Bats (Mammalia: Chiroptera) of the Eastern Mediterranean. Part 3
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Acta Soc. Zool. Bohem. 67: 245–357, 2003 ISSN 1211-376X Bats (Mammalia: Chiroptera) of the Eastern Mediterranean. Part 3. Review of bat distribution in Bulgaria Petr BENDA1, 6), Teodora IVANOVA2), Ivan HORÁČEK3), Vladimír HANÁK3), Jaroslav ČERVENÝ4), Jiří GAISLER5), Antoaneta GUEORGUIEVA2), Boyan PETROV2) & Vladimír VOHRALÍK3) 1) Department of Zoology, National Museum, Václavské nám. 68, CZ–115 79 Praha 1, Czech Republic 2) National Museum of Natural History, blvd. Tzar Osvoboditel 1, Sofia, BG–1000, Bulgaria 3) Department of Zoology, Charles University, Viničná 7, CZ–128 44 Praha 2, Czech Republic 4) Institute of Vertebrate Biology AS CR, Květná 8, CZ–603 65 Brno, Czech Republic 5) Department of Zoology and Ecology, Masaryk University, Kotlářská 2, CZ–611 37 Brno, Czech Republic 6) corresponding author: [email protected] Received October 12, 2003; accepted November 3, 2003 Published December 29, 2003 Abstract. A complete list of all bat records so far available from Bulgaria was compiled from literary references and original data. It is supplemented with distribution maps and a brief summary of the distributional status of each species, tables of measurements of newly recorded bat specimens, and some ecological or taxonomic notes. In total, at least 32 species of bats have been recorded in 2127 localities in Bulgaria, viz., Rhinolophus ferrumequinum (Schreber, 1774) (296 records), R. hipposideros (Bechstein, 1800) (274), R. euryale Blasius, 1853 (104), R. mehelyi Matschie, 1901 (29), R. blasii Peters, 1866 (60), Myotis myotis (Borkhausen, 1797) (184), M. blythii (Tomes, 1857) (117), M. bechsteinii (Kuhl, 1817) (27), M. nattereri (Kuhl, 1817) (34), M. emarginatus (Geoffroy, 1806) (73), M. mystacinus (Kuhl, 1817) or M. alcathoe Helversen et Heller, 2001 (11; plus 23 unspecified records of M. mystacinus s. l.), M. aurascens Kusjakin, 1935 (22), M. brandtii (Eversmann, 1845) (7–9), M. daubentonii (Kuhl, 1817) (36), M. dasycneme (Boie, 1825) (1), M. capaccinii (Bonaparte, 1837) (79), Vespertilio murinus Linnaeus, 1758 (23), Eptesicus serotinus (Schreber, 1774) (79), E. nilssonii (Keyserling et Blasius, 1839) (1), Hypsugo savii (Bonaparte, 1837) (67), Pipistrellus pipistrellus (Schreber, 1774) (28, plus 63 unspecified records of P. pipistrellus s. l.), P. pygmaeus (Leach, 1825) (1), P. nathusii (Keyserling et Blasius, 1839) (28), P. kuhlii (Kuhl, 1817) (9–10), Nyctalus noctula (Schreber, 1774) (92), N. leisleri (Kuhl, 1817) (12–13), N. lasiopterus (Schreber, 1780) (11), Barbastella barbastellus (Schreber, 1774) (23), Plecotus auritus (Linnaeus, 1758) (28), P. austriacus (Fischer, 1829) (101), Miniopterus schreibersii (Kuhl, 1817) (174), and Tadarida teniotis (Rafinesque, 1814) (10). Several other species, such as Plecotus kolombatovici Đulić, 1980 and P. macrobullaris Kuzjakin, 1965, were recorded in closest neighbourhood of the Bulgarian territory and are looked upon as possible candidates for the fauna of Bulgaria. Based on structural differences in bat fauna, the territory of Bulgaria was subdivided into three major faunal regions: (I) Higher mountains (Balkan and Rila-Rhodopes Massifs and adjacent karstic regions, incl. the Mediterranean Struma and Mesta valleys), (II) Karstic lowlands and uplands (parts of Danubian and Upper Thracian Lowlands and the Eastern Balkan Mts.), and (III) the Black Sea region (incl. the Strandža Mts. and Ludogorie Plateau). Composition of the bat fauna shows a strong primary W-E gradient and then only a secondary N-S one. Zoogeography, distribution, fauna, Chiroptera, Balkan peninsula, Bulgaria, Palaearctic region INTRODUCTION The territory of the present Republic of Bulgaria (110,993 km2) covers the central and eastern parts of the Balkan Peninsula and, from the biogeographic point of view, it presents a broad transitional zone between realms of the Mediterranean arboreal and the European mixed forests in south- eastern Europe. The territory of Bulgaria includes various habitat types, from semi-arid steppe and 245 coastal shrub woods to forests and alpine meadows in high mountains (Fig. 1). It lies at the zone of intergradation of the Mediterranean subtropic climate (which essentially influences the south- ern part of the country) and the temperate climates with maximum precipitation in spring and autumn. Bulgaria is adjacent and widely open towards the Black Sea and the Danubian Lowland. A large central area of the country belongs to the belt of chain-folded alpinid mountains of the Carpathian- Balkan system (the Balkan Mts. [= Stara Planina Mts.], 2376 m a. s. l.). The Balkan Mts. build mostly of the Upper Mesozoic carbonate rocks are bordered by old fault mountain system of the Rila-Rhodopes Mts. (with peaks at 2925 m