A New Species of Common Toads from the Talysh Mountains, South-Eastern Caucasus: Genome Size, Allozyme, and Morphological Evidences

Spartak N. Litvinchuk, Leo J. Borkin, Dmitry V. Skorinov, Jury M. Rosanov

Abstract


The genome size, protein (24 loci) and morphological variation in three currently recognized species of the Bufo bufo group was studied. Among common toads of the Caucasian Isthmus, two distinct groups were revealed. The main territory of the Caucasian region, including the Great Caucasian Range and the Caucasus Minor, is inhabited by the Caucasian toad, B. verrucosissimus, whereas the Talysh Mountains, geographically isolated in the south-east of the region, harbor another toad. Genetic distances between these forms were higher (D = 0.411) than that between B. verrucosissimus and European B. b. bufo (D = 0.138). Based on genome size and allozyme data, we recognized the Talysh toad as a new distinct species, Bufo eichwaldi, sp. nov. The time of origin of the species estimated on the basis of molecular data, roughly corresponds to relict character of the Hyrcanian forest of the Talysh region (the late Miocene). West Palearctic species (B. bufo, B. verrucosissimus, B. eichwaldi, sp. nov.) form joint cluster which is separated from another phylogenetic lineage with the Far Eastern B. gargarizans. Taxonomic status of B. verrucosissimus and B. b. spinosus is discussed. The Caucasian toad was treated tentatively as an allopatric species in respect to B. b. bufo of boreal Europe. Common toads of Anatolian Turkey previously recognized as B. b. spinosus were assigned to B. verrucosissimus. The Balkan common toads seems to be a distinct taxon which is the sister lineage of the B. verrucosissimus cluster. Toads from Apennine Peninsula were allocated to distinct subspecies B. b. palmarum. North African (Tunisian) B. b. spinosus was separated as a distinct phyletic lineage forming with B. eichwaldi, sp. nov., the sister group of other West Palearctic common toads. Further resolution of the taxonomic position of that and some other members of the Bufo bufo group needs careful examination of samples from Mediterranean area, including the type locality of B. b. spinosus.

Keywords


genome size; nuclear DNA content; allozymes; the Caucasus; taxonomy; Bufo bufo; Bufo bufo spinosus; Bufo eichwaldi sp. nov.; Bufo verrucosissimus; Bufo gargarizans

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References


Alekperov A. M. (1951), «Amphibia», in: A. N. Alizade, S. M. Asadov, and A. N. Derzhavin (eds.), Zhivotnyi Mir Azerbaydzhana [Animals of Azerbaijan], Izd. AN AzSSR, Baku, pp. 203 – 206 [in Russian].

Alekperov A. M. (1978), Zemnovodnye i Presmykayushchiesya Azerbaydzhana [Amphibians and Reptiles of Azerbaijan], Elm, Baku [in Russian].

Anderson S. C. (1985), «Amphibians», in: E. Yarshater (ed.), Encyclopaedia Iranica. Vol. 1. Fasc. 9, Routledge and Kegan Paul, London – Boston – Melbourne – Hensley, pp. 987 — 990.

Anderson S. C. (1999), The Lizards of Iran, Soc. for the Study of Amphibians and Reptiles, Ithaca, NY.

Avise J. C. and Aquadro C. F. (1982), «A comparative summary of genetic distances in the vertebrates. Patterns and correlations», in: Evolutionary Biology. Vol. 15, Plenum Press, New York – London, pp. 151 – 185.

Bachmann K. (1970), «Specific nuclear DNA amounts in toads of the genus Bufo», Chromosoma, 29, 365 – 374.

Bachmann K. (1972), «Nuclear DNA and developmental rate in frogs», Quart. J. Florida Acad. Sci., 35, 225 – 231.

Bachmann K., Konrad A., Oeldorf E., and Hemmer H. (1977), «Genome size in the green toad (Bufo viridis) group», Experientia, 34, 331 – 332.

Bachmann K. and Nishioka M. (1978), «Genome size and nuclear size in Palaearctic frogs (Rana)», Copeia, 1978, 225 – 229.

Baloutch M. and Kami H. G. (1995), [Amphibians of Iran], Univ. Publ. Tehran, Tehran [in Farsi].

