Rousettus Aegyptiacus (Pteropodidae) in the Palaearctic: List of Records and Revision of the Distribution Range*
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Vespertilio 15: 3–36, 2011 ISSN 1213-6123 Rousettus aegyptiacus (Pteropodidae) in the Palaearctic: list of records and revision of the distribution range* Petr BENDA1,2, Mounir ABI-SAID3, Tomáš Bartonička4, Raşit BILGIN5, Kaveh FAIZOLAHI6, Radek K. Lučan2, Haris NICOLAOU7, Antonín REITER8, Wael M. SHOHDI9, Marcel UHRIN10 & Ivan Horáček2 1 Department of Zoology, National Museum (Natural History), Václavské nám. 68, CZ–115 79 Praha 1, Czech Republic; [email protected] 2 Department of Zoology, Faculty of Science, Charles University, Viničná 7, CZ–128 44 Praha 2, Czech Republic 3 Department of Biology, American University of Beirut, Riadh El Solh 1107, 2020 Beirut, Lebanon 4 Department of Botany and Zoology, Masaryk University, Kotlářská 2, CZ–611 37 Brno, Czech Republic 5 Institute of Environmental Sciences, Bogazici University, Bebek, TR–34342 Istanbul, Turkey 6 Mohitban Society, 2010 Mitra Alley, Shariati Street, Tehran, 1963816163, Iran 7 Forestry Department, Ministry of Agriculture, Natural Resources and Environment, Louki Akrita 24, Nicosia, Cyprus 8 South Moravian Museum in Znojmo, Přemyslovců 8, CZ–669 45 Znojmo, Czech Republic 9 Nature Conservation Egypt & Department of Zoology, Faculty of Science, Al-Azhar University, Cairo, Egypt 10 Institute of Biology and Ecology, P. J. Šafárik University, Moyzesova 11, SK–040 01 Košice, Slovakia Abstract. The populations of Rousettus aegyptiacus inhabiting the Palaearctic part of the species range represent the only offshoot of the family Pteropodidae beyond tropics. In this contribution, we revised distribution status of the species in different parts of its Palaearctic range, re-examined the literature data and supplemented them with an extensive set of original records obtained in the field during the last two decades. Based on that, we compiled a complete list of the records of R. aegyptiacus from the Palaearctic Region, covering the range delineated by 12–38° N and 26–68° E. Two distribution patterns were recognised: (1) a locally continuous range in the thermo-Mediterranean zone of S Turkey, W Syria, Lebanon, N Israel, and NW Jordan, (2a) a clearly discontinuous distribution in sub-tropical deserts of the Saharo-Sindian zone (S Levant, Egypt incl. Sinai, N Sudan, Arabian Peninsula, S Iran and S Paki- stan), characterised by more or less isolated patches of distribution in desert oases and smaller areas of relatively humid habitats, and (2b) a supposedly continuous linear distribution along the Nile in Egypt, including its extensions in the El Fayum Oasis and Nile Delta. These patterns differ from the distribution characteristics of R. aegyptiacus in the sub-Saharan African range, where this fruit bat is reported to inhabit countinuous belts of rather open habitats stretching over large parts of the continent. Detailed data from the Dakhla Oasis (less than 1,000 km2) in the Western Desert of Egypt illustrate flexibility in roosting and foraging requirements of the species and its considerable capacity for long-term survival in small (2,000–3,000 inds.) and rather isolated populations. Fruit bats, distribution, range, occurrence pattern, Mediterranean, Middle East, Egypt Introduction The Egyptian fruit bat, Rousettus aegyptiacus (Geoffroy, 1810), is the only member of the Pteropo- didae family whose distribution range includes a large part in the Palaearctic Region, and also the only one whose type locality lies in the Palaearctic (the Great Pyramid of Giza, Egypt; Anderson 1902). Considering the whole distribution range, R. aegyptiacus is mostly an Afro-tropical species, * dedicated to Prof. Dr. Vladimír Hanák, one of the founders of the modern research of bats in the Mediterranean, on occasion of his 80th birthday. 3 which reaches the southwestern part of the Palaearctic (Juste & Ibañez 1993, Bergmans 1994, Kwiecinski & Griffiths 1999). In sub-Saharan Africa, it occupies areas around the Gulf of Guinea from Senegal to W Angola (including some islands in the Gulf), and savanna regions of South and East Africa from the Cape to Eritrea. In the Palaearctic, R. aegyptiacus occurs in various parts of Egypt as well as in broad areas along the sea coasts of the Middle East, from SW Turkey and Cyprus along the Levantine and Arabian shores up to S Iran and S Pakistan (Bergmans 1994). The Palaearctic range is reported to be isolated from the sub-Saharan African range by desert areas of northeastern Africa; in the Middle East (S Turkey), this bat reaches northern margin of its distribution range (Harrison & Bates 1991, Bergmans 1994, Benda et al. 2006). The complete review of distribution records of R. aegyptiacus in the Palaearctic was first provided by Kock & Nader (1979), an updated version was presented by Bergmans (1994) along with a detailed description of the whole distribution range. The Palaearctic range of R. aegyptiacus covers the area of the E Mediterannean, Egypt and the Middle