a. s. l.), which core is build by a crystalline rocks. The lowland of the Danube valley completes the relief diversity and builds the possible relay with the vast East European plain while the valleys of rivers Struma, Mesta and Marica broadly incissed into the mountain relief of southern Bulgaria interconnect the region directly with the lowlands of true Mediterranean. About one fourth of the total square of the country lies in the montane altitudinal zone (above 800 m a. s. l.) while at the same time the eastern and northern part of the country is nearly at the altitude of sea level. The effect of extreme declivity, greatly variegated mosaic relief and extensive predominace of carbonate rocks results in intesive a very diversed karstification. The karstic landscape covers about 23% of the country, and frequently takes forms of monumentous rocky cliffs and spacious natural caves (over 4200 caves are known in Bulgaria). The unique geographic position, diverse relief and climate as well as the structurally greatly variegated landscape continuously, for more than 8000 years, extensivelly impacted by postneo- lithic antropogenic rearrangements, set the pattern for the very high biodiversity (cf. Sakaljan & Majni 1993). In full this hold also for the bat fauna of Bulgaria. It should be emphasized, of course, that also amount of information on distribution, faunal and taxonomical status of bats is here much larger than in any other country of the SE Europe and/or SE Mediterranean (comp. Benda & Horáček 1998, Hanák et al. 2001, Uhrin et al. 1996, Kryštufek et al. 1992, 1998). Compared to other parts of the Balkan Peninsula where the research on mammals at the system- atic level did not start until the period between World War I and II or even later (Hanák et al. 2001), the research on the mammals in Bulgaria, including its bat fauna, had started much earlier, already in the late 19th century, that is, at the very beginning of the recent history of the Bulgarian state. The bat fauna of Bulgaria was thoroughly reviewed by Bureš (1917) for the first and by Hanák & Josifov (1959) for the second time. Since that time the amount of data increased almost explosivel- ly. In most species, the currently demonstrated distributional statuses differ much of any previous expectations and in respect to the extent of background information they can be considered quite realistic. In short, comparing to our previous survey on the Eastern Mediterranean bats (Benda & Horáček 1998, Hanák et al. 2001), this one examplifies a region which bat fauna is actually compre- hended into great details and provides the most reliable information on the subject over whole the E-Mediterranean region. Bat research in Bulgaria The first data on the recent mammal fauna of Bulgaria were published by Hristovič (1892). He stressed the very interesting zoogeographical position of the country and listed all known records of mammals, among them, two bat species, viz. Rhinolophus ferrumequinum and Myotis myotis. Later, V. Kovačev, a secondary school teacher, founded a nature collection in the town of Ruse on the basis of which he published records of the following bat species: Rhinolophus ferrumequi- num, R. hipposideros, Myotis myotis, Pipistrellus pipistrellus, Nyctalus noctula, N. leisleri, Ple- cotus auritus, and Miniopterus schreibersii (Kovačev 1894, 1906, 1908). At the beginning of the 20th century Dr. Ivan Bureš started scientific research on Bulgarian caves and founded the bat 246 collection at the National Museum of Natural History in Sofia. In his first paper specially con- cerned with bats of Bulgaria (Bureš 1917), he listed six new species for the country: Rhinolophus euryale, Myotis mystacinus, Eptesicus serotinus, Vespertilio murinus, Pipistrellus nathusii, and Nyctalus lasiopterus, and gave detailed descriptions of the external and dental morphology of all 14 Bulgarian bat species known at that time. Dr. Bureš also assisted the Russian professor P. Bahmet’ev (University of Sofia) who described for the first time the condition of anabiosis in mammals, based on experiments with bats. Bureš (1924, 1925, 1926) was also the first author who stressed the importance of “bat caves” in Bulgaria. In the period 1920–1945 the study of caves and cave fauna continued intensively under his leadership (Atanasov 1936a, b, 1942). In that period Bulgaria was also visited by a number of foreign naturalists who recorded several new bat species as well as new distributional data on already known species as Barbastella barbastellus, Myotis blythii, M. bechsteinii, M. emarginatus, M. capaccinii, and M. daubentonii (Boetticher 1925, Heinrich 1936, Wolf 1940). It was also Dr. I. Bureš who in 1940 applied, for the first time in Bulgaria, the method of bat banding in order to study their migrations (Bureš 1941, 1942). In 1955 a group of