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Lynx (Praha), n. s., 37: 9–12 (2006). ISSN 0024–7774 Professor Vladimír Hanák, 75 Profesor Vladimír Hanák pětasedmdesátníkem This volume contains the papers devoted to Vladimír Hanák, professor emeritus in Zoology at Charles University of Prague, a leading personality in Czech vertebrate zoology and one of the founding fathers of the modern study of mammals in the former Czechoslovakia and in the Czech Republic, on occassion of his 75th birthdays. Of course, it should be stressed that the senescence aspects commonly associated with that age like would not touch Vladimír in any significant extent – even in these days he continualy works in field study of bats, finishes a series of voluminous papers (among other, two volumes of a complete list of bat records in the Czech Republic and detailed summaries of distributional status of all particular species), a short time ago he coauthored immense analysis of bat banding results, a monographic survey of Palaearctic bats, wrote dozens of popularising articles, a survey on history of zoology in Central Europe and, at the same time, acts as a chief expert in various institutions of Nature Conservancy and essentially contributes to supervising of various research projects including PhD and MSc theses. Indeed, it is hard to believe that a man with so excellent mind and so manifold spectrum of activities really achieved his 75. Yet, it is the reality. Vladimír Hanák was born on 31 March 1931 in Znojmo, a town to which he paid a considerable attention during last decades, among other with his essential role in designing the National Park Podyjí-Thayatal, where he organizes a board of experts and actively promotes a large scale multidisciplinary investigations of that region. Going back to the region of his youth and/or concentration on regional faunal summaries (like e.g. a survey of bats in UNESCO Biospheric Reserve Třeboňsko he has finished just recently) is in no way a sign of regression. Since beginning of his scientific career, Hanák emphasized that a reliable primary record is an ultimate prerequisite to any further study and just because he ever put special attention to that point his results and scientific achievement, which much exceeded the regional scale of course, remain largerly valid and invariant to extensive conceptual and methodical shifts which appeared since time of his beginnings. Vladimír Hanák undoubtedly ranks among the chief personalities who designed the framework on which the Mid-European mammalogy of 20th century has arisen. At the beginning of 1950s when he started with profound critical reexamination of the taxonomical and distributional status of mammals in Central Europe and the Balkans, the state of knowledge was incredibly poor and to a considerable degree patterned by numerous misinterpretations and speculative predictions not rooted in a real evidence. Hanák started with a critical revision of that state and despite he continously worked in more groups of mammals and perforemed analyzes of mammalian communities and inter-regional comparisons of mammal fauna in a general extent, he focused, already in 1950s, together with Jiří Gaisler, his attention particularly to the group then the least known – bats. He started with large scale field studies of that group including extensive bat banding program and multidisciplinary gathering of various data on biology of particular species that was performed in cooperation with his students and colleagues of his age such as Jiří Gaisler, Karel Hůrka, Ladislav Janský, Milan Klíma, Jaroslav Figala, Leo Sigmund and others. Soon Hanák designed a large scale project of detailed analyses of taxonomic and distributional status of bat species covering not only the region of central Europe but the whole Palaearctic region. He succeeded to investigate the bat collections in many prominent European institutions and came in close contacts with almost all European specialists in that group and his approach soon became a model of a well designed research effort widely respected in an international scale. Among the achievements which particularly contributed to Hanák’s international reputation were first of all his pioneer role in discovery of sibling species Plecotus austriacus and Myotis brandtii, detailed taxonomical analyses of Myotis mystacinus group, studies on taxonomy and distribution of several least known taxa of the Palaearctic fauna such as Rhinolophus bocharicus, Eptesicus sodalis, Myotis longipes, 1970s: Vladimír Hanák among his colleagues and students during a field excursion to southern Moravia, the county of his childhood. 10 Hypsugo savii (and several other mostly in coauthorship with Jiří G aisler or Ivan Horáček) or the papers providing first comprehensive data on bats of Bulgaria, Libya and other regions. Unfortunately, only a smaller part of his achievements was properly published and the rest remained in state of manuscripts (such as his voluminous 1960 PhD thesis which summarized 2006: Vladimír Hanák with his students (and the editors of this volume) during taxonomical and field studies in southern Bohemia. distributional status of all Mid-European bat species throughout whole their ranges or a prepared monograph on the genus Myotis). Nevertheless, it was not only his research effort and publications for which Hanák became a leading personality of the post-war generation of Czech zoologists and a man who actually imprinted the development of the discipline in this country for next decades. His university teaching and the design of the vertebrate zoology program at the Charles University to which he essentially contributed present undoubtedly particularly significant part of Hanák’s achievements. Almost all Czech zoologists of today generations are direct or indirect students of Vladimír Hanák and perhaps all, similarly as many foreign zoologists who met Hanák personally were deeply impressed of his charming personality. It is not easy to identify which is source of the charm similarly as it is not easy to specify what all atributed the fact which all Hanák’s students like to stress – namely, that it was just the Hanák’s role as a university teacher what became an essential imprinting factor that stimulated their further career in zoology, their view of the specific qualities of that discipline and the meaning of the scientific study in that branch. The Hanák’s lectures, the practical courses or field excursions were, of course, traditionally performed as a top of professional standard and despite of Hanák’s sense of humor and tolerance with quite strong requirements on students’ comprehension. Yet, neither that was perhaps the most essential. What may have played a more significant role was probably the context in which Hanák the specificities of the branch and the particular scientific problems he uses to expose. Quite characteristic is his vivid, unostentatious but highly qualified interest in a broad spectrum of phenomena and processes which make together the phenomen of Nature, his deep respect to their aesthetic dimensions as well as to historical and cultural heritages which form our comprehension to them. Hanák perceives the Nature and animals like instant source area 11 of aesthetic and intelectual pleasures and the science of life grows thus in his representation in an extremelly attractive domain which opens a real chance to integrate an intelectual creativity with a meaningfull way of life rich in aesthetic and cultural issues. Vladimír has been quite consequent in respecting such a view in his own life and particularly with that, with his way to treat the pleasures and troubles of life and with his richly human interest on life if his students he has instantly demonstrated actual relevance and qualities of that view and of the way of life indexed by it. Moreover, Hanák never applied techniques that nowdays are nearly obligatory prerequisite to a scientific career: conceptual opportunism, affiliation of own view of a topic to the mainstream interpretations, overexposing or overestimation of own achievements. In consequence, his publications are not excessive either in number or in impact factors of the periodical in which they appeared but without exception they bring the concise information which reliability stays without any doubt. The ultimate index of Hanák’s scientific efforts was the actual meaning of the facts and their relevance within a broad contextual background in which his studies did exposed. The respect to his own sense of meaning of particular facts and the deep meaning of the world was valued for him much more than aspects of his academic career, secular honours or public fame. His straightforward way of presenting his opinions and deeply rooted sense of justice brought him many troubles, particularly in the period after occupation of Czechoslovakia by foreign armies in 1968. His career as well as contacts with the colleagues abroad were stopped for long years. The situation dramatically changed after the “velvet revolution” in 1989 when Hanák was immediately appointed associate professor and elected to various academic positions in which he considerably help in transformation of the university study of biology into a vivid modern institution. We do not wish, however, to repeat here the biographic data of Vladimír Hanák, neither enumerate his particular research achievements nor will provide list of his publications, by the way extensivelly enlarged during the recent decades. The reader can find these information in the volume Prague Studies in Mammalogy edited on occassion of Hanák’s 60th birthdays. Here we would like only to express, on behalf of numerous students and friends, warm thanks to Vladimír for all the gifts which he provided us and to wish him to the decades to come a good health, optimistic mind and lot of pleasure which he uses so generously to diseminate around throughout his life. The Editors 12 Lynx (Praha), n. s., 37: 13–17 (2006). ISSN 0024–7774 Holocene remains of the Pine marten (Martes martes) from the Tatra Mts. (Poland) – skull morphology and population structure (Carnivora: Mustelidae) Zmienność morfologiczna czaszek holoceńskiej populacji kuny leśnej (Martes martes) z jaskiń tatrzańskich i populacji współczesnej (Carnivora: Mustelidae) Justyna Bachanek & Bronisław W. Wołoszyn Polish Academy of Sciences, Institute of Systematics and Evolution of Animals, ul. Sławkowska 17, PL–31-016 Kraków, Poland received on 6 June 2006 Abstract. A morphological analysis of 36 skulls of the pine marten Martes martes (Linnaeus, 1758) col lected in caves in the Tatra Mts. (southern Poland) was carried out. Special attention was paid to age and sex structure of this Holocene population. The results of skull measurements of the subfossil population were compared with those of recent pine marten populations from Poland. INTRODUCTION The paper is aimed at the analysis of skull morphology of subfossil remains of the pine marten Martes martes (Linnaeus, 1758) from Poland regarding age and sex structure of the population. So far, no studies have been focused particularly on this aspect. Most publications are devoted to morphological comparison of skulls between Martes martes and Martes foina (Erxleben, 1777). The differences between the two species were presented by Anderson (1970), Altuna (1973), Steiner & Steiner (1986) using non-metrical parameters, and Gerasimov (1985) using a multivariate analysis of variance. In Europe, the skull morphology of martens was studied in various aspects and according to various criteria: age classes – Röttcher (1965), Habermehl & Röttcher (1967), sex dimorphism Bree et al. (1970), Rossolimo & Pavlinov (1974), as well as variability and asymmetry of dentition of pine and beech martens in Poland (Wolsan 1985, 1986, 1989) and in Europe (Reig 1989). The above mentioned publications were aimed only at comparisons of recent populations of the two marten species, whereas there are no articles comparing recent and subfossil populations. MATERIAL AND METHODS Several hundreds of caves are known in the Tatra Mts., mostly in their western part. These caves are characterized by specific ecoclimate and fauna. The material of 36 pine marten skulls was collected in the following Tatra caves: Czarna (7 specimens), Zimna (4), Mała Świstówka (1), Miętusia (1), Cho chołowska (2), Bielska (1), Ptasia Studnia (6), Niedźwiedzia (1), Za siedmioma Progami (12), Wysoka (1). The studied material was collected between 1961 and 1981, mostly by K. Kowalski, A. Woźnica, 13 M. Zagórny, M. Gębołys, B. W. Wołoszyn and has been deposited in the Institute of Systematics and Evolution of Animals, PAS, in Kraków. The subfossil skulls of the pine marten were found on the surface of cave floor and most probably are of the Holocene age. The following measurements were taken: condylobasal length (CBL), alveoli condylobasal length (CBLa); zygomatic width (ZYW); postorbital width (PorW); mastoid width (MstW). Males and females were distinguished on the basis of differences in alveoli condylobasal length (CBLa) and zygomatic width (ZYW). We used the simplified age classification presented by Buchalczyk & Ruprecht (1989), based on the following characters: (1) state of preservation of sutura internasalis, s. nasomaxillaris, s. nasofrontalis, and s. maxillofrontalis; (2) degree of tooth wear; and (3) crista sagittalis development. Three age classes were distinguished: (I) young individuals, contours of s. nasomaxillaris, s. nasofronta lis, and s. maxillofrontalis visible; beginning of crista sagittalis formation; rough surface of the toothrow; presumed age: 5–8 months; (II) mature specimens, crista sagittalis situated below the line, rough surface of the toothrow; presumed age: between 9 months and 2 years; (III) old individuals, long crista sagittalis, smooth surface of the toothrow, some teeth may be missing; age: 2 years or more. The data on skull morphology of the studied Holocene population were compared with the results of measurements of the recent marten population taken from the publication by Reig & Ruprecht (1989). These authors studied Martes martes from four regions of Poland: Masurian and Pomeranian Lakelands, Wielkopolska-Kujawy Lowland, Białowieża Forest, and Silesia. Since the subfossil material was damaged, it was impossible to take some of the measurements usually taken on recent material. The CBL length of the skull of recent specimens is measured from the anterior margin of incissivi to the posterior margin of condylus occipitalis. However, incisors are missing in most subfossil skulls, therefore the CBLa was taken. Results of both measurements taken on the same recent Table 1. Range and mean values of skull measurements of subfossil and recent martens (males and females separately). Explanations: CBLa – alveolar condylobasal length; CBL – condylobasal length; ZYW – zy gomatic width; PorW – postorbital width Tab. 1. Zróżnicowanie holoceńskiej populacji kun ze względu na różnorodność badanych parametrów. measurement mean±SD Holocene min–max females CBLa 74.97±1.83 72.15–78.33 CBL ZYW 42.68±2.75 36.47–48.69 PorW 18.25±1.38 16.35–21.50 mean±SD Recent min–max 78.5±4.46 75.0±4.27 45.3±2.40 17.3±6.30 72.15–88.15 75.50–81.60 43.00–48.00 15.30–20.00 males CBLa 83.18±2.22 79.80–88.15 CBL 85.6±1.70 ZYW 47.29±2.62 43.23–50.70 51.1± 3.20 PorW 19.24±0.83 17.77–21.01 19.1±6.50 82.0–87.90 47.6–55.20 15.8–21.30 all individuals CBLa 78.50±4.46 72.15–88.15 CBL 82.2±1.43 ZYw 44.20± 3.46 36.50–50.70 48.2±1.35 PorW 18.50±1.55 15.25–22.55 18.2±1.16 78.75–84.75 45.30–51.60 15.50–20.65 14 specimens were compared. An average difference between the two measurements was 0.6 mm, i.e. 0.01%, being within statistical error limits. An analysis regarding sex and age structure of the population was carried out. Statistical significance level of all differences was below 0.005. Values were transformed before the statistical analysis. All P-values reported are two tailed, the T-test of the differences between two means was performed. RESULTS Sexual dimorphism and population structure Both the Holocene and recent marten populations exhibit a distinctive sex dimorphism: males are larger than females. The distinguishing characters are: condylobasal length (CBL) or alveoli condylobasal length (CBLa), and zygomatic width (ZYW). In the subfossil population, differences in CBL and ZYW between the two sexes are statistically significant (p<0.001). Based on these characters, we classified 15 specimens as females, and 11 as males (Fig. 1), while skull dimensions of the remaining 10 specimens did not allow their classification. Comparison of subfossil and recent material Within the sex groups, the Holocene marten skulls are shorter than the recent ones by 3.85 mm on average (statistically significant difference, p<0.005; Table 1). Fig. 1. Skull dimensions of Holocene martens – sex groups. Rys. 1. Zróżnicowanie holoceńskiej populacji kun ze względu na płeć. 15 Fig. 2. Sex and age structure of the studied Holocene marten population. Rys. 2. Struktura populacji holoceńskiej pod względem płci i wieku. Age of individuals The Holocene marten sample comprises 36 specimens. Most of them are adults from the II age class (n=24). Older specimens (of III age class) are less numerous (n=10), and only two individuals were classified as young. DISCUSSION Trogloxenes – animals inhabiting caves incidentally or using them as seasonal shelters – are the most numerous group in the fauna of caves. The pine marten Martes martes is one of such species: it occurs in caves sporadically but gets through cave corridors far from the entrance. The authors analysed the structure of a subfossil marten population on the basis of skull morphology. Our results show that the Holocene martens had on average by 3.85 mm smaller skulls than the recent ones. This difference may be related to thermoregulation requirements in different climatic conditions: according to Bergmann’s rule animals living in warmer climate tend to have a relatively higher body surface to body mass ratio (e.g. smaller body size) than animals living in colder climate where reduction of relative body surface (e.g. larger body size) is favored. It is likely that the remains of martens from the Tatra caves originate from the Holocene optimal climatic period, Atlantic or Subatlantic, when smaller individuals could easily radiate excess of heat. The results of radiocarbon dating (C14) of the subfossil material could bring a definitive answer to this question. The age structure analysis was carried out using simplified age categories proposed by Bu chalczyk & Ruprecht (1989). These authors assessed the age of 596 marten specimens, grou ping them into five age groups. For the small Holocene sample (n=36) we reduced age groups to three. Mature specimens prevailed in our material, while old animals (of 2 or more years of age) were by half less numerous. Young individuals (of less than 8 months) were rare – only two specimens out of the 36. The low proportion of young animals indicates that caves were used by martens as temporary shelters or hunting grounds, not for raising offspring (Fig. 2). 16 Podsumowanie Zbadano holoceńską populację kun leśnych (Martes martes) z jaskiń tatrzańskich. Wybrane parametry pomiarów czaszki porównano z pomiarami wykonanymi na materiale współczesnym. Mniejsze rozmiary populacji holoceńskiej (Tab.1) wskazują zgodnie z Regułą Bergmanna, że badana populacja zasiedlała ten teren w cieplejszym klimacie, prawdopodobnie w optimum klimatycznym Holocenu. REFERENCES Altuna J., 1973: Distincion craneal entre la Marta (Martes martes) y la Foina (Martes foina) (Mammalia). Munibe, 25: 33–38. Anderson E., 1970: Quaternary evolution of the genus Martes (Carnivora, Mustelidae). Acta Zool. Fenn., 130: 1–132. Buchalczyk T. & Ruprecht A. L., 1977: Skull variability of Mustela putorius Linnaeus, 1758. Acta The riol., 22: 87–120. van Bree P. J. H., van Mensh P. J. A. & van Utrecht W. L., 1970: Sur le dimorphisme sexuel des canines chez la Foine, Martes foina (Erxleben, 1777). Mammalia, 34: 676–682. Gerasimov S., 1985: Species and sex determination of Martes martes and Martes foina by use of systems of craniometrical indices developed by stepwise discriminant analysis. Mammalia, 49: 235–248. Habermehl K. H. & Röttcher D., 1967: Die Möglichkeiten der Altersbestimmung beim Marder und Iltis. Ztschr. Jagdwiss., 13: 89–102. Reig S., 1989: Morphological variability of Martes martes and Martes foina in Europe. Ph.D. Thesis, Jagiellonian University, Cracow, 145 pp. Reig S. & Ruprecht A. L., 1989: Skull variability of Martes martes and Martes foina from Poland. Acta Theriol., 34: 595–624. Rossolimo O. & Pavlinov I., 1974: Sexual dimorphism in the development, size and proportions of the skull in the pine marten (Martes martes Linn.: Mammalia, Mustelidae). Pp.: 180–191. In: King C. M. (ed.): Biology of the Mustelids: some Soviet Research. Volume 1. British Library, Boston Spa, Yorkshire, 266 pp. Röttcher D., 1965: Beträge zur Altersbestimmung bei Nerz, Steinmarder und Iltis. Inaugural-Dissertati on zur Erlangung des Doktorgrades. Veterinärmedizinischen Fakultät der Justus Liebieg-Universität, Giessen, 83 pp. Steiner H. M. & Steiner F. M., 1986: Die nicht-metrische Unterscheidung von Schädeln mitteleuropäischer Baum- und Steinmarder (Martes martes und Martes foina, Mammalia). Ann. Naturhist. Mus. Wien, 88/89 B: 267–280. Wolsan M., Ruprecht A. L. & Buchalczyk T., 1985: Variation and asymmetry in the dentition of the pine and stone martens (Martes martes and M. foina) from Poland. Acta Theriol., 30: 79–114. Wolsan M., 1986: Morphological variation of the first upper molar in the genus Martes (Carnivora, Mus telidae). Mem. Mus. Natn. Hist. Natur. Paris (S. C), 53: 241–254. Wolsan M., 1989: Dental polymorphism in the genus Martes (Carnivora: Mustelidae) and its evolutionary Significance. Acta Theriol., 34: 545–593. Wołoszyn W., 1996: Fauna jaskiń tatrzańskich [Fauna of the Tatra caves]. Pp.: 525–533. In: Mirek Z., Głowaciński Z., Klimek K. & Pięko-Mirkowa H. (eds.): Przyroda Tatrzańskiego Parku Narodowego [Nature of the Tatra National Park]. Tatrzański Park Narodowy, Kraków-Zakopane. 