East, incl. S Pakistan (delineated by 12–38° N and 26–68° E), but it is obviously quite discontinuous (see also Fig. 1): resident occurrence is confirmed for (1) Cyprus, (2) central parts of the Levant (W Lebanon and N Israel), and (3) northern parts of Egypt (Nile Delta and the Cairo Region), while status of the species in S Turkey, S Egypt, S Arabia, S Iran and S Pakistan, from where isolated records are available, is unclear and in no way suggests continuous occupancy of the whole range. In short, although R. aegyptiacus represents a top curiosity in the bat fauna of the western Palaearctic, the actual extent of its range, its history and character of local occurrence has not been sufficiently comprehended until recently. In response to scarcity of this information we re- -examined the distributional status of the species in some of the regions least known from this point of view. During the last two decades a number of new records were obtained which refine the previous picture both in the geographical and chorological respects. Although some of these records have already been published (see the List of Records), a vast majority of them have not been reported until now and/or not surveyed in regard to the distributional status of the species under study. The density of records and current geographical coverage of the range open a possibility of a detailed quantitative analysis of the range structure and its contextual setting to address the im- plicite questions on the enigmatic phenomenon of the Palaearctic offshoot of the family range. The critical re-examination and a standardised account of all available records are an essential prerequi- site for such analyses and the first step to be undertaken. Just this is the aim of the present study. Material and Methods The distribution data were obtained by screening of available literature reporting local bat records in the respective areas and by direct field studies in particular countries of the Middle East incl. Egypt (see List of Records). Field data were collected with the help of common techniques of field bat study, i.e. direct observations of flying bats, mist-nettings and visits of potential or already known roosts. Besides the direct findings of fruit bat individuals, also their abandoned roosts with undoubted traces of former presence were recorded (see e.g. Hadjisterkotis 2006) as well as their feeding places, i.e. accumulated remnants of food (mainly fruit stones) or the typical splodges from faeces and urine in sheltered places (such as monuments or abandoned rooms, see Figs. 2–5 and/or Benda et al. 2007: 76, Figs. 2, 3). The complete list of distributional records of R. aegyptiacus in the Palaearctic, provided in the following chapter, is arranged according to particular countries and classified in three categories; (1) localities summarised by Bergmans (1994) presented as mere site names; (2) new data published after or overlooked by Bergmans (1994) presented as site names complemented with the quotation of published source of the data; and (3) original records obtained by the authors of this contribution (both new and published), complemented with province name and coordinates of the site, method of data collection and a recorded number of fruit bats as well as the name/s of the collector/s of the data. In this paper we neither report the numerous negative records nor discuss the contextual aspects of the Palaearctic range of the species. These topics will be analysed elsewhere. 4 The list of records is supposedly complete for the ten countries which were in focus of our field study. For Israel, where R. aegyptiacus has been repeatedly intensively investigated as a model species during the last decades (e.g. Korine & Arad 1993, 1999, Korine et al. 1994, 1999, 2004, Krasnov et al. 2007, Tsoar et al. 2011, etc.), the present list is to be looked upon as provisional and obviously incomplete. It summarises just the available literature data (plus some original observations by the authors), including the map records surveyed by Mendelssohn & Yom-Tov (1999). It provides reliable comparative information sufficient for purpose of the present paper (see Figs. 1 and 6). List of Records Bergmans (1994) reported 123 record sites of Rousettus aegyptiacus from its Palaearctic range (Fig. 1). The other literature data not included in Bergmans’ (1994) review, mostly those published during the last two decades, report the species from 91 localities. With further 134 new record sites, the total number of localities where R. aegyptiacus was recorded within the Palaearctic range rose to 348. A majority of them (57.3%) was evidenced during the last two decades (see also Table 1 and Figs. 6, 11, 17). The minimum number of records in the particular countries is as follows: Turkey – 30 sites, Cyprus – 51, Syria – 5, Lebanon – 39, Israel and the Palestinian Territories – 32, Jordan – 20, Egypt – 64 (without new records from the Dakhla Oasis, see below), North Sudan – 2, Saudi Arabia – 5, Yemen – 19, Oman – 47, United Arab Emirates – 1, Iran – 31, Pakistan – 4.