Bannikov A. G., Darevsky I. S., Rustamov A. K., and Szczerbak N. N. (1977), Opredelitel’ Zemnovodnykh i Presmykayushchikhsya Fauny SSSR [Guide Book of Amphibians and Reptiles of the USSR Fauna], Prosveshchenie, Moscow [in Russian].

Birschtein V. J. and Mazin A. L. (1982), «Chromosomal polymorphism of Bufo bufo: karyotype and C-banding pattern of B. b. verrucosissima», Genetica, 59, 93 – 98.

Blair W. F. (1972), «Evidence from hybridization», in: W. F. Blair (ed.), Evolution in the Genus Bufo, Univ. of Texas, Austin – London, pp. 196 – 232.

Blanford W. T. (1876), Eastern Persia. An Account of the Journeys of the Persian Boundary Commission 1870 – 71 – 72. Vol. II. The Zoology and Geology, Macmillan and Co., London.

Bobrinsky N. A. and Gladkov N. A. (1961), Geografiya Zhivotnykh [Animal Geography], 2nd Edition, Moscow [in Russian].

Boettger O. (1886), «Die Reptilien und Amphibien des Talysch-Gebietes. Nach den neuesten Materialien bearbeitet», in: G. Radde, Die Fauna und Flora des südwestlichen Caspi-Gebietes. Wissenschaftliche Beiträge zu den Reisen an der Persisch-Russischen Grenze, F. A. Brockhaus, Leipzig, pp. 30 – 81.

Boettger O. (1892), Katalog der Batrachier-Sammlung im Museum der Senckenbergischen Naturforschenden Gesellschaft in Frankfurt am Main, Gebrüder Knauer, Frankfurt am Main.

Boettger O. (1899), «Reptilia et Batrachia», in: Die Sammlungen des Kaukasischen Museums, im Vereine mit Special-Gelehrten Bearbeitet, und Herausgegeben von Dr. Gustav Radde. Bd. 1. Zoologie, Tiflis, S. 271 – 290 [in Russian and German].

Borkin L. J. (1984), «European – Far Eastern disjunctions in amphibian distributions: new analysis of the problem», in: L. J. Borkin and N. B. Ananjeva (eds.), Ecology and Faunistics of Amphibians and Reptiles of the USSR and Adjacent Countries. Proc. Zool. Inst., USSR Acad. Sci., Leningrad, Vol. 124, pp. 55 – 88 [in Russian].

Borkin L. J. (1986), «Pleistocene glaciations and western – eastern Palearctic disjunction in amphibian distribution», in: Z. Roček (ed.), Studies in Herpetology. Proc. Eur. Herpetol. Meeting, Charles Univ., Prague, pp. 63 – 66.

Borkin L. J. (1987), «On the systematics and zoogeography of amphibians of the Caucasus», in: L. J. Borkin (ed.), Herpetological Investigations in the Caucasus. Proc. Zool. Inst., USSR Acad. Sci., Leningrad, Vol. 158 for 1986, pp. 47 – 58 [in Russian].

Borkin L. J. (1998), «Amphibia», in: N. B. Ananjeva, L. J. Borkin, I. S. Darevsky, and N. L. Orlov, Zemnovodnye i Presmykayushchiesya [Amphibians and Reptiles], The Encyclopedia of Nature of Russia, ABF, Moscow, pp. 19 – 174 [in Russian].

Borkin L. J. (1999), «Distribution of amphibians in North Africa, Europe, Western Asia, and the former Soviet Union», in: W. E. Duellman (ed.), Patterns of Distribution of Amphibians: a Global Perspective, The Johns Hopkins Univ. Press, Baltimore – London, pp. 329 – 420.

Borkin L. J. and Darevsky I. S. (1987), «The list of amphibians and reptiles of the USSR fauna», in: I. S. Darevsky and V. G. Krever (eds.), Amphibii i Reptilii Zapovednykh Territoriy [Amphibians and Reptiles of Protected Territories], Moscow, pp. 128 – 143 [in Russian].

Borkin L. J., Korshunov A. V., Lada G. A., Litvinchuk S. N., Rosanov J. M., Shabanov D. A., and Zinenko A. I. (2004), «Mass occurrence of polyploidy green frogs (Rana esculenta complex) in eastern Ukraine», Russ. J. Herpetol., 11(3), 194 – 213.

Borkin L. J. and Litvinchuk S. N. (2008), «Genetic distance and speciation in amphibians», in: N. B. Ananjeva and G. A. Lada (eds.), The Problems of Herpetology, St. Petersburg, pp. 41 – 52 [in Russian, with English abstract].