17 Lynx (Praha), n. s., 37: 19–21 (2006). ISSN 0024–7774 Savi’s pipistrelle (Hypsugo savii): bat species breeding in the Czech Republic (Chiroptera: Vespertilionidae) Netopýr Saviův (Hypsugo savii): další v Česku se rozmnožující druh netopýra (Chiroptera: Vespertilionidae) Tomáš Bartonička1 & Peter Kaňuch2,3 1 Institute of Botany and Zoology, Faculty of Science, Masaryk University, Kotlářská 2, CZ–611 37 Brno, Czech Republic; bartonic@sci.muni.cz 2 Institute of Forest Ecology, Slovak Academy of Sciences, Štúrova 2, SK–960 53 Zvolen, Slovakia; kanuch@netopiere.sk 3 Institute of Vertebrate Biology, Academy of Sciences CR, CZ–675 02 Studenec 122, Czech Republic received on 25 November 2006 Abstract. On 17 August 2006, an adult female Savi’s pipistrelle (Hypsugo savii) was mist-netted above a park fountain in the Brno city. The female showed signs of postlactation like the presence of bare patches around its bulgy nipples. It was measured, banded and next night released in the same park. Previous records of the species concerned two males and a subadult female. This is the fourth record of H. savii but the first reliable one indicating reproduction of this species in the territory of the Czech Republic. Savi’s pipistrelle, Hypsugo savii (Bonaparte, 1837) has a West Palaearctic distribution range and in Europe it is widespread mostly in the Mediterranean region (Horáček & Benda 2004). Previous records of the species in the Czech Republic up to 2003 were made outside the bat’s breeding period, thus irrelevant to the problem of reproduction of the species. First male was found 20 km southwards of Brno in the village of Žabčice and a subadult female and an adult male just within the territory of Brno (Gaisler 2001, Gaisler & Vlašín 2003). At the evening of 17 August 2006, two mist-nets (7 and 12 m long) were set over a historical stone fountain (five meters in diameter) in the Lužánky park in the city of Brno. The park, ca. 400×400 meters, has a perimeter of ca. 1.8 km. Full-grown and hollow trees in the park as well as old historical buildings surrounding the park provide high number of potential roosts for bats. An artificial pond was built recently in the western part of the park and is also important to the bats. However, the shallow water body in the fountain was the only water habitat of the park and its surroundings until the vegetation season of 2005. Mist-netting lasted between 8:00pm and 11:30pm. We caught 13 individuals of five bat species – mostly Pipistrellus pipistrellus (Schreber, 1774) s. str. and per one individual of Eptesicus serotinus (Schreber, 1774), Plecotus auritus (Linnaeus, 1758) and Myotis daubentonii (Kuhl, 1817). The female of H. savii was caught at 9:10 pm. She had signs of postlactation such as the presence of bare patches around its bulgy nipples and swollen external genitals. Wings, ears and face were dark in contrast to the brown dorsal fur and grey belly. There were two cusps on I2, between I3 and C1 was a gap, C1 and P4 were in contact. The peak frequency of the search sequence of echolocation calls was 19 between 33.0–35.5 kHz. External measurements: body 47.8 mm, tail 32.8 mm (free tip of the tail was 3 mm long), forearm 35.1 mm, hind foot 6.7 mm, ear 10.2, tragus 4.3 mm, the fifth finger 42.6 mm, the third finger 57 mm, body mass 6.8 g. A sample of wing membrane was preserved in 96% ethanol for later DNA analysis. A photograph was made using a digital camera (Fig. 1). The bat was marked with the band N. Museum Praha No X16019, kept in captivity and next night released within the same park. It flew away without any obvious difficulties. In addition to the two published records (Gaisler 2001, Gaisler & Vlašín 2003), an adult male was found in March 2006 within the city of Brno (leg. P. Koutný). It was seriously injured, with its left forearm broken. The bat was deposited in the collections of the Institute of Botany and Zoology, Faculty of Science, Masaryk University. Fast train of records of H. savii during the last five years shows spreading of the species towards the north. During the last two years several individuals of H. savii, in winter as well as in summer, were found also in Slovakia (Le hotská & Lehotský 2006, Lehotská 2006, Ceľuch & Ševčík 2006). It can be assumed that the species is an active immigrant to the Czech Republic and Slovakia. The records from Slovakia are located more to the south that these from Brno. At present, Brno and its environs seems to be the northernmost point of the species’ regular occurrence (Gaisler & Vlašín 2003) and the potential reproduction. Two individuals found in northern Germany and another from England were probably transported passively (Ohlendorf et al. 2000, Gaisler 2001). Low spectral parameters and, in particular, low peak frequency of pipistrelle like echolocation calls recorded in Nitra and Brno (lower than in signals of Pipistrellus nathusii) suggest more often occurrence of H. savii in urban areas than expected (T. Bartonička and M. Ceľuch, unpublished). Bats can Fig. 1. Postlactating female of Savi’s pipistrelle (Hypsugo savii) mist-netted in the Brno city. Obr. 1. Postlaktační samice netopýra Saviova (Hypsugo savii) odchycená v Brně. 20 occupy the prefab houses similarly as Nyctalus noctula (Schreber, 1774) or Vespertilio murinus (Linnaeus, 1758). Possible existence of a maternity colony will be investigated in near future. SOUHRN V centru Brna byla dne 17. 8. 2006 odchycena dospělá samice netopýra Saviova (Hypsugo savii). Samice vykazovala jasné postlaktační znaky jako bezchlupé okolí zvětšených prsních bradavek a zduřelé vnější genitálie. Celkově se jedná již o čtvrtý záznam druhu na území České republiky avšak jde o první doklad jeho reprodukce na tomto území. Acknowledgements We are very grateful to J. Gaisler for valuable comments to the manuscript. Female was captured during the activities involving the study of pipistrelle’s populations in central Europe. The study was supported by the grant of Grant Agency of the Czech Republic No 206/06/0954 and the grant of Ministry of Education, Youth and Sports of the Czech Republic No MSM0021622416. References Gaisler J., 2001: A mammal species new to the Czech Republic – Savi’s pipistrelle Hypsugo savii. Folia Zool., 50: 231–233. Gaisler J. & Vlašín M., 2003: Second record of the Savi’s pipistrelle (Hypsugo savii) in the Czech Re public. Vespertilio, 7: 181–182. Horáček & Benda P., 2004. Hypsugo savii (Bonaparte, 1837) – Alpenfledermaus. Pp.: 911–941. In: Krapp F. (ed.): Handbuch der Säugetiere Europas. Band 4: Fledertiere. Teil II: Chiroptera II. Vespertilionidae 2, Molossidae, Nycteridae. AULA-Verlag, Wiebelsheim, x+605–1186 pp. Lehotská B., 2006: Second record of the Savi’s pipistrelle (Hypsugo savii) in Slovakia. Vespertilio, 9–10: 225–226. Lehotská B. & Lehotský R., 2006: First record of Hypsugo savii (Chiroptera) in Slovakia. Biológia, Bratislava, 61: 192. Ohlendorf B., Vierhaus H., Heddergott M. & Bodino F., 2000: Korrektur: Fund einer Nordfledermaus (Eptesicus nilssoni) in Hamburg (ds. Z. Bd. 5, p. 220) betraf eine Alpenfledermaus (Hypsugo savii). Nyctalus (N. F.), 7: 454. Ceľuch M. & Ševčík M., 2006: Netopiere (Chiroptera) v meste Nitra (predbežná správa) [Bats (Chiroptera) in the City of Nitra (Preliminary Report)]. Skupina pre ochranu netopierov, Nitra, 11 pp (in Slovak). 21 Lynx (Praha), n. s., 37: 23–44 (2006). ISSN 0024–7774 On the occurrence of Eptesicus bobrinskoi in the Middle East (Chiroptera: Vespertilionidae) K výskytu netopýra turanského (Eptesicus bobrinskoi) na Blízkém východě (Chiroptera: Vespertilionidae) Petr Benda1,2 & Antonín Reiter3 1 Department of Zoology, National Museum (Natural History), Václavské nám. 68, CZ–115 79 Praha 1, Czech Republic; petr.benda@nm.cz 2 Department of Zoology, Charles University, Viničná 7, CZ–128 44 Praha 1, Czech Republic 3 South Moravian Museum Znojmo, Přemyslovců 8, CZ–669 45 Znojmo, Czech Republic; reiter@znojmuz.cz received on 23 September 2006 Abstract. A profound morphologic comparison of smaller representatives of the genus Eptesicus inha biting the western Palaearctic (E. bobrinskoi, E. gobiensis, E. nasutus, and E. nilssonii) confirmed the occurrence of Eptesicus bobrinskoi in NW Iran. Although it was previously not fully accepted, a revision of the collection material as well as of specimens newly recorded in Iran brought evidence showing that the doubts were unjustified. INTRODUCTION The Bobrinskoy’s serotine, Eptesicus bobrinskoi Kusjakin, 1935, was described on the basis of three individuals collected by S. P. Naumov in May and June 1928 in a region close to the north-eastern shore of the Aral Sea, central Kazakhstan, which were primarily assigned by Bobrinskoi (1931) to the form Eptesicus caucasicus (Satunin, 1901) [= Hypsugo savii (Bo naparte, 1837)]. Kusjakin (1935) characterised E. bobrinskoi to be in its external appearance close to Eptesicus alashanicus Bobrinskoj, 1926 [= Hypsugo alashanicus] but with a larger skull, although both forms were of similar body size. He mentioned E. bobrinskoi to be typical with its flattened braincase, a relatively small second upper incisor, and a very pale coloration of the pelage and naked parts. The Tjulek well in the Aral part of the Karakum Desert [= Priaralskye Karakumy D.], 65 km E of the town of Aral’skoe more [= Aral’sk], Kazakhstan, was designated as the type locality of this species (Kusjakin 1935, Rossolimo & Pavlinov 1979). Ellerman & Morrison-Scott (1951) provisionally arranged E. bobrinskoi as a subspecies of E. nasutus (Dobson, 1877) but this view has not generally been followed (see also Harrison 1963 and Hanák & Gaisler 1971). E. bobrinskoi remains the only bat species described from Kazakhstan (Butovskij et al. 1985). The essential part of the distribution range of E. bobrinskoi covers the zone of colder deserts and semi-deserts of central Kazakhstan, between the Aral Sea in the west and the western Betpak-Dala Desert in the east (Strelkov 1980), i.e. between 45° 30’ – 49° N and 59° – 70° 15’ E 23 (Butovskij et al. 1985). About 90% of the records ascribed to this species come from this region (cf. Strelkov 1980, Strelkov & Šajmardanov 1983). A few of records of E. bobrinskoi were described from outside this main range. Kuzjakin (1950) reported a young male specimen from the vicinity of Faskal, North Ossetia, northern slope of the Greater Caucasus Mts He also mentioned a female specimen deposited in the alcohol collection of the Zoological Institute of the Academy of Science [St. Petersburg] labelled “Jakutsk, Middendorf” suggesting its origin in Yakutia [= Sakha, northern Siberia, Russia]. These records seem to be unusual from the biogeographical point of view but have been largely considered rather correctly identified (Vereščagin 1959, Strelkov 1963, 1981, Tavrovskij et al. 1971). However, Kuzjakin (1965) labelled both records with question marks (?) and noted the preservation state of these specimens as not quite perfect and mainly, the age of the individuals as juvenile. In this way, Kuzjakin (1965) indirectly doubted his previous identifications. Therefore, these records were considered clearly doubtful by Hanák & Horáček (1986) who regarded them to be of juvenile individuals of E. nilssonii (Keyserling et Blasius, 1839). This view was followed by Nowak (1994), Borisenko & Pavlinov (1995), Pavlinov & Rossolimo (1998), Horáček et al. (2000), and Simmons (2005), but not by Koopman (1993, 1994) or Duff & Lawson (2004). Anyway, E. bobrinskoi has not recently been considered a member of the Yakutian bat fauna (Revin et al. 2004). Strelkov (1980) described a finding of several individuals of E. bobrinskoi in the northern Caspian Region in western Kazakhstan (the Mus Tomb, 18–20 km W of Kosčagyl, ca. 10 km S of the lower Emba River). Although geographically isolated, this record has been accepted beyond any doubt by later authors (Butovskij et al. 1985, Strelkov 1986, Rybin et al. 1989, Ilyin 2000). Moreover, a new record of E. bobrinskoi has been recently reported from a region close to the latter one (Davygora et al. 1998, Ilyin 2000), from a site 20–25 km S of Krasnojar, Novoalekseevka Dist., Aktjubinskaja Region, NW Kazakhstan. This record represents the northwesternmost verified spot of occurrence of this species in the Palaearctic, and because the Emba River is considered by some authors to be the Euro-Asian border in the region between the Caspian Sea and the Ural Mts, it is now appropriate to place E. bobrinskoi on the list of European bat species (Benda & Hutterer 2005). The only record of E. bobrinskoi in the Middle East (and also outside the former USSR) was published by Harrison (1963) from north-western Iran. He reported a finding of seven dead individuals (four males, three females) in “the Sulphur caves at Guter-Su, north of Mt. Sabalan, […] Iranian Azerbaijan, 38° 10’ N, 47° 40’ E”. Harrison (1963) supported this species identifi cation by his observation of the typical morphologic characters according to Kuzjakin (1950), and also by a comparison with the collection material of E. nilssonii and E. nasutus which both markedly differed from the Guter-Su specimens in size and in all important cranial traits. The Harrison’s (1963) identification was confirmed by Hanák & Gaisler (1971) who used his data in their broad comparison of the genus Eptesicus. E. bobrinskoi was for a long time considered to be a part of the fauna of Iran (Lay 1967, Etemad 1969, Hanák & Gaisler 1971, Corbet 1978, DeBlase 1980, Allison & Gaisler 1982, Butovskij et al. 1985). However, Hanák & Horáček (1986) revised the series of skulls of E. bobrinskoi from the Guter-Su finding, which is deposited in the collection of the Natural History Museum, London (BMNH), under the Nos. 63.1186–1192. They concluded that: “(a) Alle unter suchten Stücke einschlißlich der größten (Nr. 63 1189) sind diesjährige Jungen mit auffallend niedrigeren Werten der Schädelbreite, mit einer unvollendeten Ossifikation der Terminalränder des Processus coronoideus und mit einer unvollendeten Eruption der Zähne, u. ä. (b) Die metri24 schen Werte und proportionen des Rostrums, welche eher der Art E. bobrinskoi entsprechen, sind aus den oben angeführten Gründen wenig nachweisbar; nach wichtigsten anderen Merkmalen (Anwesenheit der Foramina cavernosa, Fellfährung) entspricht das Material vielmehr der Art E. nilssoni. (c) Aus den angeführten Tatsachen ergibt sich, daß dieses Material offensichtlich juvenile Stücke der zentralasiatischen Population E. nilssoni darstelt. […] (d) Die angeführte Interpretation des Fundes aus Guter-Su steht im Einklang mit der bisherigen Ansicht über die zoogeographischen Bewertung von E. bobrinskoi.” According to this revision, E. bobrinskoi has no longer been regarded as a member of the bat fauna of the Middle East (although not absolutely, see e.g. Nowak 1994, Sharifi et al. 2000), and the Harrison’s (1963) record has been considered to be of E. nilssonii gobiensis Bobrinskoj, 1926 (Koopman 1993, 1994, Bori senko & Pavlinov 1995, Pavlinov & Rossolimo 1998, Horáček et al. 2000, Duff & Lawson 2004, Simmons 2005, cf. Rybin et al. 1989, Rydell 1993, Benda & Horáček 1998, Albayrak 2003). However, based on a profound morphologic analysis, Strelkov (1986) showed the latter form to represent a separate species, E. gobiensis. This result has been widely accepted (Pavlinov & Rossolimo 1987, Rydell 1993, Borisenko & Pavlinov 1995, Bates & Harrison 1997, Roberts 1997, Pavlinov & Rossolimo 1998, Horáček et al. 2000, Šajmardanov 2001, Simmons 2005, etc.). During a field trip to Iran in the late spring of 2006 we visited the abandoned sulphuric mines in the oriental thermal bath resort of Qutur Su (= Guter-Su by Harrison 1963). In these caverns we found several dead birds and three partly mummified cadavers of bats. We preliminarily identified these bats as small representatives of the genus Eptesicus Rafinesque, 1820, but differing from E. nilssonii or E. gobiensis by a shorter forearm in two individuals: LAt 34.4 and 36.7 mm (the third cadaver was without wings) in Qutur Su bats vs. 37–44 mm in E. nilssonii s. l. (Hanák & Horáček 1986). These bats thus well resembled E. bobrinskoi according to Harrison (1963). The aim of this paper is to compare our specimens from Qutur Su with those identified as E. bobrinskoi by Harrison (1963) as well as with representative samples of E. bobrinskoi, E. na sutus, E. nilssonii, and E. gobiensis, to identify and/or revise our and Harrison’s (1963) findings and thus, to contribute to the knowledge of smaller Eptesicus bats in the Middle East. THE RECORD The three bats were found in two small abandoned sulphuric mines above the village of Qutur Su ( [qwtwr sw]), ca. 20 km ESE of Meshgin Shahr ( [mšgin šhr]), Province of Ardabil, 38° 20’ N, 47° 51’ E; 2545 m a. s. l. (Fig. 1). Although slightly differing in the geographical coordinates given, this site is undoubtedly identical with the locality of Guter-Su reported by Harrison (1963). The site is situated on the northern slope of the main peak of the Sabalan Range ( [kwh sblan]) which rises to the altitude of 4811 m from the flat steppe plateau of about 1100 m a. s. l. The village and mines lie in the zone of alpine meadows at the altitude of about 2500 m (Fig. 2). In these small artificial rocky caverns, which act as a natural trap being filled by an unbreathable atmosphere, the bat cadavers were found under smaller stones covering the mine floor. MATERIAL AND METHODS For a morphologic comparison of the newly collected bats in Qutur Su, we used museum material of all smaller species of the genus Eptesicus occurring in the western Palaearctic; viz. E. bobrinskoi Kusjakin, 1935 (11 specimens) from central Kazakhstan (i.e. from its “topo-type” area), E. nasutus (Dobson, 1877) (18) from the Middle East, including all currently recognised subspecies [E. n. nasutus from Afghanistan 25 Figs. 1, 2. Qutur Su, NW Iran. 1 – Abandoned sulphuric mines, the only site of (repeated) finding of E. bobrinskoi in Iran (above). 2 – Alpine meadows at the altitude of ca. 2500 m surrounding the site of Fig. 1, situated on the northern slope of Mount Sabalan (4811 m a. s. l., in the background) (below). Obr. 1, 2. Qutur Su, SZ Iran. 1 – Opuštěné sirné dobývky, jediná lokalita (opakovaného) nálezu netopýra turanského (E. bobrinskoi) v Iranu (nahoře). 2 – Alpinské louky v nadmořské výšce zhruba 2500 m obklopující lokalitu na obr. 1, ležící na severním svahu hory Sabalan (4811 m n. m., v pozadí) (dole). 26 and from Persian Baluchestan, E. n. matschiei (Thomas, 1905) from SW Arabia, E. n. pellucens (Thomas, 1906) from Mesopotamia, and E. n. batinensis Harrison, 1968 from Oman], E. nilssonii (Keyserling et Blasius, 1839) (24) from Central Europe, E. gobiensis Bobrinskoj, 1926 (6) from northern Kirghizia, as well as the BMNH series of seven bats from Qutur Su identified by Harrison (1963) as E. bobrinskoi and by Hanák & Horáček (1986) as E. nilssonii. Since Hanák & Horáček (1986) found the Qutur Su series to be partially composed of immature individuals, we used also sets mixed from adult and subadult indi viduals for dimensional comparisons. For the data associated with the specimens see the Appendix. For comparative purposes we used mainly skulls, from external dimensions we took only the forearm length (LAt). The specimens were measured in a standard way using mechanical or optical callipers. We evaluated 38 dimensions in each skull (17 measurements in skull and maxillar tooth-rows, 7 measurements in the mandible and mandibular tooth-rows, see also Abbreviations) including 14 indices that described the skull shape. The baculum was extracted with the use of a 4% solution of NaOH and coloured by the Alizarin red. Statistical analyses were performed using the Statistica 6.0 software. ABBREVIATIONS Collections. BMNH = Natural History Museum, London, United Kingdom; CUP = Department of Zoology, Charles University, Praha, Czech Republic; JOC = private collection of Ján Obuch, Blatnica, Slovakia; NMP = National Museum (Natural History), Praha, Czech Republic; ZIN = Institute of Zoology of the Russian Academy of Science, Sankt Peterburg, Russia. Measurements. LAt = forearm length; LCr = greatest length of skull; LCb = condylobasal length of skull; LaZ = zygomatic width of skull; LaI = width of interorbital constriction; LaInf = rostral width between the foramina infraorbitalia; LaN = neurocranium width; LaM = skull width at the mastiodal processes; ANc = neurocranium height; ACr = skull height (incl. the tympanic bullae); CC = rostral width between the canines (incl.); P4P4 = rostral width between the upper premolars (incl.); M3M3 = rostral width between the third upper molars (incl.); IM3 = length of upper toothrow between the first incisor and the third molar, I1M3 (incl.); CM3 = length of upper toothrow between the canine and the third molar, CM3 (incl.); P4M3 = length of upper toothrow between the premolar and the third molar, P4M3 (incl); M1M3 = length of upper toothrow between the first and third molars, M1M3 (incl.); CP4 = length of upper toothrow between the canine and the premolar, CP4 (incl.); LMd = condylar length of mandible; ACo = height of the coronoid process; IM3 = length of lower toothrow between the first incisor and the third molar, I1M3 (incl.); CM3 = length of lower toothrow between the canine and the third molar, CM3 (incl.); P4M3 = length of lower toothrow between the second premolar and the third molar, P4M3 (incl.); M1M3 = length of lower toothrow between the first and the third molar, M1M3 (incl.); CP4 = length of lower toothrow between the canine and the second premolar, CP4 (incl.). Others. m = male; f = female; ind. = individual of undetermined sex; a = adult; s = immature; A = alcohol preparation; B = skin (balg); S = skull; Sk = skeleton. RESULTS AND DISCUSSION The two series of bats from Qutur Su, that published by Harrison (1963) and that newly collec ted, are very similar in dimensions (Table 1) and seem to comprise very close or even identical morphotypes. To mention the most important characters, these specimens have similar forearm lengths, lengths of skull and tooth-rows and mainly, similar heights of braincase (ANc, ACr). Therefore both sets of specimens from Qutur Su were compared as one group of samples. Unlike Hanák & Horáček (1986), we found a part of the Harrison’s (1963) samples to be adult individuals, which was also true for two of the newly collected bats (see Table 1). From the whole set of the Qutur Su bats, at least five specimen bear signs of adult age on their skulls; i.e. a fully ossified basilar suture in the skull base (the connection between basioccipital and ba sisphenoidal bones) and moreover, a noticeable teeth abrasion. Thus, skulls of these specimens 27 28 sex age LAt NMP, 5 June 2006 90890 f a 36.7 15.2414.88 9.56 4.14 4.86 7.95 8.11 4.74 5.88 4.32 