Bats (Mammalia: Chiroptera) of the Eastern Mediterranean and Middle East
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Acta Soc. Zool. Bohem. 79: 267–458, 2015 ISSN 1211-376X Bats (Mammalia: Chiroptera) of the Eastern Mediterranean and Middle East. Part 12. Bat fauna of Afghanistan: revision of distribution and taxonomy* Petr Benda & Jiří Gaisler† Department of Zoology, National Museum (Natural History), Václavské nám. 68, CZ–115 79 Praha 1, Czech Republic; e-mail: [email protected] Department of Zoology, Faculty of Science, Charles University in Prague, Viničná 7, CZ–128 44 Praha 2, Czech Republic † deceased. Received 20 October 2015; accepted 25 November 2015 Published 21 December 2015 Abstract. A complete list of bat records available from Afghanistan was compiled from literature and from examination of museum specimens. The record review is complemented by distribution maps, summaries of distributional status of the particular species, supplemented by observations on morphology and evaluation of taxonomic status of the Afghanistani populations. From the territory of Afghanistan, at least 224 records of 40 bat species belonging to eight families are known; viz. Rhinopoma microphyllum (Brünnich, 1782) (15 record sites), R. muscatellum Thomas, 1903 (4), R. hardwickii Gray, 1831 (10), Lyroderma lyra (Geoffroy, 1810) (2), Rhinolophus ferrumequinum (Schreber, 1774) (20), R. bocharicus Kaŝenko et Akimov, 1918 (5), R. hipposideros (Borkhausen, 1797) (7), R. lepidus Blyth, 1844 (6), R. blasii Peters, 1867 (10), Hipposideros fulvus Gray, 1838 (2), Asellia tridens (Geoffroy, 1813) (10), Taphozous nudiventris Cretzschmar, 1830 (3), Myotis blythii (Tomes, 1857) (14), M. emarginatus (Geoffroy, 1806) (2), M. formosus (Hodgson, 1835) (1), M. davidii (Peters, 1869) (5), M. longipes (Dobson, 1873) (4), Submyotodon caliginosus (Tomes, 1859) (1), Vespertilio murinus Linnaeus, 1758 (1), Eptesicus serotinus (Schreber, 1774) (2), E. pachyomus (Tomes, 1857) (2), E. ognevi Bobrinskoj, 1918 (1?), E. gobiensis Bobrinskoj, 1926 (1), Rhyneptesicus nasutus (Dobson, 1877) (5), Hypsugo savii (Bonaparte, 1837) (4), Pipistrellus pipistrellus (Schreber, 1774) (35), P. kuhlii (Kuhl, 1817) (9), P. coromandra (Gray, 1838) (6), P. tenuis (Temminck, 1840) (3), Nyctalus noctula (Schreber, 1774) (1), N. montanus (Barrett-Hamilton, 1906) (2), N. leisleri (Kuhl, 1817) (4), Otonycteris leucophaea (Severcov, 1873) (6), Barbastella darjelingensis (Hodgson, 1855) (3), Plecotus strelkovi Spitzenberger, 2006 (8), Scotophilus heathii (Horsfield, 1831) (3), Miniopterus fuliginosus (Hodgson, 1835) (1), M. pallidus Thomas, 1907 (4), Tadarida teniotis (Rafinesque, 1814) (1), and Nyctinomus aegyptiacus Geoffroy, 1818 (1). While Nyctalus noctula is here reported from the country for the first time, Rhinolophus mehelyi Matschie, 1901, Myotis bucharensis Kuzâkin, 1950, and Pipistrellus javanicus (Gray, 1838), reported erroneously to occur in Afghanistan by some previous authors, have been deleted from the list of the Afghanistani bat fauna. Key words. Distribution, taxonomy, Rhinopomatidae, Megadermatidae, Rhinolophidae, Hipposideridae, Emballonuridae, Vespertilionidae, Miniopteridae, Molossidae, Afghanistan, Middle East, Turkestan, Indian subcontinent, Palaearctic, Oriental region. INTRODUCTION The territory of the State of Afghanistan (647,000–653,000 km2, according to various sources) lies at the easternmost margin of the Middle East and westernmost margin of the broader Indian region (Fig. 1). It is situated on the crossroads of environmental as well as cultural influences between the Mediterranean, Arabia, Turkestan, Tibet, and India. The Hindu Kush range, with the highest * Dedicated to late Professor Dalibor Povolný (1924–2004), Brno, a reputable dipteran entomologist and a leader of the Czechoslovak Biological Expeditions to Afghanistan in 1965–1967. 267 Afghanistani peak of Nowshaq (7,492 m a. s. l.) forms the geographical axis of the country running from the south-west to the north-east; it also creates a border between two principal zoogeographi- cal regions in south-western Asia, the Palaearctic and Orient. From the biogeographical point of view, the area of Afghanistan represents an extremely interesting and also important region, where several faunal influences meet and coexist in a very variable selection of sub-tropical habitats. The bat fauna of this country has never been thoroughly reviewed and its knowledge currently rema- ins far behind that in the surrounding countries (see Bates & Harrison 1997, Benda et al. 2012). So, although Afghanistan only neighbours the Middle East (but this is rather a political term and geographical definition) and with a limited biogeographical influence from the Mediterranean, we still find it useful to assess the bat fauna of this country in this review series. Since