Borkin L. J., Litvinchuk S. N., Rosanov J. M., and Milto K. D. (2001), «Cryptic speciation in Pelobates fuscus (Anura, Pelobatidae): evidence from DNA flow cytometry», Amphibia–Reptilia, 22(4), 387 – 396.

Borkin L. J., Shabanov D. A., Brandler O. V., Kukushkin O. V., Litvinchuk S. N., Mazepa G. A., and Rosanov J. M. (2007), «A case of natural triploidy in European diploid green toad (Bufo viridis), with some distributional records of diploid and tetraploid toads», Russ. J. Herpetol., 14(2), 121 – 132.

Borkin L. J. and Roshchin V. V. (1981), «Electrophoretic comparison of proteins between European and Far Eastern toads of the Bufo bufo complex», Zool. Zh., 60(12), 1802 – 1812 [in Russian].

Busack S. D. (1986), «Biogeographic analysis of the herpetofauna separated by the formation of the strait of Gibraltar», Natl. Geogr. Res., 2(1), 17 – 36.

Cavalier-Smith T. (1985), «Introduction: the evolutionary significance of genome size», in: T. Cavalier-Smith (ed.), The Evolution of Genome Size, John Wiley and Sons, Chichester, pp. 1 – 36.

Cavalli-Sforza L. L. and Edwards A. W. F. (1967), «Phylogenetic analysis: Model and estimation procedures», Am. J. Hum. Genet., 19, 233 – 257.

Chkhikvadze V. M. (1984), «Review of fossil urodeles and anurans of the USSR», Izv. AN GrSSR Ser. Biol., 10(1), 5 – 13 [in Russian].

Cuvier G. (1829), Le Règne Animal distribué d’après son organisation, pour servir de base à l’histoire naturelle des animaux et d’introduction à l’anatomie comparée. Nouvelle édition, revue et augmentée. Tome II, Paris, chez Déterville.

Darevsky I. S. (1987), «Protection of amphibians and reptiles in reserves of the Caucasus», in: I. S. Darevsky and V. G. Krever (eds.), Amphibii i Reptilii Zapovednykh Territoriy [Amphibians and Reptiles of Protected Territories], Moscow, pp. 85 – 101 [in Russian].

Daudin F. M. An XI [1803], Histoire Naturelle, génerale et particulière des Reptiles. T. 8, Imprimerie de F. Dufart, Paris.

De Lange L. (1973), «A contribution to the intraspecific systematics of Bufo bufo (Linnaeus, 1758) (Amphibia)», Beaufortia. Ser. Misc. Publ., 21(280), 99 – 116.

De Smet W. H. O. (1981), «The nuclear Feulgen-DNA content of the vertebrates (especially reptiles), as measured by fluorescence cytophotometry, with notes on the cell and chromosome size», Acta Zool. Pathol. Antverp., 76, 119 – 167.

Dwigubsky I. (1832), Opyt Estestvennoy Istorii vsekh Zhivotnykh Rossiyskoy Imperii. Gady, ili zhivotnye presmykayushchiesya [An Attempt of Natural History of All Animals of Russian Empire. Reptiles], Izd. Mosk. Univ., Moscow.

Eichwald E. (1831), Zoologia specialis quam expositis animalibus tum vivis, tum fossibilibus potissimum Rossiae in universum, et Poloniae in specie, in usum lectionum publicorum in Universitate Caesare Vilnensi habendarum edidit D. Eduardus Eichwald, P. P. O. [T. 3], Pars posterior specialem expositionem Spondylozoorum continens, Typis Josephi Zawadski, Vilnae.

Eichwald E. (1841), Fauna Caspio-Caucasia nonnullis observationibus novis illustravit. Litteris typographiae diarii gall, Politic. Petropol., Petropoli.

Eiselt J. and Schmidtler J. F. (1973), «Froschlurche aus dem Iran unter Berücksichtigung außeriranischer Populationsgruppen», Ann. Naturhist. Mus. Wien, 77, 181 – 243.

Engelmann W.-E., Fritzsche J., Günther R., and Obst F. J. (1993), Lurche und Kriechtiere Europas, 2. Auf. Neumann, Radebeul.

Felsenstein J. (2004), PHYLIP v. 3.6b, Univ. Washington, Seattle.