6.04 5.21 3.38 1.9110.75 3.14 5.61 3.64 1.79 90891 f s 34.4 – – – 4.02 4.31 – – – – 3.88 5.63 5.07 3.37 1.8810.14 2.81 5.48 3.76 1.90 90892 m a – 14.5714.43 – 3.95 4.48 7.16 7.90 4.61 5.88 4.00 5.88 5.02 3.39 1.8710.29 2.86 5.42 3.78 1.91 7.91 4.34 5.48 3.90 5.78 5.28 3.57 1.9610.51 2.91 5.62 3.85 1.84 8.22 4.47 – 3.79 5.94 5.38 3.55 2.12 – 3.12 – – – 7.77 4.52 5.51 3.74 5.61 5.08 3.47 1.8210.29 3.00 5.67 3.68 2.07 8.05 4.28 5.81 – – 5.07 3.35 1.81 – – – – – LCr LCb LaZ LaILaInf LaN LaM ANc ACr CC M3M3 CM3 M1M3 CP4 LMd ACo CM3 M1M3 CP4 BMNH, 21 August 1961 63.1186 m a 35.0 14.6714.35 – 3.92 4.37 7.42 63.1187 m s 35.4 14.8214.28 – 4.11 – 7.82 63.1188 f s 35.1skull broken 63.1189 m a 35.1 15.1714.87 8.38 4.19 4.28 7.64 63.1190 f a 35.7 15.0714.53 – 4.08 4.48 7.87 63.1191 f s 33.9skull broken 63.1192 m s 34.7skull entire but crushed No. Table 1. Dimensions of E. bobrinskoi collected at Qutur Su, NW Iran. For dimension abbreviations see the text. Forearm lengths (LAt) of the BMNH specimens were taken from DeBlase (1980) Tab. 1. Rozměry jedinců E. bobrinskoi nalezených v Qutur Su, SZ Iran. Vysvětlivky zkratek rozměrů viz text. Délky předloktí (LAt) jedinců z londýnské sbírky (BMNH) byly převzaty z přehledu DeBlaseova (1980) (BMNH 63.1186, 1189, 1190, NMP 90890, 90892) were available for a relevant morphologic comparison. As pointed out in Introduction, the Qutur Su bats are typical with relatively short forearms (LAt 33.9–36.7 mm). In this character they are very close to the samples of E. bobrinskoi from Kazakhstan (LAt 34.0–36.7 mm) and overlap also with the samples of E. nasutus (LAt 35.5–39.9 mm). On the other hand, the Qutur Su bats clearly differ in their forearm lengths from the samples of E. nilssonii and E. gobiensis (Table 2). Based on the comparison of skull dimensions, the skulls of the Qutur Su bats are most similar to the samples of E. bobrinskoi; they broadly overlap in all measurements taken (Table 2), although the Kazakh samples are on average slightly larger. The Qutur Su bats also partly overlap in their skull lengths (LCr, LCb, CM3, P4M3, M1M3, LMd, CM3, P4M3, M1M3) and widths (LaZ, LaI, LaInf, LaN, LaM, M3M3) with E. nilssonii and slightly also with E. gobiensis, but clearly differ from both sample sets in skull heights (ANc, ACr). However, the Qutur Su bats clearly exceed in their skull lengts all samples of E. nasutus but overlap them in the skull heights. These relations are shown also in Fig. 3; the Qutur Su bats are identical in skull size (represented by its condylobasal length, LCb) with Kazakh samples of E. bobrinskoi and very similar to the samples of E. nilssonii and E. gobiensis. Concerning skull height (represented by neurocranium height, ANc), the Qutur Su bats are identical with the samples of E. bobrinskoi and also of E. nasutus. The samples of the latter species, however, completely differ by their skull lengths. In this comparison the samples from Qutur Su and from Kazakhstan create one Fig. 3. Scatter plot of the condylobasal length of skull (LCb) against the height of neurocranium (ANc) in the compared samples of smaller Eptesicus species of the western Palaearctic; adopted from Strelkov (1986). Data in millimetres. Obr. 3. Srovnání kondylobasální délky lebky (LCb) proti výšce neurokrania (ANc) jedinců malých druhů rodu Eptesicus západní Palearktidy; podle Strelkova (1986). Rozměry v milimetrech. 29 Table 2. Basic biometric data on the comparative samples of small representatives of the genus Eptesicus of the western Palaearctic (see Appendix). For dimension abbreviations see the text. Tab. 2. Základní biometrické údaje srovnávacích vzorků malých zástupců rodu Eptesicus západní Paleark tidy (viz Appendix). Vysvětlivky zkratek rozměrů viz text E. bobrinskoi NW Iran M min max E. bobrinskoi C Kazakhstan LAt 9 35.11 33.9 36.7 0.799 10 35.19 34.0 36.7 0.909 6 42.22 40.2 43.5 1.202 LCr LCb LaZ LaI LaInf LaN LaM ANc ACr 6 14.9214.5715.24 0.277 6 14.5614.2814.88 0.260 2 8.97 8.38 9.56 0.834 7 4.06 3.92 4.19 0.100 6 4.46 4.28 4.86 0.212 6 7.64 7.16 7.95 0.303 6 7.99 7.77 8.22 0.164 6 4.49 4.28 4.74 0.170 5 5.71 5.48 5.88 0.200 6 15.7515.1716.04 0.317 6 15.4314.9215.59 0.258 5 10.09 9.6110.28 0.276 6 4.11 3.92 4.28 0.136 6 4.96 4.75 5.11 0.148 6 8.07 7.82 8.42 0.216 6 8.59 8.27 8.81 0.201 6 5.20 4.97 5.47 0.207 6 6.47 6.32 6.68 0.121 CC P4P4 M3M3 I1M3 CM3 P4M3 M1M3 6 6 6 7 7 7 7 LMd ACo I1M3 CM3 P4M3 M1M3 5 10.4010.1410.75 0.238 6 2.97 2.81 3.14 0.137 5 6.49 6.41 6.58 0.072 5 5.56 5.42 5.67 0.105 5 4.40 4.34 4.47 0.048 5 3.74 3.64 3.85 0.083 11 10.3610.0010.76 0.272 11 3.25 3.02 3.48 0.157 10 6.46 6.27 6.68 0.134 11 5.63 5.47 5.83 0.133 11 4.51 4.42 4.74 0.095 11 3.85 3.70 4.02 0.094 6 11.1510.6711.46 0.278 6 3.41 3.29 3.52 0.095 6 6.95 6.74 7.09 0.138 6 6.09 5.93 6.28 0.116 6 4.70 4.12 4.93 0.298 6 4.10 4.02 4.17 0.061 CM3/LCb LaInf/LCb LaN/LCb LaM/LCb ANc/LCb ACr/LCb M3M3/LCb CC/CM3 M3M3/CM3 LaInf/LaI M3M3/LaI LaM/LaI LaM/LaN ACo/LMd 6 0.355 5 0.308 6 0.512 6 0.549 6 0.309 5 0.392 5 0.400 6 0.762 6 1.124 6 1.102 6 1.436 6 1.967 6 1.047 5 0.283 11 0.363 11 0.316 11 0.526 10 0.571 11 0.310 11 0.399 11 0.427 11 0.807 11 1.176 11 1.155 11 1.558 10 2.076 10 1.061 11 0.314 6 0.361 6 0.321 6 0.512 6 0.557 6 0.337 6 0.419 6 0.429 6 0.833 6 1.189 6 1.206 6 1.612 6 2.090 6 1.064 6 0.306 30 0.35 0.29 0.49 0.53 0.29 0.37 0.39 0.70 1.09 1.02 1.38 1.89 1.02 0.28 4.32 0.208 5.01 0.222 6.04 0.172 6.27 0.126 5.38 0.134 4.34 0.121 3.57 0.090 0.37 0.008 0.33 0.014 0.53 0.014 0.57 0.013 0.32 0.011 0.41 0.016 0.41 0.010 0.83 0.045 1.17 0.033 1.17 0.052 1.49 0.040 2.01 0.045 1.10 0.029 0.29 0.008 SD 11 14.7614.4215.27 0.313 11 14.3713.9714.73 0.300 11 9.54 9.20 9.86 0.228 11 3.94 3.68 4.08 0.137 11 4.55 4.33 4.88 0.180 11 7.75 7.58 7.98 0.108 10 8.23 7.93 8.56 0.199 11 4.45 4.20 4.73 0.169 11 5.73 5.52 5.94 0.126 11 11 11 11 11 11 11 4.21 5.12 6.13 6.12 5.21 4.32 3.54 4.08 4.96 6.00 5.95 5.08 4.20 3.41 0.36 0.30 0.51 0.55 0.30 0.39 0.41 0.76 1.13 1.08 1.49 1.98 1.04 0.30 4.39 0.120 5.27 0.115 6.23 0.078 6.27 0.104 5.35 0.089 4.48 0.091 3.73 0.100 0.37 0.004 0.33 0.010 0.54 0.009 0.59 0.011 0.32 0.009 0.41 0.006 0.44 0.009 0.85 0.025 1.20 0.020 1.23 0.043 1.66 0.050 2.17 0.054 1.08 0.018 0.33 0.011 n M min max n 3.74 4.49 5.61 5.94 5.02 3.97 3.35 n M min max 3.94 4.73 5.81 6.08 5.16 4.16 3.44 SD E. gobiensis N Kirghizia 6 6 6 6 6 6 6 4.64 5.45 6.63 6.63 5.58 4.54 3.67 4.52 5.31 6.47 6.47 5.38 4.41 3.58 0.35 0.31 0.50 0.54 0.32 0.41 0.42 0.80 1.15 1.15 1.52 2.00 1.04 0.29 SD 4.83 0.110 5.61 0.108 6.78 0.116 6.74 0.105 5.71 0.115 4.64 0.094 3.77 0.061 0.37 0.009 0.34 0.012 0.52 0.009 0.57 0.009 0.35 0.012 0.43 0.008 0.44 0.005 0.87 0.024 1.23 0.029 1.30 0.059 1.70 0.061 2.21 0.080 1.10 0.027 0.33 0.013 Table 2. (continued) Tab. 2. (pokračování) E. nilssonii C Europe n M min max E. n. nasutus Baluchestan SD n M min max E. n. pellucens Mesopotamia SD n M min max SD LAt 11 40.09 38.3 42.7 1.448 9 38.09 35.7 39.9 1.205 3 36.27 35.5 37.2 0.862 LCr LCb LaZ LaI LaInf LaN LaM ANc ACr 25 15.3514.8715.92 0.312 24 14.9914.4415.50 0.320 19 10.04 9.5110.44 0.238 25 4.05 3.71 4.28 0.173 25 4.91 4.62 5.27 0.194 25 7.91 7.57 8.38 0.229 25 8.54 8.21 8.84 0.210 25 5.19 4.96 5.54 0.151 24 6.64 6.23 6.88 0.161 8 13.0512.6513.39 0.255 8 12.7112.2813.18 0.317 6 8.79 8.53 8.98 0.151 8 2.99 2.75 3.13 0.123 8 4.30 4.02 4.43 0.131 8 6.30 6.17 6.42 0.075 8 6.90 6.75 7.08 0.105 8 4.53 4.42 4.75 0.127 8 5.68 5.49 5.87 0.139 6 13.2412.8813.95 0.393 6 12.9112.4813.65 0.443 5 8.87 8.39 9.83 0.564 6 2.89 2.69 3.13 0.165 6 4.40 4.02 5.09 0.396 5 6.34 6.00 6.64 0.269 6 7.15 6.91 7.54 0.249 6 4.52 4.34 4.74 0.158 6 5.57 5.36 5.82 0.169 5.11 0.153 5.86 0.160 6.63 0.191 6.88 0.142 5.74 0.120 4.61 0.129 3.73 0.082 8 8 8 8 8 8 8 4.24 4.85 5.90 5.61 4.88 4.00 3.38 3.95 4.68 5.68 5.23 4.42 3.63 2.97 4.40 0.167 4.94 0.101 6.17 0.175 5.84 0.210 5.13 0.227 4.16 0.175 3.64 0.209 6 6 6 6 6 6 6 4.34 4.98 6.01 5.69 4.81 4.00 3.40 4.04 4.66 5.68 5.42 4.64 3.82 3.27 25 11.0510.7311.44 0.208 24 3.30 3.03 3.57 0.139 25 6.98 6.61 7.28 0.162 25 5.97 5.65 6.31 0.150 25 4.67 4.42 4.92 0.118 25 3.96 3.75 4.22 0.111 8 8 8 8 8 8 9.31 3.20 5.64 5.13 4.13 3.58 8.64 3.08 5.21 4.64 3.72 3.33 9.59 0.300 3.37 0.098 5.91 0.233 5.40 0.252 4.45 0.225 3.78 0.163 6 6 6 6 6 6 9.53 3.28 5.69 5.16 4.16 3.59 9.1110.32 0.421 3.17 3.43 0.105 5.42 6.29 0.340 4.82 5.76 0.347 3.93 4.61 0.263 3.35 4.03 0.238 8 0.384 8 0.339 8 0.483 8 0.543 8 0.356 8 0.447 8 0.464 8 0.868 8 1.209 8 1.442 8 1.977 8 2.310 8 1.095 8 0.344 0.36 0.33 0.47 0.53 0.34 0.42 0.45 0.84 1.17 1.37 1.85 2.23 1.08 0.33 0.40 0.012 0.35 0.007 0.50 0.009 0.56 0.010 0.38 0.014 0.47 0.015 0.48 0.013 0.89 0.019 1.29 0.035 1.60 0.082 2.24 0.127 2.48 0.091 1.13 0.017 0.36 0.009 6 0.372 6 0.341 5 0.478 6 0.554 6 0.351 6 0.432 6 0.466 6 0.904 6 1.251 6 1.525 6 2.085 6 2.480 5 1.123 6 0.344 CC25 P4P4 25 M3M3 25 I1M3 25 CM3 25 P4M3 25 M1M3 25 LMd ACo I1M3 CM3 P4M3 M1M3 4.84 5.50 6.39 6.63 5.54 4.40 3.57 CM3/LCb 24 0.370 LaInf/LCb 24 0.328 LaN/LCb 25 0.515 LaM/LCb 24 0.570 ANc/LCb 24 0.346 ACr/LCb 23 0.444 M3M3/LCb 24 0.427 CC/CM3 25 0.875 M3M3/CM3 25 1.153 LaInf/LaI 25 1.213 M3M3/LaI 25 1.577 LaM/LaI 25 2.111 LaM/LaN 25 1.081 ACo/LMd 24 0.299 4.51 5.26 6.02 6.36 5.21 4.02 3.38 0.36 0.30 0.49 0.55 0.33 0.43 0.41 0.83 1.11 1.09 1.45 1.97 1.04 0.27 0.39 0.006 0.35 0.011 0.53 0.010 0.59 0.008 0.36 0.008 0.47 0.009 0.45 0.009 0.92 0.026 1.21 0.027 1.33 0.062 1.75 0.067 2.34 0.086 1.13 0.020 0.32 0.012 0.37 0.32 0.47 0.53 0.33 0.42 0.45 0.85 1.20 1.40 1.97 2.40 1.07 0.33 4.74 0.258 5.47 0.274 6.58 0.319 6.12 0.271 5.18 0.209 4.39 0.211 3.74 0.181 0.38 0.006 0.37 0.020 0.49 0.009 0.57 0.013 0.37 0.015 0.45 0.009 0.48 0.012 0.95 0.036 1.29 0.039 1.67 0.102 2.23 0.093 2.63 0.087 1.15 0.030 0.37 0.012 31 cluster differing from the cluster of specimens of E. nasutus and the cluster of samples of E. nilssonii and E. gobiensis. The same or very similar results were obtained also by the comparisons of skull indices (Figs. 4, 5). Two indices describing the shape of braincase (relative width, LaN/LCb, vs. relative height, ANc/LCb) grouped the compared samples into three clusters (Fig. 4); (1) Qutur Su bats and E. bobrinskoi (relatively low and wide braincase), (2) E. nilssonii and E. gobiensis (relatively high and wide braincase), and (3) E. nasutus (relatively high and narrow braincase). Two indices describing the shape of rostrum (relative length, I1M3/LCb, vs. relative width, CC/CM3) grouped the samples into two main clusters (Fig. 5); (1) Qutur Su bats, E. bobrinskoi and E. gobiensis (relatively short and narrow rostrum), and (2) E. nasutus and E. nilssonii (relative long and wide rostrum). However, the Qutur Su bats, having a relatively shortest and narrowest rostrum within the compared samples, showed to be most similar to the samples of E. bobrinskoi (together making the only dimensional overlap), while the other samples displayed a relatively much wider and mostly also longer rostrum. Finally, the analysis of principal components (PCA) showed very similar results as the above mentioned comparisons (Fig. 6). The PCA of all 38 skull dimensions (PC1 56.84% of variance, PC2 20.42%) selected a group of dimensions with a higher significance for variance (>70%) within the compared set of samples; all measurements (except for ACo) and the folowing indices: LaInf/LCb, ANc/LCb, ACr/LCb, CC/CM3, M3M3/LaI, LaM/LaI, and ACo/LMd. The PCA of these 28 dimensions (PC1 65.85% of variance; PC2 19.00%) clearly separated three clusters of samples (Fig. 6); (1) Qutur Su bats and E. bobrinskoi (PC2>0.55), (2) E. nasutus (PC1< –0.5), and (3) E. nilssonii and E. gobiensis (PC1>0.1; PC2<0.55). Fig. 4. Scatter plot of the relative width of neurocranium (LaN/LCb) against the relative height of neuro cranium (ANc/LCb) in the compared samples of smaller Eptesicus species of the western Palaearctic. Obr. 4. Srovnání relativní šířky mozkovny (LaN/LCb) proti relativní výšce mozkovny (ANc/LCb) jedinců malých druhů rodu Eptesicus západní Palearktidy. 