Afghanistan is currently inaccessible for field research and most probably will remain inaccessible for a long time, we tried to review bat fauna of this country based on a literature survey and mainly on a thorough examination of as many specimens available in museum collections as possible. Although the mammal fauna of Afghanistan started to be investigated in a similar period as in other South-Asian areas (see Hutton 1845, Blanford 1881, Scully 1887a, Thomas 1889, etc.), the bats of this country remained almost unknown for a long time. With the exception of very few old individual and perhaps accidental records (Hutton 1845, Dobson 1878, Anderson 1881, Scully 1881b, Ogneff & Heptner 1928, Dupree 1958, Gaisler 1971), the bat fauna of Afghanistan started to be studied in the second half of the twentieth century. Moreover, majority of the bat field studies were made along with a research of other mammal/animal groups and only few campaigns were focused directly on bats. The first collection of bats was gathered by Johann Friederich Klapperich (1913–1987), an ento- mologist and preparator at the ZFMK museum, Bonn. He spent more than a year in Afghanistan in 1952–1953 and carried out three long-time trips through the north-eastern part of the country; the description of the trips as well as the gazetteer was published by Klapperich (1954). His collection, comprising 23 bat specimens of five species, was evaluated and published by Zimmermann (1956); however, this collection was spread over several museums in Europe and the US (see Neuhauser 1969), although most of its content remains in the SMF at Frankfurt am Main. Another collection of bats from Afghanistan was gathered by Knut Lindberg (1892–1962), a Swedish physician who worked between 1927–1947 at the Barsi Light Railways in Kurduvadi near Bombay, India. After his retirement in 1947, Lindberg made three field trips to Afghanistan (1947, 1957–1960, 1962) and during the latter two he collected cave fauna including bats (Lin- droth 1963, Löwegren 1964). Most of Lindberg’s bat records made until 1959 (99 specimens of 12 species) were published by Aellen (1959a), some others by Lindberg (1961, 1962), several specimens remained unpublished until now; the respective specimens are housed in the MHNG and MZLU collections. Based mainly on the Lindberg’s collection, Aellen (1959a) first offered a complete review of bat fauna of Afghanistan then composed of 15 species (Table 1). In the 1960s, several groups and individuals studied bats in Afghanistan; concerning the en- richment of knowledge of the Afghanistani bat fauna, these years were the most fruitful at all. Jochen Niethammer (1935–1998) from ZFMK, Bonn, visited Afghanistan in the periods 1962–1966 and 1972–1975; he collected an extensive series of small terrestrial mammals (see e.g. Nietham- mer 1965, 1967, 1969a, b, 1970, 1973, 1975, 1982, 1983, Nauman & Niethammer 1973, 1974, etc.) and along with them also some bats. Including the cave deposit remains, this bat collection comprises some 360 specimens of 18 currently recognised species. Although Meyer-Oehme (1968) and Niethammer (1968, 1983) published some records from this collection, most of the specimens have remained unpublished. Together with J. Niethammer, another German zoologist worked in Afghanistan in the 1960s, Detlef Meyer-Oehme (*1929). As a high school teacher at Kabul, he studied bats extensivelly (sometimes in co-operation with Niethammer, see below) and 268 published their findings in two rather short publications (Meyer-Oehme 1965, 1968); however, these reports brought records of several species new for the fauna of Afghanistan. As stressed by Niethammer (1968), thanks to the records made by D. Meyer-Oehme, at the late 1960s the bat fauna of Afghanistan was known better than that of any of the neighbouring countries. The bat collection created by Meyer-Oehme, represented by more than 500 specimens, is currently housed in the SMF and although it has not been published as a whole, particular specimens were used in many comparative studies (see e.g. Kock et al. 1972, Felten et al. 1977, Kock 1980, 1999, Nader & Kock 1980, 1990, Van Cakenberge & De Vree 1994, Benda et al. 2006, 2012, etc.). A group of zoologists from Czech scientific institutes visited Afghanistan at least four times in 1965–1967 to collect various animals, but mostly mammals and their parasites (see Daniel 1966, 1969, and Gaisler et al. 1967 for basic descriptions of the trips). The group included Milan Daniel (*1931), Jiří Gaisler (1934–2014), Dalibor Povolný (1924–2004), Pavel Rödl (*1941), Zdeněk Šebek (*1925), and František Tenora (1930–2011). Their effort resulted in collection of ca. 750 bat specimens belonging to 18 species, originating mainly from the eastern regions of Afghanistan and now housed mostly in the IVB and NMP collections. The