Frost D. R., Grant T., Faivovitch J. N., Bain R. H., Haas A., Haddad C. L. F. B., De Sa R. O., Channing A., Wilkinson M., Donnellan S. C., Raxworthy C. J., Cambell J. A., Blotto B. L., Moler P., Drewes R. C., Nussbaum R. A., Lynch J. D., Green D. M., and Wheeler W. C. (2006), «The amphibian tree of life», Bull. Am. Mus. Nat. Hist., 297, 1 – 370.

Frost D. R. and Hillis D. M. (1990), «Species in concept and practice: herpetological applications», Herpetologica, 46, 87 – 104.

Garsia-Paris M., Montori A., and Herrero P. (2004), Fauna Iberica. Vol. 24. Amphibia, Lissamphibia, Museo Nacional de Ciencias Naturales, Consejo Superior de Investigaciones Científicas, Madrid.

Grossheim A. A. (1926), Flora Talysha [The Flora of Talysh], Tiflis [in Russian].

Harrison T. M., Copeland P., Kidd W. S. F., and Yin A. (1992), «Raising Tibet», Science, 255, 1663 – 1670.

Hemmer H. (1975), «Serumeiweissunterschied zweischen B. bufo bufo und B. b. spinosus (Salentia, Bufonidae)», Salamandra, 11, 61 – 63.

Hemmer H. and Böhme W. (1976), «Zwischenbericht über die innerartliche Variabilität der Erdkröte (Bufo bufo L.) (Amphibia: Salentia: Bufonidae)», Salamandra, 12, 194 – 201.

Highton R. (1989), «Biochemical evolution in the slimy salamanders of the Plethodon glutinosus complex in the eastern United States», Univ. Illinois Biol. Monogr., 57, 1 – 78.

Highton R. (1990), «Taxonomic treatment of genetically differentiated populations», Herpetologica, 46(1), 114 – 121.

Igawa T., Kurabayashi A., Nishioka M., and Sumida M. (2006), «Molecular phylogenetic relationship of toads distributed in the Far East and Europe inferred from the nucleotide sequences of mitochondrial DNA genes», Mol. Phyl. Evol., 38, 250 – 260.

Inger R. F. (1972), «Bufo of Eurasia», in: W. F. Blair (ed.), Evolution in the Genus Bufo, Univ. Texas Press, Austin – London, pp. 102 – 118.

Kessler K. (1878), Puteshestviye po Zakavkazskomu Krayu v 1875 Godu s Zoologicheskoy Tsel’yu [Travels in the Transcaucasia in 1875 for Zoological Purpose], Trudy S.-Petersburg. Obshch. Estestvoisp. Issue 8. Pril., St. Petersburg [in Russian].

Kirichenko A. N. (1910), «Report about trip on the Talysh in 1909 for collecting for Zoological Museum of Imperial Academy of Sciences», Ann. Mus. Zool. Acad. Imp. Sci. St.-Pétersbourg, 15, 0139 – 0152 [in Russian].

Krassowsky D. B. (1933), «[Materials on the study of amphibians and reptiles in the Caucasian State Reserve]», Nachrichten des 2-en Nord-Kaukasus Pädagogischen Instituts Gadief Namens, Ordjonikidze, 10, 89 95 [in Russian].

Kumar S., Tamura K., Jakobsen I. B., and Nei M. (2001), MEGA2: Molecular Evolutionary Genetics Analysis software, Arizona State Univ., Tempe.

Kutrup B., Yilmaz N., Canakci S., Belduz A. O., and Doglio S. (2006), «Intraspecific variation of Bufo bufo, based on 16S Ribosomal RNA sequences», Amphibia–Reptilia, 27, 268 – 273.

Kuzmin S. L. (1999), Zemnovodnye Byvshego SSSR [The Amphibians of the Former Soviet Union], KMK, Moscow [in Russian].

Licht L. E. and Lowcock L. A. (1991), «Genome size and metabolic rate in salamanders», Comp. Biochem. Physiol., 100B, 83 – 92.

Litvinchuk S. N., Borkin L. J., and Rosanov J. M. (2004), «Intra- and interspecific genome size variation in hynobiid salamanders of Russia and Kazakhstan: determination by flow cytometry», Asiatic Herpetol. Res., 10, 282 – 294.