32 Fig. 5. Scatter plot of the relative length of rostrum (I1M3/LCb) against the relative width of rostrum (CC/CM3) in the compared samples of smaller Eptesicus species of the western Palaearctic. Obr. 5. Srovnání relativní délky rostra (I1M3/LCb) proti relativní šířce rostra (CC/CM3) jedinců malých druhů rodu Eptesicus západní Palearktidy. Fig. 6. Results of the principal component analysis of skull dimensions in the compared samples of smaller Eptesicus species of the western Palaearctic. For details see Results. Fig. 6. Výsledky analysy hlavních proměnných lebečních rozměrů jedinců malých druhů rodu Eptesicus západní Palearktidy. 33 All presented morphometric comparisons showed the Qutur Su bats to be very similar in their skull characters to the samples of E. bobrinskoi from central Kazakhstan, and clearly distinguished them from all other comparative samples of Eptesicus from the western Palaearctic. Fig. 7. Skulls of small representatives of the genus Eptesicus of the western Palaearctic; in dorsal views. Scale bar = 5 mm. Obr. 7. Lebky malých zástupců rodu Eptesicus západní Palearktidy; v dorsálním pohledu. Měřítko = 5 mm. Legend / legenda: a – E. bobrinskoi (NMP 90890, female, Qutur Su, NW Iran); b – E. bobrinskoi (ZIN 61694, male, Aryskumy Desert, C Kazakhstan); c – E. nasutus (NMP 48404, female, Pir Sohrab, Balu chestan, SE Iran); d – E. gobiensis (CUP CT84/25, female, Ala-Arča NR, N Kirghizia); e – E. nilssonii (NMP 91122, female, Vrbno near Blatná, SW Bohemia, Czech Republic). 34 Figs. 7, 8 show examples of skulls of the five compared Eptesicus populations and/or taxa. The differences among the morphotypes resulting from the above analyses are well observable also in these drawings. Besides the distinctions in dimensions (Table 2), the particulars in the shapes of braincase and rostrum in the Qutur Su bats and E. bobrinskoi (an extremely low braincase, a distinct areal ratio between frontal and parietal parts of the braincase, a relatively small external auditory orifice, flattened frontal bones, narrow zygomatic arches, relative slender teeth, a flattened facial part of the skull, a relatively very narrow mesial part of the rostrum, shapes of supraorbital ridges, shapes of orbital processes of the zygomata) clearly differ from other compared bats. E. nasutus is most distinct, in comparison with the previous morphotype it has relatively much wider zygomatic arches but a much narrower braincase with smaller frontal and larger parietal bones, much narrower zygomata, a more massive lambda, more massive teeth (and of course, a unicuspidal first upper incisor) and a very short and high ante-orbital part of the rostrum with distinct supracanine swellings. Some of these differences were also observed by Harrison (1963), see this paper also for distinctions in some external features. The skulls of two very similar morphotypes, E. nilssonii and E. gobiensis, differ from the Qutur Su bats and E. bobrinskoi mainly in the relatively and absolutely much higher braincase and rostrum, distinct frontal concavities, a relatively large external auditory orifice, relatively wider mesial Fig. 8. Skulls of small representatives of the genus Eptesicus of the western Palaearctic; in lateral views. Scale bar = 5 mm. For legend see Fig. 7. Obr. 8. Lebky malých zástupců rodu Eptesicus západní Palearktidy; v laterálním pohledu. Měřítko = 5 mm. Legenda viz obr. 7. 35 parts of the rostrum, more developed orbital processes of the zygomata and more massive teeth (Figs. 7, 8). As stressed above, these two pairs of morphotypes also clearly differ in their lengths of forearm (Table 2). A baculum was extracted (Fig. 9a) from the only male among the newly collected Qutur Su bats (NMP 90892). It is a small bone, having a shape of a ‘cognac bottle’ in the dorsal aspect, 0.70 mm long and 0.45 mm broad in most wide (proximal) part; the constriction of the ‘bottle-neck’ is 0.17 mm broad. The baculum of the Qutur Su series was earlier described by Harrison (1963: 307) [No. 5] and Hill & Harrison (1987: 296) [BMNH 63.1187], however, drawings of these descriptions differ. The newly prepared baculum rather corresponds in its shape and size to that depicted by Hill & Harrison (1987), and also to one of the preparations of E. bobrin skoi from central Kazakhstan published by Strelkov (1986, 1989) (Fig. 9). Strelkov (1989) gave the following length and width ranges of baculum in E. bobrinskoi: 0.62–0.72 mm, and 0.37–0.50 mm, respectively. However, the bacula of the Qutur Su bats (Harrison 1963, Hill & Harrison 1987, own data) as well as of E. bobrinskoi (Strelkov 1986, 1989) resemble in their Fig. 9. Bacula of small representatives of the genus Eptesicus of the western Palaearctic; in dorsal views. Scale bar = 1 mm. a – original, b – after Harrison (1963), c, g – after Hill & Harrison (1987), d–f and h–q – after Strelkov (1989). Obr. 9. Bakula malých zástupců rodu Eptesicus západní Palearktidy; v dorsálním pohledu. Měřítko = 1 mm. a – originál, b – podle Harrisona (1963), c, g – podle Hilla & Harrisona (1987), d–f, h–q – podle Strelkova (1989). Legend / legenda: a–c – E. bobrinskoi (Qutur Su, Iran); d–f – E. bobrinskoi (Kazakhstan); g, h – E. nasutus (Oman, ‘Near East’); i–k – E. nilssonii (Russia, Mongolia); l–q – E. gobiensis (Mongolia, East Turkestan, Tajikistan). 36 shape (but not in size) also those of E. gobiensis (Strelkov 1986, 1989), for which Strelkov (1986) gave the dimension ranges 1.20–1.60 mm and 0.72–0.90 mm, respectively. On the other hand, the bacula of E. nilssonii and E. nasutus completely differ both in their size and shape from the above taxa (Topál 1958, Hill & Harrison 1987, Strelkov 1989; Fig. 9). It seems to be clear from the above gathered arguments that the series of the Qutur Su bats, both that published by Harrison (1963) and that newly collected by us, belong to the only morphotype. Moreover, this morphotype was found to be identical in many aspects with that of the samples of E. bobrinskoi from central Kazakhstan. The original identification of the Qutur Su bats as E. bobrinskoi by Harrison (1963) appears to be proved as well as the occurrence of this species in the Middle East. However, the most important argument from the findings by Hanák & Horáček (1986) (see Introduction) remains, considering the geographical isolation of the region of NW Iran from the core distribution area of E. bobrinskoi, i.e. the western and central parts of Kazakhstan (e.g. Strelkov 1980). The shortest distance between Qutur Su and the westernmost Kazakh site (near Kosčagyl, which is, however, 400 km west of the core area, see above) is about 1100 km, taken across the Caspian Sea. Taken round this lake, the distance can be some 1800 km along its eastern coast or 1400 km along the western coast. However, no records of E. bobrinskoi are known either in the similar landscape of eastern Azerbaijan on the western coast or in the deserts of Turkmenistan on the eastern coast (Strelkov et al. 1978, Rahmatulina 2005). It should be mentioned that the habitats of these distant regions are quite different. The Kazakh habitats of occurrence of E. bobrinskoi are situated in the zone of northern lowland deserts and semi-deserts (Butovskij et al. 1985), while the Persian locality lies in the high montane zone of alpine meadows in the altitude of around 2500 m (Figs. 1, 2). These regions represent quite distinct landscape types, sharing only some general features as the absence of trees and presumably also a harsh continental climate. The occurrence of one species in two restricted regions differing by their habitats remains enigmatic. A profound faunal investigation in the areas between them, i.e. in the northern parts of the Middle East as well as in Transcaucasia and West Turkestan, is needed to shed light on this phenomenon. However, several isolated records assigned to the smaller species of Eptesicus from the Mid dle East and adjacent areas have been discussed and/or doubted in the literature (see Hanák & Horáček 1986, Nader & Kock 1990). It is possible that some of these records may pertain to E. bobrinskoi. Only two records of E. nilssonii are known from Transcaucasia*; both were published by K. A. Satunin at the end of the 19th century and were tentatively assigned to E. n. nilssonii by Hanák & Horáček (1986). The closer record to Qutur Su comes from the approximately 70 km distant Viljaž-Čaj River near Lenkoran in south-eastern Azerbaijan, where E. nilssonii was found in 1888 (Satunin 1910). Although this record has been widely considered beyond any doubt (Ognev 1927, 1928, Kuzjakin 1950, 1965, Vereščagin 1959, Strelkov 1963, Ha nák & Horáček 1986, Rahmatulina 1990, 2005), it may well be a misidentified specimen. The regions of south-eastern Azerbaijan are covered by similar habitat types as the adjacent parts of Iran, and even by hot deserts in the lowland areas of the lower Kura River. Although * Ševčenko & Zolotuhina (2005) mentioned a third record of E. nilssonii from Transcaucasia: adult male, Georgia, Teberda, 5 July 1978, leg. E. Javrujan. However, the site Teberda lies on the northern slope of the Greater Caucasus in the Kabardino-Balkarskaja Republic, Russia, and not in Georgia. Thus, this record belongs to the main distribution range of E. nilssonii in the Caucasus Region (see Hanák & Horáček 1986) and not to the Transcancasianone. 