceedings of the National Academy of Sciences of the USA, 2007, 104, 50, 19903–19907.

    Vieites, D.R., Wollenberg, K.C., Andreone, F., Köhler, J., Glaw, F. & Vences, M. (2009) Vast underestimation of Madagascar's biodiversity evidenced by an integrative amphibian inventory. Proceedings of the National Academy of Sciences of the USA, 106, 8267–8272.

    Vinogradov, A.E. (1998) Genome size and GC-percent in vertebrates as determined by flow cytometry: the triangular relationship. Cytometry, 31, 100–109.

    Wang, H.-H., Li, L.-Y., Wang, L.-W. & Liang, Ch.-Ch. (2007) Morphological and histological studies on the telencephalon of the salamander Onychodactylus fischeri. Neuroscience Bulletin, 23, 3, 170–174. (In Chinese)

    Wang, H.-H., Li, L.-Y., Wang, L.-W. & Liang, Ch.-Ch. (2008a) Histology of telencephalon of adult and subadult Onychodactylus fischeris [sic!]. Sichuan Journal of Zoology, 27, 2, 223–226. (In Chinese)

    Wang, L.-W., Liang, Ch.-Ch., Huang, W. & Peng, X. (2008b) Influence of the ambient temperature on body temperature and energy metabolism of the salamander Onychodactylus fischeri. Acta Zoologica Sinica, 54, 4, 640–644. (In Chinese)

    Wang, L.-W., Zhao, Y.-Y. & Liang, Ch.-Ch. (2004) Studies on the Skeletal System of Onychodactylus fischeri ( Boulenger, 1886). Sichuan Journal of Zoology, 23, 3, 172–177, pl. V. (In Chinese)

    Wei, G., Xiong, J.-L. & Zeng, X.-M. (2009) A new species of hynobiid salamander (Urodela: Hynobiidae: Pseudohynobius) from southwestern China. Zootaxa, 2149, 62–68.

    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 divergent lineages in Mertensiella luschani associated with the rise of Anatolia. Molecular Phylogenetics and Evolution, 18, 3, 434–448.

    Won, H.-G. (1971) Choson Ryangso Pyachyun-gryuchji [Amphibians and Reptiles of Korea]. Pyongyang, 170 pp. (In Korean)

    Wu, Y., Jiang, K. & Hanken, J. (2010b) A new species of newt of the genus Paramesotriton (Salamandridae) from southwestern Guangdong, China, with a new northern record of P. longliensis from western Hubei. Zootaxa, 2494, 45–58.

    Wu, Y., Wang, Y., Jiang, K., & Hanken, J. (2010a) Homoplastic evolution of external colouration in Asian stout newts (Pachytriton) inferred from molecular phylogeny. Zoologica Scripta, 39, 9–22.

    Xia, Y., Gu, H.-F., Peng, R., Chen, Q., Zheng, Y.-C., Murphy, R.W. & Zeng, X.-M. (2012) COI is better than 16S rRNA for DNAbarcoding Asiatic salamanders (Amphibia: Caudata: Hynobiidae). Molecular Ecology Resources, 12, 48–56.

    Xu, N., Zeng, X.-M. & Fu, J.-Zh. (2007) A new species of the genus Pseudohynobius (Caudata, Hynobiidae) from China. Acta Zootaxonomica Sinica, 32, 230–233. (In Chinese with English abstract)

    Yang, S.-Y., Kim, J.-B., Min, M.-S., Suh, J.-H. & Kang, Y.-J. (2001) Hanguk ui yangseooryu [Monograph of Korean Amphibia], Seoul: Tukpyolsi: Akademi Academybook Sojok, 187 pp. (In Korean)

    Ye, Ch.-Y., Fei, L. & Hu, Sh.-Q. (1993) [Rare and Economic Amphibians of China.] Chengdu: Sichuan Publ. House of Science and Technology, 412 pp. (In Chinese)

    Yoon, I.-B., Lee, S.-J. & Yang, S.-Y. (1996) The study on prey resource and life history of Hynobius leechii Boulenger and Onychodactylus fischeri Boulenger (Amphibia: Salientia). Korean Journal of Environmental Biology, 14, 195–204. (In Korean)

    Yoshikawa, N., Matsui, M. & Nishikawa, K. (2010b) Allozymic variation and phylogeography of two genetic types of Onychodactylus japonicus (Amphibia: Caudata: Hynobiidae) sympatric in the Kinki District, Japan. Zoological Science, 27, 4, 344–355.

    Yoshikawa, N., Matsui, M., Nishikawa, K., Kim, J.-B. & Kryukov, A. (2008) Phylogenetic relationships of the Japanese clawed salamander, Onychodactylus fischeri (Amphibia: Caudata: Hynobiidae), and its congener inferred from the mitochondrial cytochrome b gene. Molecular Phylogenetics and Evolution, 49, 249–259.

    Yoshikawa, N., Matsui, M., Nishikawa, K., Misawa, Y. & Tanabe, S. (2010a) Allozymic variation in the Japanese clawed salamander, Onychodactylus japonicus (Amphibia: Caudata: Hynobiidae), with special reference to the presence of two sympatric genetic types. Zoological Science, 27, 33–40.

    Yoshizawa, S. (1914) [Tailed Amphibians in Etsu (Toyama-ken).] Doubutsugaku zasshi [Zoological Magazine, Tokyo], 26, 451–454. (In Japanese)

    Zeng, X.-M., Fu, J.-Zh., Chen, L., Tian, Y. & Chen, X. (2006) Cryptic species and systematics of the hynobiid salamanders of the Liua Pseudohynobius complex: molecular and phylogenetic perspectives. Biochemical systematics and Ecology, 34, 467–477.

    Zhang, P. & Wake, D.B. (2009) Higher-level salamander relationships and divergence dates inferred from complete mitochondrial genomes. Molecular Phylogenetics and Evolution, 53, 492–508.

    Zhang, P., Chen, Y.-Q., Zhou, H., Liu, Y.-F., Wang, X.-L., Papenfuss, T.J., Wake, D.B. & Qu, L.-H. (2006) Phylogeny, evolution, and biogeography of Asiatic salamanders (Hynobiidae). Proceedings of the National Academy of Sciences of the USA, 103, 19, 7360–7365.

    Zhao, E.-M. & Adler, K. (1993) Herpetology of China. Society for the Study of Amphibians and Reptiles, 522 pp.

    Zhao, E.-M. & Hu, Q.-X. (1988) Studies on Chinese Tailed Amphibians. In: Studies on Chinese Salamanders. Society for the Study of Amphibians and Reptiles, pp. 1-39.

    Zhao, E.-M. (1998) China Red Data Book of Endangered Animals. Amphibia & Reptilia. Beijing, Hong Kong, New York: Science Press, 330 pp.

    Zhao, W.-G., Liu, X.-L. & Fang, J.-J. (1991) [Preliminary study of karyotype of clawed salamander (Onychodactylus fischeri).] Harbin Normal University Journal of Natural Sciences, 7, 2, 78–82. (In Chinese)

    Zhao, Y.-Y., Wang, L.-W., Liang, Ch.-Ch. & Zhang, J.-X. (2004) Preliminary Studies on Anatomy and Histology of Digestive System of Onychodactylus fischeri (Caudata: Hynobiidae). Sichuan Journal of Zoology, 24, 3, 290–293. (In Chinese)

    Zhou, F., Jiang, A.-W. & Jiang, D.-B. (2006) A new species of the genus Hynobius from Guangxi Zhuang Autonomous Region, China (Caudata, Hynobiidae). Acta Zootaxonomica Sinica, 31, 670–674. (In Chinese with English abstract)