Dmitriy G. Smirnov, Dr.Sc., Professor of the Department of Zoology and Ecology in the Penza State University (440026, Russia, Penza, Krasnaya, 40); e-mail:
Vladimir P. Vekhnik, Senior Researcher, Zhiguli State Nature Biosphere Reserve (445362, Russia, Zhigulevsk, Bakhilova Polyana village,1 Zhigulevskaya street); e-mail:
Gadzhibek S. Dzhamirzoyev, PhD, Senior Researcher, Deputy Director for Science of the Dagestan State Nature Reserve; Senior Researcher of the Institute of Ecology of Mountain Territories of RAS (367010, Russia, Makhachkala, 120 Gagarin Street); e-mail:
Sergey V. Titov, Dr.Sc., Professor, Head of the Department of Zoology and Ecology in the Penza State University (440026, Russia, Penza, Krasnaya Street, 40); e-mail:

Reference to article

Smirnov D.G., Vekhnik V.P., Dzhamirzoyev G.S., Titov S.V. 2020. On the taxonomic status of species from the group «Myotis nattereri» (Chiroptera, Vespertilionidae) in the Eastern Caucasus. Nature Conservation Research 5(4): 30–42.

Section Research articles

In Russia, Myotis nattereri was considered as a monotypic species, of which its range is divided into two parts.. One of them is the northern fragment consisting of Central Russia and the Urals, while the southern part of its range includes the Caucasus and Crimea. On the basis of morphological and molecular-genetic data, the taxonomic status of its Caucasian forms is discussed. In the Eastern Caucasus (Republic of Dagestan, Russia), we caught Myotis animals related to the group of «Myotis nattereri» species, similar to specimens of the West Asian Myotis cf. tschuliensis on the basis of skull size measurements. A comparison analysis showed that, outwardly, M. cf. tschuliensis 2 individuals are highly similar to specimens from Central Russia. However, these animals have some differences in the ear structure, os penis, attachment of the posterior edge of the wing membrane to the foot, and in the ratio of cilia lengths to bristles on the free edge of the tail membrane. The more reliable differences between M. cf. tschuliensis 2 animals and M. nattereri s.str. specimens were revealed on the basis of craniometric studies, but the complete confirmation of the species' independence of Caucasian specimens was obtained using the analysis of mtDNA sequences. According to phylogenetic constructions, Dagestan specimens designated by us as M. cf. tschuliensis 2, formed a well separated sister line in relation to the clade of Caucasian forms: M. schaubi and M. cf. tschuliensis 1 (p-distance: 7.4–7.0%). There are no reliable morphological differences between M. cf. tschuliensis 2 and M. cf. tschuliensis 1. Prior to genetic studies of M. tschuliensis specimens, we suggest using an open nomenclature for the East Caucasian form. If specimens designated as M. cf. tschuliensis 1, originated from the Near East, are genetically identical to the M. cf. tschuliensis specimens from its type location in Turkmenistan, then Dagestan bats designated as M. cf. tschuliensis 2 should be considered as a new, undescribed species.


cryptic species, Myotis cf. tschuliensis, Natterer's bat, Republic of Dagestan, taxonomy

Artice information

Received: 03.04.2020. Revised: 05.08.2020. Accepted: 14.08.2020.