Litvinchuk S. N., Borkin L. J., Rosanov J. M., and Skorinov D. V. (2006), «Allozyme and genome size variation in tree frogs from the Caucasus, with description of a new subspecies Hyla arborea gumilevskii from the Talysh Mountains», Russ. J. Herpetol., 13(3), 183 – 206.

Litvinchuk S. N., Borkin L. J., Rosanov J. M., Skorinov D. V., Khalturin M. D., Dћukić G., Kalezić M. L., and Mazanaeva L. F. (2005a), «Geographic differentiation in newts (Triturus) of eastern Europe: genome size, allozymes, and morphology», in: N. Ananjeva and O. Tsinenko (eds.), Herpetologia Petropolitana, St. Petersburg, pp. 51 – 54.

Litvinchuk S. N., Rosanov J. M., and Borkin L. J. (2001a), «Natural autotriploidy in the Danube newt, Triturus dobrogicus (Salamandridae)», Russ. J. Herpetol., 8(1), 74 – 76.

Litvinchuk S. N., Rosanov J. M., Borkin L. J., Khalturin M. D., Timofeev B. I., Dћukić G., and Kalezić M. L. (2001b), «Genome size and some problems of urodelan systematics (Salamandridae and Hynobiidae)», in: N. B. Ananjeva et al. (eds.), The Problems of Herpetology, Pushchino – Moscow, pp. 168 – 170 [in Russian, with English abstract].

Litvinchuk S. N., Rosanov J. M., Borkin L. J., and Skorinov D. V. (2008), «Molecular, biochemical and cytogenetic aspects of microevolution in anurans of Russia and adjacent countries», in: N. B. Ananjeva and G. A. Lada (eds.), The Problems of Herpetology, St. Petersburg, pp. 247 – 257 [in Russian, with English abstract].

Litvinchuk S. N., Zuiderwijk A., Borkin L. J., and Rosanov J. M. (2005b), «Taxonomic status of Triturus vittatus (Amphibia: Salamandridae) in western Turkey: trunk vertebrae count, genome size and allozyme data», Amphibia– Reptilia, 26(3), 305 – 323.

Lizana M., Marquez R., Martin-Sanchez R., Ciudad J., Lopez A., and Orfao A. (2000), «Determination of cellular DNA content of Iberian salamanders by flow cytometry», Amphibia–Reptilia, 21, 411 – 418.

Lüscher B., Grossenbacher K., and Scholl A. (2001), «Genetic differentiation of the common toad (Bufo bufo) in the Swiss Alps», Amphibia–Reptilia, 22, 141 – 154.

McCulloch R. D., Upton D. E., and Murphy R. W. (1996), «Trends in nuclear DNA content among amphibians and reptiles», Comp. Biochem. Physiol., 113B, 601 – 605.

Macey J., Schulte J., Larson A., Fang Z., Wang Y., Tuniyev B., and Papenfuss T. (1998), «Phylogenetic relationships of toads in the Bufo bufo species group from the eastern escarpment of the Tibetan Plateau: a case of vicariance and dispersal», Mol. Phyl. Evol., 9, 80 – 87.

Matsui M. (1986), «Geographic variation in toads of the Bufo bufo complex from the Far East, with a description of a new subspecies», Copeia, 1986(3), 561 – 579.

Matsui M., Seto T., Kohsaka Y., and Borkin L. J. (1985), «Bearing of chromosome C-banding pattern on the classification of Eurasian toads of the Bufo bufo complex», Amphibia–Reptilia, 6(1), 23 – 33.

Mazin A. L. (1980), «Amounts of nuclear DNA in anurans of the USSR», Experientia, 36, 190 – 191.

Ménétries E. (1832), Catalogue raisonné des Objets de Zoologie recueillis dans un Voyage au Caucase et jusquaux Frontières actuelles de la Perse entrepris par Ordre de S. M. LEmpereur, Imprim. Acad. Imp. Sci., St.-Pétersbourg.

Mertens R. and Wermuth H. (1960), Die Amphibien und Reptilien Europas (Dritte Liste, nach dem Stand vom 1. Januar 1960), Waldemar Kramer, Frankfurt am Main.

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

Nei M. (1978), «Estimation of average heterozygosity and genetic distance from a small number of individuals», Proc. Natl. Acad. Sci., 70, 3321 – 3323.