37 E. nilssonii, rather a boreal bat species, rarely occurs in the Greater Caucasus Range (see the review by Hanák & Horáček 1986), south-eastern Transcaucasia belongs to a distinct faunal zone isolated by semi-deserts and steppes from the north (Rakhmatulina 1995). The second Transcaucasian specimen of E. nilssonii was reported by Satunin (1896) from Tiflis (= Tbilisi, Georgia), however, this isolated record seems to be more probable from the zoogeographical point of view than the former one. Since both individuals are kept in scientific collections, see Bukhnikashvili et al. (2004) and Rahmatulina (2005), these records could be revised and their species identification confirmed or disproved, respectively. In question of a possible confusion still remains also the isolated record assigned to E. nilssonii gobiensis (= E. gobiensis) by Lay (1967) and DeBlase (1980), collected at Sama in the Elborz Mts, N Iran. According to DeBlase (1980), this specimen has a relatively large skull (LCr 15.9 mm) but a relatively short forearm (LAt 37.7 mm), i.e. ratio of these dimensions in a state close to that found in E. bobrinskoi (see above). However, this record was accepted by Hanák & Horáček (1986) under DeBlase’s (1980) species affiliation. Nevertheless, a real example of confusion of species identification in the genus Eptesicus in the Middle East is a report of ‘Eptesicus nilssoni nilssoni’ by Hatt (1959) from Baghdad, Iraq, i.e. from a semi-desert lowland region of Mesopotamia. This record was accepted under Hatt’s (1959) species affiliation by several authors (Al-Robaae 1966, Lay 1967, Allison & Gaisler 1982, Hanák & Horáček 1986, Rydell 1993, Duff & Lawson 2004, Simmons 2005), although Harrison (1972) re-identifed the respective specimen as E. bottae (Peters, 1869) (see also Nader & Kock 1990 and Harrison & Bates 1991). Similarly, Pocock (1935: 442) determined a small individual of Eptesicus from Shanna, Saudi Arabia, as “a specimen apparently referable to Eptesicus hingstoni [= E. bottae hingstoni Thomas, 1919]”, while later on this bat was re-examined as E. nasutus (Nader & Kock 1990, Harrison & Bates 1991, this paper). An individual of E. anatolicus Felten, 1971 from Mala-i-Mir (= Izeh), SW Iran, was first reported by Thomas (1906) under the name “Vespertilio sp., near V. serotinus”. Later on, it was considered to pertain to E. bottae by Lay (1967) and Etemad (1969) and to E. serotinus shiraziensis (Dobson, 1871) by Gaisler (1970). Finally, it was correctly identified by DeBlase (1980); see also the analysis by Benda et al. (2006). Other examples of confusion in species identification within the genus Eptesicus are mentioned by DeBlase (1980) and Nader & Kock (1990). In conclusion, we have confirmed the occurrence of E. bobrinskoi in Iran, which seems to be isolated from the main distribution range of the species for the time being. However, the extent of this isolation may prove to be smaller after all specimens available are revised and/or more profound faunal explorations are made in the areas filling the geographical gap in between these isolated spots. SOUHRN Netopýr turanský (Eptesicus bobrinskoi Kusjakin, 1935) je obyvatelem především stepí a polopouští sever ní části Turanské nížiny mezi Aralským jezerem a Hladovou pouští (zvanou též Betpak-Dala) ve středním Kazachstanu (Butovskij et al. 1985) – z této oblasti pochází asi 90 % všech nálezů druhu. Několik nálezů tohoto netopýra bylo však hlášeno i z jiných oblastí: ze Severní Osetie (severní svah Velkého Kavkazu) a z Jakutska (Kuzjakin 1950), ze severozápadního Iranu (Harrison 1963) a ze západního a severozápadního Kazachstanu (Strelkov 1980, Davygora et al. 1998). Zatímco poslední (kazašské) nálezy jsou bez výhrad akceptovány, zpochybněn byl kavkazský, jakutský a perský nález, přičemž bylo souzeno – většinou zřejmě právem – že se jedná nejspíše o juvenilní jedince netopýra severního, E. nilssonii (Keyserling et Blasius, 1839) (Hanák & Horáček 1986). 38 Autorům se podařilo v červnu 2006 navštívit lázeňské letovisko Qutur Su (38° 20’ N, 47° 51’ E; 2545 m n. m) v severozápadním Iranu, jedinou známou lokalitu sběru E. bobrinskoi v Iranu a na Blízkém východě. Jedinci zde sebraní v roce 1961 jsou uloženi v londýnském Přírodovědeckém museu, jejich nález a určení bylo zveřejněno Harrisonem (1963); později však byli přeurčeni jako mladí jedinci E. nilssonii (Hanák & Horáček 1986). Autoři tohoto příspěvku na téže lokalitě nalezli tři mrtvé kusy drobných netopýrů rodu Eptesicus, zjevně však nikoli E. nilssonii. Proto byli jedinci určení Harrisonem (1963), jakož i nově nalezení, zevrubně porovnáni se srovnávacím materialem všech malých netopýrů rodu Eptesicus, obývajících západní Palearktidu: E. bobrinskoi, E. nilssonii, E. gobiensis a E. nasutus. Z tohoto srovnání jasně vyplynulo, že zmíněné dvě serie (celkem 10 kusů) netopýrů z Qutur Su představují nepochybně jedince E. bobrinskoi. Potvrzení výskytu netopýra turanského, který představuje zejména faunový prvek nížinných stepí a polopouští, v severozápadním Iranu, navíc na lokalitě ležící v alpinském velehorském pásmu, je velmi překvapivý. Lokalita nálezu v Iranu je nejméně 1400 km vzdálená od nejbližšího místa potvrzeného nálezu E. bobrinskoi v Kazachstanu. V příspěvku je diskutována možnost nesprávných určení některých nálezů rodu Eptesicus v oblasti vyplňující tuto mezeru (Satunin 1896, 1910, Lay 1967). V případě, že některé z nich ve skutečnosti představují nález netopýra turanského, jeho známý výskyt v Iranu by nemusel být natolik isolován, výskyt druhu v alpinské poloze velehor však záhaným zůstává nepochybně. ACKNOWLEDGEMENTS We thank Jiří Mlíkovský for his help in the field. We are obliged to Pauline Jenkins & Daphne Hill (BMNH) for enabling us to revise the Persian material of E. bobrinskoi (cf. Harrison 1963) as well as of other bats deposited in the BMNH collection, to Petr P. Strelkov (ZIN) for providing us with the comparative samples of E. bobrinskoi from Kazakhstan, and to Ivan Horáček (CUP) for enabling us to compare the material of E. gobiensis. We thank Ivan Horáček also for valuable comments on the topics. This study was supported by the Grant Agency of the Czech Republic (grants Nos. 206/02/D041 and 206/05/2334) and the Ministry of Culture of the Czech Republic (grants Nos. RK01P03OMG006, DE06P04OMG008, and MK00002327201). REFERENCES Albayrak İ., 2003: The bats of the Eastern Black Sea Region in Turkey (Mammalia: Chiroptera). Turk. J. Zool., 27: 269–273. Allison F. R. & Gaisler J., 1982: Eptesicus Rafinesque, 1820. Pp.: 172–176. In: Honacki J. H., Kinman K. E. & Koeppl J. W. (eds.): Mammal Species of the World. A Taxonomic and Geographic Reference. Allen Press, Inc. & The Association of Systematic Collections, Lawrence, ix+694 pp. Al-Robaae K., 1966: Untersuchungen der Lebensweise irakischer Fledermäuse. Säugetierk. Mitt., 14: 177–211. Anděra M. & Hanák V., 2004: Savci (Mammalia): výsledky výzkumu 1957–2004 [Mammals (Mamma lia): results of research 1957–2004]. Pp.: 227–247. In: Papáček M. (ed.): Biota Novohradských hor: modelové taxony, společenstva a biotopy [Biota of the Novohradské hory Mts: Model Taxa, Communities and Biotopes]. Jihočeská univerzita, České Budějovice, 304 pp (in Czech). Bárta Z., Červený J., Gaisler J., Hanák P., Hanák V., Horáček I., Hůrka L., Miles P., Nevrlý M., Rumler Z., Sklenář J. & Žalman J., 1981: Výsledky zimního sčítání netopýrů v Československu: 1969–1979 [Results of the winter census of bats in Czechoslovakia; 1969–1979]. Sborn. Okr. Muz. v Mostě, Ř. Přírodověd., 3: 171–116 (in Czech, with a summary in English). Bates P. J. J. & Harrison D. L., 1997: Bats of the Indian Subcontinent. Harrison Zoological Museum, Sevenoaks, 258 pp. Benda P. & Horáček I., 1998: Bats (Mammalia: Chiroptera) of the Eastern Mediterranean. Part 1. Review of distribution and taxonomy of bats in Turkey. Acta Soc. Zool. Bohem., 62: 255–313. 39 Benda P. & Hutterer R., 2005: Composition and nomenclature of the European bat fauna. Pp.: 17–22. In: Hutterer R., Ivanova T., Meyer-Cords C. & Rodrigues L. (eds.): Bat Migrations in Europe. A Review of Banding Data and Literature. Natursch. Biol. Vielfalt, 28: 1–162. Benda P., Andreas M., Kock D., Lučan R., 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 Soc. Zool. Bohem., 70 (in press). Bejček V., 1975: Nález letní kolonie netopýra severního (Eptesicus nilssoni Keyserling et Blasius, 1893) v Javorníkách [Ein Sommerquartier der Nordfledermaus (Eptesicus nilssoni Keyserling et Blasius, 1893) in Javorníky-Gebirge]. Lynx, n. s., 17: 7–9 (in Czech, with a summary in German). Bobrinskoi N., 1931: Neue Angaben über die geographische Verbreitung der Fledermäuse (Chiroptera) Rußlands. Zool. Anz., 96: 265–268. Borisenko A. V. & Pavlinov I. Ja., 1995: Podotrjad Microchiroptera [Sub-Order Microchiroptera]. Pp.: 72–120. In: Pavlinov I. Ja., Borisenko A. V., Kruskop S. V. & Jahontov E. L. (eds.): Mlekopitajuščie Evrazii. II. Non-Rodentia. Sistematiko-geografičeskij spravočnik [Mammals of Eurasia. II. Non-Roden tia. Systematic-geographical Review]. Sborn. Trud. Zool. Muz. MGU, 33: 1–334 (in Russian). Bukhnikashvili A. K., Kandaurov A. S. & Natradze I. M., 2004: Nahodki rukokrylyh v Gruzii za poslednie 140 let [Records of bats in Georgia over the last 140 years]. Plecotus et al., 7: 41–57 (in Russian, with a summary in English). Butovskij P. M., Šajmardanov R. T. & Strelkov P. P., 1985: Otrjad Rukokrylye – Chiroptera Blumenbach, 1779 [Order Bats – Chiroptera Blumenbach, 1779]. Pp.: 125–260. In: Bekenov A., Butovskij P. M., Kasabekov B. B., Lankin P. M., Strelkov P. P., Stogov I. I., Fedosenko A. K., Šajmardanov R. T. & Šubin I. G. (eds.): Mlekopitajuščie Kazahstana v četyreh tomah. Tom četvertyj. Nasekomojadnye i rukokrylye [Mammals of Kazakhstan in Four Volumes. Fourth Volume. Insectivores and Bats]. Nauka Kazahskoj SSR, Alma-Ata, 280 pp (in Russian, with a subtitle in English). Corbet G. B., 1978: The Mammals of the Palaearctic Region: a Taxonomic Review. British Museum (Natural History) and Cornell University Press, London & Ithaca, 314 pp. Davygora A. V., Il’in V. Yu. & Smirnov D. G., 1998: Novye nahodki rukokrylyh (Chiroptera, Vesperti lionidae) na juge Orenburgskoj oblasti i severo-zapade Kazahstana [New sittings of bats (Chiroptera, Vespertilionidae) in southern Orenburg District and north-western Kazakhstan]. Zool. Žurnal, 77: 984–985 (in Russian, with a summary in English). DeBlase A. F., 1980: The bats of Iran: systematics, distribution, ecology. Fieldiana Zool., N. S., 4: xvii+424 pp. Duff A. & Lawson A., 2004: Mammals of the World. A Checklist. A & C Black, London, 312 pp. Ellerman J. R. & Morrison-Scott T. C. S., 1951: Checklist of Palaearctic and Indian Mammals 1758 to 1946. British Museum (Natural History), London, 810 pp. Etemad E., 1969: Psiandaran Airan – Khafašha. Khafašhai [The bats of Iran, and the keys to identify them]. University of Tehran, Tehran, 201 pp (in Farsi, with a summary in English, paginated separately: 25 pp.). Gaisler J., 1970: The bats (Chiroptera) collected in Afghanistan by the Czechoslovak Expeditions of 1965–1967. Acta Sci. Natur. Acad. Sci. Bohemoslov. Brno, 4(6): 1–56. Gaisler J. & Hanák V., 1972: Netopýři podzemních prostorů v Československu [Bats of underground spaces in Czechoslovakia]. Sborník – Příroda (Plzeň), 7: 1–47 (in Czech, with a summary in English). Hanák V., 1959: K rozšíření netopýra severního (Eptesicus nilssoni Keyserling et Blasius 1893) v Čechách [Zur Verbreitung der nordischen Fledermaus (Eptesicus nilssoni Keyserling et Blasius 1893) in Böhmen]. Sborn. Severočes. Mus. Liberec, Přír., 1: 147–151 (in Czech, with a summary in German). Hanák V. & Gaisler J., 1971: The status of Eptesicus ognevi Bobrinskii, 1918, and remarks on some other species of this genus (Mammalia: Chiroptera). Věst. Čs. Společ. Zool., 35: 11–24. Hanák V. & Horáček I., 1986: Zur Südgrenze des Areals von Eptesicus nilssoni (Chiroptera: Vespertilio nidae). Ann. Naturhistor. Mus. Wien, 88–89 B: 297–308. Harrison D. L., 1963: Report on a collection of bats (Microchiroptera) from N. W. Iran. Ztschr. Säuge tierk., 28: 301–308. 