The full text of the article

Artaev O.N., Smirnov D.G. 2016. The bats (Chiroptera; Mammalia) of Mordovia: specific structure and features of distribution. Nature Conservation Research 1(1): 38–51. DOI: 10.24189/ncr.2016.004 [In Russian]
Baker R.J., Bradley R.D. 2006. Speciation in mammals and the genetic species concept. Journal of Mammalogy 87(4): 643–662. DOI: 10.1644/06-MAMM-F-038R2.1
Benda P., Andreas M., Kock D., Lučan R.K., Munclinger P., Nová P., Obuch J., Ochman K., Reiter A., Uhrin M., Weinfurtová D. 2006. Bats (Mammalia: Chiroptera) of the Eastern Mediterranean. Part 4. Bat fauna of Syria: distribution, systematics, ecology. Acta Societatis Zoologicae Bohemicae 70: 1–329.
Benda P., Hanák V., Červený J. 2011. Bats (Mammalia: Chiroptera) of the Eastern Mediterranean and Middle East. Part 9. Bats from Transcaucasia and West Turkestan in collection of the National Museum, Prague. Acta Societatis Zoologicae Bohemicae 75: 159–222.
Benda P., Faizolahi K., Andreas M., Obuch J., Reiter A., Ševčík M., Uhrin M., Vallo P., Ashraf S. 2012. Bats (Mammalia: Chiroptera) of the Eastern Mediterranean and Middle East. Part 10. Bat fauna of Iran. Acta Societatis Zoologicae Bohemicae 76: 163–582.
Benda P., Gazaryan S.V., Vallo P. 2016. On the distribution and taxonomy of bats of the Myotis mystacinus morphogroup from the Caucasus region (Chiroptera: Vespertilionidae). Turkish Journal of Zoology 40: 842–863. DOI: 10.3906/zoo-1505-47
Bogdarina S.V., Strelkov P.P. 2003. Distribution of bats (Chiroptera) in the north of European Russia. Plecotus et al. 6: 7–28. [In Russian]
Bogdanowicz W., Van Den Bussche R.A., Gajewska M., Postawa T., Harutyunyan M. 2009. Ancient and Contemporary DNA Sheds Light on the History of Mouse-Eared Bats in Europe and the Caucasus. Acta Chiropterologica 11(2): 289–305. DOI: 10.3161/150811009X485530
Clement M., Posada D., Crandall K.A. 2000. TCS: a computer program to estimate gene genealogies. Molecular Ecology 9(10): 1657–1659. DOI: 10.1046/j.1365-294x.2000.01020.x
Çoraman E., Dietz C., Hempel E., Ghazaryan A., Levin E., Presetnik P., Zagmajster M., Mayer F. 2019. Reticulate evolutionary history of a Western Palaearctic bat complex explained by multiple mtDNA introgressions in secondary contacts. Journal of Biogeography 46(2): 343–354. DOI: 10.1111/jbi.13509
Darriba D., Taboada G.L., Doallo R., Posada D. 2012. jModelTest 2: more models, new heuristics and parallel computing. Nature Methods 9(8): 772. DOI: 10.1038/nmeth.2109
Frilei C.E. 1947. Preparation and preservation of the baculum of mammals. Journal of Mammalogy 28(4): 395–397. DOI: 10.2307/1375362
García-Mudarra J.L., Ibáñez C., Juste J. 2009. The Straits of Gibraltar: barrier or bridge to Ibero-Moroccan bat diversity. Biological Journal of the Linnean Society 96(2): 434–450. DOI: 10.1111/j.1095-8312.2008.01128.x
Gazaryan S.V. 2002. Ecological-faunal analysis of bats (Chiroptera) in the West Caucasus. PhD Thesis. Moscow. 225 p. [In Russian]
Grimmberger E., Rudolf K. 2009. Atlas der Säugetiere Europas, Nordafrikas and Vorderasian. Münster: Natur und Tier Verl. 495 p.
Hall T.A. 1999. BioEdit: a user-friendly biological sequence alignment editor and analysis program for Windows 95/98/NT. Nucleic Acids Symposium Series 41: 95–98.
Horáček I., Hanak V. 1984. Comments on the systematics and phylogeny of Myotis nattereri (Kuhl, 1818). Myotis 21: 20–29.
Horáček I., Hanák V., Gaisler J. 2000. Bats of the Palearctic region: a taxonomic and biogeographic review. In: B.W. Wołoszyn (Ed.): Proceedings of the VIII European Bat Research Symposium. Vol. 1: Approaches to Biogeography and Ecology of Bats. Kraków: Chiropterological Information Center, Institute of Systematics and Evolution of Animals PAS. P. 11–157.
Ibáñez C., García-Mudarra J.L., Ruedi M., Stadelmann B., Juste J. 2006. The Iberian contribution to cryptic diversity in European bats. Acta Chiropterologica 8(2): 277–297. DOI: 10.3161/1733-5329(2006)8%5B277:TICTCD%5D2.0.CO;2
Ilyin V.Yu., Smirnov D.G., Krasilnikov D.B., Yanaeva N.M. 2002. Materials to the cadastre of bats (Chiroptera) in European Russia and adjacent regions. Penza: Penza State Pedagogical University. 64 p. [In Russian]
Jones G., Parsons S., Zhang S., Stadelmann B., Benda P., Ruedi M. 2006. Echolocation calls, wing shape, diet and phylogenetic diagnosis of the endemic Chinese bat Myotis pequinius. Acta Chiropterologica 8(2): 451–463. DOI: 10.3161/1733-5329(2006)8[451:ECWSDA]2.0.CO;2
Juste J., Ruedi M., Puechmaille S.J., Salicini I., Ibáñez C. 2019. Two new cryptic bat species within the Myotis nattereri species complex (Vespertilionidae, Chiroptera) from the Western Palaearctic. Acta Chiropterologica 20(2): 285–300. DOI: 10.3161/15081109ACC2018.20.2.001