Nikolsky A. M. (1905), Herpetologia Rossica. Mém. l’Acad. Imp. Sci. St.-Pétersbourg. VIIIe Sér. Classe physico-mathématique, Vol. 17. No. 1 [in Russian].

Nikolsky A. M. (1913), Herpetologia Caucasica, The Caucasian Museum, Tiflis [in Russian].

Nikolsky A. M. (1916), Reptilia. Vol. II. Ophidia. Faune de la Russie et des Pays limitrophes, Imp. Acad. Sci., Petrograd [in Russian].

Nikolsky A. M. (1918), Amphibia. Faune de la Russie et des Pays limitrophes, Acad. Sci. Russie, Petrograd [in Russian].

Nishioka M., Sumida M., Ueda H., and Wu Z. (1990), «Genetic relationships among 13 Bufo species and subspecies by the method of electrophoretic analysis», Sci. Rep. Lab. Amphibian Biol., Hiroshima Univ., 10, 53 – 91.

Oeldorf E., Nishioka M., and Bachman K. (1978), «Nuclear DNA amounts and development rate in Holarctic Anura», Zeitschr. Zool. Syst. Evolutionforsch., 16, 216 – 224.

Olmo E. (1973), «Quantitative variations in the nuclear DNA and phylogenesis of the Amphibia», Caryologia, 26(1), 43 – 68.

Olmo E. (1983), «Nucleotype and sell size in vertebrates: a review», Basic Appl. Histochem., 27, 227 – 256.

Orlova V. F. and Tuniev B. S. (1989), «On the taxonomy of the Caucasian common toads belonging to the group Bufo bufo verrucosissimus (Pallas) (Amphibia, Anura, Bufonidae)», Bull. Mosk. Obshch. Isp. Prirody Ser. Biol., 94(3), 13 – 24 [in Russian, with English abstract].

Pallas P. S. (1814), Zoographia Rosso-Asiatica, sistens omnium animalium in extenso Imperio Rossico et adjacentibus maribus observatorum recensionem, domicilia, mores et descriptiones, anatomen atque icones plurimorum. Tomus III. Animalia monocardia seu frigidi sanguinis Imperii Rosso-Asiatici. Ex officina Caes, Academiae Scientarum, Petropoli.

Pisanets E. M. (2001), «Taxonomic relationships of the common toads (Bufo bufo complex) and some theoretical and practical problems of systematics. Communication 1», Vestn. Zool. Kiev, 35(5), 37 – 44 [in Russian, with English abstract].

Pisanets E. M. (2002), «Taxonomic relationships of the common toads (Bufo bufo complex) and some theoretical and practical problems of systematics. Communication 2», Vestn. Zool. Kiev, 36(1), 61 – 68 [in Russian, with English abstract].

Pryalkina T. I. (1985), «Comparative analyses of miogene specters of some species and subspecies of genus Bufo», in: I. S. Darevsky (ed.), The Problems of Herpetology, Leningrad, p. 173 [in Russian].

Pryalkina T. I. (1989), Study of Systematic Relationships between Anurans of the USSR Fauna with Use of Comparative Analysis of Proteins. Authors Abstract of Candidates Thesis, Izd. Zool. Inst. AN SSSR, Leningrad [in Russian].

Rosanov J. M. and Vinogradov A. E. (1998), «Precise DNA cytometry: investigation of individual variability in animal genome size», Tsitologiya, 40, 792 – 800 [in Russian].

Saitou N. and Nei M. (1987), «The neighbor-joining method: a new method for reconstructing of phylogenetic trees», Mol. Biol. Evol., 4, 406 – 425.

Sanchiz B. (1998), Salientia. Handbuch der Paläoherpetologie. Teil 4, Verlag Dr. Friedrich Pfeil, München.

Schmidtler J. J. and Schmidtler J. F. (1969), «Über Bufo surdus; mit einem Schlüssel und Anmerkungen zu den übrigen Kröten Irans und West-Pakistans», Salamandra, 5, 113 – 123.

Schreiber E. (1875), Herpetologia europaea. Eine systematische Bearbeitung der Amphibien und Reptilien, welche bisher in Europa aufgefunden sind, Druck und Verlag von Friedrich Vieweg und Sohn, Braunschweig.

Schreiber E. (1912), Herpetologia europaea. Eine systematische Bearbeitung der Amphibien und Reptilien, welche bisher in Europa aufgefunden sind. 2. Auflage, Friedrich Vieweg und Sohn, Braunschweig.