Kerth G., Petrov B., Conti A., Anastasov D., Weishaar M., Gazaryan S., Jaquiéry J., König B., Perrin N., Bruyndonckx N. 2008. Communally breeding Bechstein's bats have a stable social system that is independent from the postglacial history and location of the populations. Molecular Ecology 17(10): 2368–2381. DOI: 10.1111/j.1365-294X.2008.03768.x
Kozhurina E.I. 2009. Conspectus of the Russian bat fauna: systematics and distribution. Plecotus et al. 11–12: 71–105. [In Russian]
Kruskop S.V. 2012. Order Chiroptera. In: I.Ya. Pavlinov, A.A. Lissovsky (Eds.): Mammals of Russia: systematic and geographical handbook. Proceedings of the Zoological Museum of the Moscow State University. Vol. 52. P. 73–126. [In Russian]
Kumar S., Stecher G., Tamura K. 2016. MEGA7: Molecular Evolutionary Genetics Analysis Version 7.0 for Bigger Datasets. Molecular Biology and Evolution 33(7): 1870–1874. DOI: 10.1093/molbev/msw054
Kuzyakin A.P. 1935. New data on the taxonomy and geographical distribution of bats (Chiroptera) in USSR. Bulletin of Moscow Society of Naturalists 44(7–8): 428–438. [In Russian]
Kuzyakin A.P. 1950. Bats. Moscow: Sovetskaya nauka. 443 p. [In Russian]
Leigh J.W., Bryant D., Nakagawa S. 2015. POPART: full-feature software for haplotype network construction. Methods in Ecology and Evolution 6(9): 1110–1116. DOI: 10.1111/2041-210x.12410
Mayer F., Dietz C., Kiefer A. 2007. Molecular species identification boosts bat diversity. Frontiers in Zoology 4: 4. DOI: 10.1186/1742-9994-4-4
Ognev S.I. 1928. Mammals of East Europe and Northern Asia. Vol. 1. Moscow; Leningrad: Gospolitizdat. 631 p. [In Russian]
Pavlinov I.Ya., Kruskop S.V., Varshavskiy A.A., Borisenko A.V. 2002. Terrestrial mammals of Russia. Handbook-determinant. Moscow: KMK Scientific Press Ltd. 298 p. [In Russian]
Puechmaille S.J., Allegrini B., Boston E.S., Dubourg-Savage M.J., Evin A., Knochel A., Le Bris Y., Lecoq V., Lemaire M., Rist D., Teeling E.C. 2012. Genetic analyses reveal further cryptic lineages within the Myotis nattereri species complex. Mammalian Biology 77(3): 224–228. DOI: 10.1016/j.mambio.2011.11.004
Red Data Book of the Republic of Crimea. Animals. Simferopol: IT ARIAL, 2015. 440 p. [In Russian]
Ronquist F., Huelsenbeck J.P. 2003. MRBAYES 3: Bayesian phylogenetic inference under mixed models. Bioinformatics 19(12): 1572–1574. DOI: 10.1093/bioinformatics/btg180
Ruedi M., Mayer F. 2001. Molecular systematics of bats of the genus Myotis (Vespertilionidae) suggests deterministic ecomorphological convergences. Molecular Phylogenetics and Evolution 21(3): 436–448. DOI: 10.1006/mpev.2001.1017
Salicini I., Ibáñez C., Juste J. 2011. Multilocus phylogeny and species delimitation within the Natterer's bat species complex in the Western Palearctic. Molecular Phylogenetics and Evolution 61(3): 888–898. DOI: 10.1016/j.ympev.2011.08.010
Salicini I., Ibáñez C., Juste J. 2013. Deep differentiation between and within Mediterranean glacial refugia in a flying mammal, the Myotis nattereri bat complex. Journal of Biogeography 40(6): 1182–1193. DOI: 10.1111/jbi.12062
Sambrook J., Fritsch E.F., Maniatis T. 1989. Molecular Cloning: A Laboratory Manual. New York: Cold Spring Harbor Laboratory Press. 1546 p.
Sikes R.S., Gannon W.L. 2011. Guidelines of the American Society of Mammalogists for the use of wild mammals in research. Journal of Mammalogy 92(1): 235–253. DOI: 10.1644/10-MAMM-F-355.1
Simmons N.B. 2005. Order Chiroptera. In: D.E. Wilson, D.M. Reeder (Eds.): Mammal species of the World: a taxonomic and geographic reference. Baltimore: Johns Hopkins University Press. P. 312–529.
Smirnov D.G. 2000. Variability of bakulum in bats (Chiroptera, Vespertilionidae) in the Middle Volga region and adjacent areas. Plecotus et al. 3: 20–34. [In Russian]
Smirnov D.G., Dzhamirzoev G.S., Gazaryan S.V., Vekhnik V.P., Bykov Yu.A. 2019a. Bats (Chiroptera) of Dagestan: review of fauna resulted from research in 2017–2019. Plecotus et al. 22: 3–48. [In Russian]
Smirnov D.G., Vekhnik V.P., Dzhamirzoev G.S., Bykov Yu.A. 2019b. New data on the distribution of bats (Chiroptera) in the flat part of the Republic of Dagestan. Russian Journal of Ecosystem Ecology 4(4). DOI: 10.21685/2500-0578-2019-4-5 [In Russian]
Snitko V.P. 2011. Naterrer's bat (Myotis nattereri) at its eastern range boundary (the Urals). Zoologicheskii Zhurnal 90(10): 1245–1254. [In Russian]
Topál G. 1958. Morphological studies on the os penis of bats in the Carpathian Basin. Annales Musei Historico-Naturalis Hungarici 50(Series nova 9): 331–342.
White J. 1951. A Practical Method for Mounting the Bacula of Small Mammals. Journal of Mammalogy 32(1): 125. DOI: 10.1093/jmammal/32.1.125