Shcherbak N. N. (1982), «Grundzüge einer herpetogeographischen Gliederung der Paläarktis», Vertebr. Hung., 21, 227 – 239.

Sobolevsky N. I. (1929), «The herpetofauna of the Talysh and the Lenkoran Lowland», Mem. Soc. Amis Sci. Nat. d’Antrop. d’Ethnogr. Moscou, 5, 1 – 143 [in Russian].

Sokal R. R. and Sneath P. H. A. (1963), Principles of Numerical Taxonomy, Freeman, San Francisco.

Swofford D. L. and Selander R. B. (1981), BIOSYS-1. A Computer Program for the Analysis of Allelic Variation in Population Genetics and Biochemical Systematics, Illinois Nat. Hist. Survey, Champaigh.

Takhtajan A. (1978), Floristicheskiye Oblasti Zemli [The Floristic Regions of the World], Nauka, Leningrad [in Russian].

Tarkhnishvili D. N. and Gokhelashvili R. K. (1999), «The Amphibians of the Caucasus», Adv. Amphibian Res. Former Soviet Union, 4, 1 – 233.

Terentjev P. V. and Chernov S. A. (1936), Kratkiy Opredelitel Zemnovodnykh i Presmykayushchikhsya SSSR [Brief Guide Book of Amphibians and Reptiles of the USSR], Moscow – Leningrad [in Russian].

Terentjev P. V. and Chernov S. A. (1949), OpredelitelPresmykayushchikhsya i Zemnovodnykh [Guide Book of Reptiles and Amphibians], 3rd Edition, Sovetskaya Nauka, Moscow – Leningrad [in Russian].

Tosunoğlu M. and Taşkavak E. (2001), «A serological investigation of the Bufo bufo (Anura, Bufonidae) populations in Southern Marmara (Manyas, Balikesir) and Eastern Black Sea (Camlihemşin, Rize) regions», Ital. J. Zool., 68, 165 – 168.

Tuck R. G. (1975), «Bufo bufo (Common toad)», Herpetol. Rev., 6, 115.

Tuniyev B. (1999 [1998]), «A systematic list of amphibians from the Alpides of the Caucasus and Middle Asia, with comments on their taxonomy», Adv. Amphibian Res. Former Soviet Union, 3, 43 — 75.

Veith M. (1996), «Molecular markers and species delimitation: examples from the European batrachofauna», Amphibia–Reptilia, 17, 303 – 314.

Velieva Z. D. (1975), Fauna and Ecology of Amphibia of Southeastern Azerbaijan. Author’s Abstract of Candidate’s Thesis, Izd. AzGPI, Baku [in Russian].

Velieva Z. D. (1981), «On new records and ecology of the common toad in Azerbaijan», in: I. S. Darevsky (ed.), The Problems of Herpetology, Leningrad, p. 30 [in Russian].

Vialli M. (1957), «Volume et contenu en ADN par noyau», Exp. Cell Res., Suppl. 4, 284 – 293.

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

Wake D. B. (1981), «The application of allozyme evidence to problems in the evolution of morphology», in: Evolution Today. Proc. 2nd Int. Congr. Syst. Evol. Biol., Carnegue-Mellon Univ. Press., Pittsburgh, pp. 257 – 270.

Wright S. (1978), Variability Within and Among Natural Populations, Univ. Chicago Press, Chicago.

Ullerich F.-H. (1966), «Karyotyp und DNS-Gehalt von Bufo bufo, B. viridis, B. bufo × B. viridis und B. calamita (Amphibia, Anura)», Chromosoma, 18, 316 – 342.

Ullerich F.-H. (1970), «DNA-Gehalt und Chromosomenstruktur bei Amphibien», Chromosoma, 30, 1 – 37.

Yilmaz I. and Kumlutas Y. (1995), «Turkiye’de Yasayan Bufo bufo (Linnaeus) 1758’nun dagilisi ve taksonomik durumu hakkinda bir inceleme», Tur. J. Zool., 19, 277 – 284.

Zhordania R. G. (1960), «List of collections of amphibians of Zoological Department of S. N. Dzhanashia State Museum of Georgia AN GrSSR», Trudy Gos. Muz. Gruzii, 20-A, 159 – 179 [in Russian].




DOI: https://doi.org/10.30906/1026-2296-2008-15-1-19-43

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