40 Harrison D. L., 1964: The Mammals of Arabia. Volume I. Introduction, Insectivora, Chiroptera, Primates. Ernest Benn Limited, London, xx+192 pp. Harrison D. L., 1968: On three mammals new to the fauna of Oman, Arabia, with the description of a new subspecies of bat. Mammalia, 32: 317–325. Harrison D. L., 1972: The Mammals of Arabia. Volume III. Lagomorpha, Rodentia. Ernest Benn Limited, London, xvii+[288] pp. Harrison D. L. & Bates P. J. J., 1991: The Mammals of Arabia. Second edition. Harrison Zoological Museum, Sevenoaks, xvi+354 pp. Hatt R. T., 1959: The Mammals of Iraq. Miscell. Publ., Mus. Zool., Univ. Michigan, 106: 1–113, vi pl. Hill J. E. & Harrison D. L., 1987: The baculum in the Vespertilionidae (Chiroptera: Vespertilionidae) with a systematic review, a synopsis of Pipistrellus and Eptesicus, and the descriptions of a new genus and subgenus. Bull. Brit. Mus. (Natur. Hist.), Zool., 52(7): 225–305. Horáček I., Hanák V. & Gaisler J., 2000: Bats of the Palearctic region: a taxonomic and biogeographic review. Pp.: 11–157. In: Wołoszyn B. W. (ed.): Proceedings of the VIIIth European Bat Research Symposium. Vol. I. Approaches to Biogeography and Ecology of Bats. Chiropterological Information Center, Institute of Systematics and Evolution of Animals PAS, Kraków, 280 pp. Ilyin V. Ju., 2000: Dinamika arealov treh vidov rukokrylyh na krajnem jugo-vostoke Evropy [Dynamics of the ranges of three bat species in south-easternmost Europe]. Plecotus et al., 3: 43–49 (in Russian, with a summary in English). Koopman K. F., 1993: Order Chiroptera. Pp.: 137–241. In: Wilson D. E. & Reeder D. M. (eds.): Mammal Species of the World. A Taxonomic and Geographic Reference. Smithsonian Institution Press, Washing ton & London, 1206 pp. Koopman K. F., 1994: Chiroptera: Systematics. Pp.: 1–217. In: Niethammer J., Schliemann H. & Starck D. (eds.): Handbuch der Zoologie. Band VIII. Mammalia. Teilband 60. Walter de Gruyter, Berlin & New York, vii+224 pp. Kusjakin A. P. 1935: Novye dannye po sistematike i geografičeskomu rasprostraneniju letučih myšej (Chiroptera) v SSSR [Neue Angaben über Systematik und geographische Verbreitung der Fledermäuse (Chiroptera) der U. d. S. S. R.]. Bjull. Mosk. Obšč. Ispyt. Prir., Otd. Biol., 44: 428–438 (in Russian, with a summary in German). Kuzjakin A. P., 1950: Letučie myši (Sistematika, obraz žizni i pol’za dlja sel’skogo i lesnogo hozjajstva) [Bats (Systematics, Life History and Utility in Agriculture and Forestry)]. Sovetskaja Nauka, Moskva, 444 pp (in Russian). Kuzjakin A. P., 1965: Otrjad Rukokrylye. Ordo Chiroptera [Order Bats. Ordo Chiroptera]. Pp.: 79–116. In: Bobrinskij N. A., Kuznecov B. A. & Kuzjakin A. P. (eds.): Opredelitel’ mlekopitajuščih SSSR [Key to Identification of Mammals of the USSR]. Prosveščenie, Moskva, 384 pp (in Russian). Lay D. M., 1967: A study of the mammals of Iran, resulting from the Street Expedition of 1962–63. Fiel diana Zool., 54: 1–282. Nader I. A. & Kock D., 1990: Eptesicus (Eptesicus) bottae (Peters 1869) in Saudi Arabia with notes on its subspecies and distribution (Mammalia: Chiroptera: Vespertilionidae). Senckenberg. Biol., 70: 1–13. Nevrlý M., 1963: Zimoviště netopýrů v Jizerských horách [Ein Winterquartier der Fledermäuse im Iser gebirge]. Severočeské museum, Liberec, 46 pp (in Czech, with a summary in German). Nowak R. M., 1994: Walker’s Bats of the World. The Johns Hopkins University Press, Baltimore and London, 287 pp. Ognev S. I., 1927: A synopsis of the Russian bats. J. Mammal., 8: 140–157. Ognev S. I., 1928: Zveri vostočnoj Evropy i severnoj Azii. Tom I [The Mammals of the Eastern Europe and of the Northern Asia. Volume I]. Gosudarstvennoe Izdatel’stvo, Moskva & Leningrad, 631 pp (in Russian). Pavlinov I. Ja. & Rossolimo O. L., 1987: Sistematika mlekopitajuščih SSSR [Systematics of the mammals of the USSR]. Sborn. Trud. Zool. Muz. MGU, 25: 1–285 (in Russian). Pavlinov I. Ja. & Rossolimo O. L., 1998: Sistematika mlekopitajuščih SSSR. Dopolnenija [Systematics of the mammals of the USSR. Addenda]. Sborn. Trud. Zool. Muz. MGU, 38: 1–193 (in Russian). 41 Pocock R. I., 1935: The mammals collected in S.E. Arabia by Mr. Bertram Thomas and Mr. H. St. J. Philby. Ann. Mag. Natur. Hist., (10)15: 441–467. Rahmatulina I. K., 1990: Zoogeografičeskaja harakteristika fauny rukokrylyh Azerbajdžana [Zoogeo graphical characteristics of the bat fauna of Azerbaijan]. Pp.: 53–57. In: Il’in V. Ju., Strelkov P. P. & Rodionov V. A. (eds.): Rukokrylye. Materialy pjatogo Vsesojuznogo soveščanija po rukokrylym (Chiroptera) [Bats. Proceedings of the Fifth Federal Bat Conference]. Vsesojuznoe teriologičeskoe obščestvo & Penzenskij gosudarstvennyj pedagogičeskij institut imeni V. G. Belinskogo, Penza, 174 pp (in Russian, with a summary in English). Rahmatulina I. K., 2005: Rukokrylye Azerbajdžana (fauna, ekologija, zoogeografija) [Bats of Azerbai jan (Fauna, Ecology, Zoogeography)]. Nacional’naja Akademija Nauk Azerbajdžana, Baku, 480 pp (in Russian). Rakhmatulina I. K., 1995: Zoogeography of bats in the eastern Transcaucasia. Myotis, 32–33: 135–144. Revin Yu. V., Anufriev A. I. & Boeskorov G. G., 2004: Letučie myši (Mammalia, Chiroptera) Jakutii [Bats (Mammalia, Chiroptera) of Yakutia]. Plecotus et al., 7: 83–95 (in Russian, with a summary in English). Roberts T. J., 1997: The Mammals of Pakistan. Oxford University Press, Karachi, 252 pp. Rossolimo O. L. & Pavlinov I. Ja., 1979: Katalog tipovyh ekzempljarov mlekopitajuščih, hranjaščihsja v Zoologičeskom muzee MGU [Catalogue of type specimens of mammals in the collection of the Zoologi cal Museum of the Moscow State University]. Sborn. Trud. Zool. Muz. MGU, 18: 5–43 (in Russian). Rybin S. N., Horáček I. & Červený J., 1989: Bats of southern Kirghizia: distribution and faunal status. Pp.: 421–441. In: Hanák V., Horáček I. & Gaisler J. (eds.): European Bat Research 1987. Charles University Press, Praha, 720 pp. Rydell J., 1993: Eptesicus nilssonii. Mammal. Species, 430: 1–7. Šajmardanov R. T., 2001: Novye nahodki (1985–1993) redkih i maloizvestnyh dlja fauny Kazahstana vidov rukokrylyh (Chiroptera) [New records (1985–1993) of bat species rare and little investigated in the fauna of Kazakhstan]. Plecotus et al., 4: 82–83 (in Russian, with a summary in English). Satunin K. A., 1896: Vorläufige Mittheilungen über die Säugethierfauna der Kaukasusländer. Zool. Jahrb., Syst., 9: 277–314. Satunin K. A., 1910: Materialy k’ poznaniju mlekopitajuščih Kavkazskago kraja i Zakaspijskoj oblasti. XVI. Mlekopitajuščija Kavkaza i priležaščih’ stran’, hranjaščijasja v’ Zoologičeskom’ muzee Impera torskoj Akademii Nauk’ [Materials to the exploration of mammals of the Caucasus and Transcaspian Regions. XVI. Mammals of the Caucasus and adjacent regions deposited in the Zoological Museum of the Imperial Academy of Science]. Izv. Kavkaz. Muz. (Tiflis), 4: 278–284 (in Russian). Ševčenko L. S. & Zolotuhina S. I., 2005: Katalog kollekcij Zoologičeskogo muzeja NNPM NAN Ukrainy. Mlekopitajuščie. Vyp. 2. Nasekomojadnye (Insectivora), Rukokrylye (Chiroptera), Zajceobraznye (Lago morpha), Gryzuny (Rodentia) [Catalogue of the collections of the Zoological Museum of the National Museum of Natural History, National Academy of Sciences of Ukraine. Mammals. Volume 2. Insectivores (Insectivora), Bats (Chiroptera), Lagomorphs (Lagomorpha), Rodents (Rodentia)]. Zoomuzej NNPM NAN Ukrainy, Kiev, 238 pp (in Russian, with an abstract in English). Sharifi M., Hemmati Z. & Rahimi P., 2000: Distribution and conservation status of bats in Iran. Myotis, 38: 61–68. Simmons N. B., 2005: Order Chiroptera. Pp.: 312–529. In: Wilson D. E. & Reeder D. M. (eds.): Mammal Species of the World. A Taxonomic and Geographic Reference. Third Edition. Volume 1. The John Hopkins University Press, Baltimore, xxxviii+743 pp. Sklenář J., 1969: Poznámky k rozšíření netopýrů (Chiroptera) ve východních Čechách [Bemerkungen zur Verbreitung der Fledermäuse (Chiroptera) in Ostböhmen]. Acta Mus. Reginaehradec. S. A.: Sci. Natur., 10: 79–87 (in Czech, with a summary in German). Souček J., 1970: Netopýři (Microchiroptera) Chráněné krajinné oblasti Jeseníky (CHKO) [Bats (Micro chiroptera) of the Protected Landscape Area of Jeseníky]. Campanula, 1: 29–47 (in Czech). 42 Strelkov P. P., 1963: II. Otrjad Chiroptera – Rukokrylye [II. Order Chiroptera – bats]. Pp.: 122–218. In: Sokolov I. I. (ed.): Mlekopitajuščie fauny SSSR. Čast’ 1 [Mammals of the fauna of the USSR. Part 1]. Izdatel’stvo Akademii nauk SSSR, Moskva & Leningrad, 640 pp (in Russian). Strelkov P. P., 1980: Letučie myši (Chiroptera, Vespertilionidae) central’nogo i zapadnogo Kazahstana [The bats (Chiroptera, Vespertilionidae) of Central and West Kazakhstan]. Trudy Zool. Inst. Akad. Nauk SSSR, 99: 99–123 (in Russian, with a subtitle in English). Strelkov P. P., 1981: Otrjad Chiroptera Blumenbach, 1779 – Rukokrylye [Order Chiroptera Blumenbach, 1779 – Bats]. Pp.: 31–53. In: Gromov I. M. & Baranova G. I. (eds.): Katalog mlekopitajuščih SSSR. Pliocen–sovremennost’ [Catalogue of Mammals of the USSR. Pliocene–Recent]. Nauka, Leningrad, 456 pp (in Russian). Strelkov P. P., 1986: Gobijskij kožanok (Eptesicus gobiensis Bobrinskoy, 1926) – novyj vid rukokrylyh fauny Palearktiki [The Gobi bat (Eptesicus gobiensis Bobrinskoy, 1926) – a new species of bat for the fauna of the Palaearctic]. Zool. Žurnal, 65: 1103–1108 (in Russian, with a summary in English). Strelkov P. P., 1989: New data on the structure of baculum in Palaearctic bats. II. Genus Eptesicus. Pp.: 95–100. In: Hanák V., Horáček I. & Gaisler J. (eds.): European Bat Research 1987. Charles University Press, Praha, 720 pp. Strelkov P. P. & Šajmardanov R. T., 1983: Novye dannye o rosprostranenii letučih myšej (Chiroptera) v Kazahstane [New data on the distribution of bats (Chiroptera) in Kazakhstan]. Trudy Zool. Inst. Akad. Nauk SSSR, 119: 3–37 (in Russian). Strelkov P. P., Sosnovceva V. P. & Babaev N. V., 1978: Letučie myši (Chiroptera) Turkmenii [The bats of Turkmenia]. Trudy Zool. Inst. Akad. Nauk SSSR, 79: 3–71 (in Russian, with a subtitle in English). Tavrovskij V. A., Egorov O. V., Krivošeev V. G. & Popov M. V. & Labutin Ju. V., 1971: Mlekopitajuščie Jakutii [Mammals of Yakutia]. Nauka, Moskva, 660 pp (in Russian). Thomas O., 1905: A new genus and two new species of bats. Ann. Mag. Natur. Hist., (7)16: 573. Thomas O., 1906: On a collection of mammals from Persia and Armenia presented to the British Museum by Col. A. C. Bailward. Proc. Zool. Soc. London, 1905(2): 519–527. Thomas O., 1919: Some new mammals from Mesopotamia. J. Bombay Natur. Hist. Soc., 26: 745–749. Topál G., 1958: Morphological studies on the os penis of bats in the Carpathian Basin. Ann. Hist.-Natur. Mus. Natl. Hungar., 50: 331–342. Vereščagin N. K., 1959: Mlekopitajuščie Kavkaza. Istorija formirovanija fauny [The Mammals of the Caucasus. The History of Forming of the Fauna]. Izdatel’stvo Akademii Nauk SSSR, Moskva & Leningrad, 704 pp (in Russian). Zukal J. & Gaisler J., 1991: K výskytu a změnám početnosti netopýra severního, Eptesicus nilssoni (Key serling et Blasius, 1839) v Československu [On the occurrence and changes in abundance of Eptesicus nilssoni (Keyserling et Blasius, 1839) in Czechoslovakia]. Lynx, n. s., 25[1989]: 83–95 (in Czech, with a summary in English). APPENDIX List of the material examined Eptesicus nilssonii (Keyserling et Blasius, 1839) Czech Republic (19). 1 ma (NMP 91133 [S]), Dlouhá Ves, Franz-Franz Mine, Šumperk Dist., 30 January 1959, leg. V. Hanák (cf. Gaisler & Hanák 1972); – 1 fa (NMP 91144 [S+B]), Malé Karlovice, Tísňavy, school attic, Vsetín Dist., 5 June 1973, leg. V. Bejček (cf. Bejček 1975); – 3 ma, 1 ms, 2 fa, 1 fs (NMP 91136, 91138, 91139 [S+B], 91123, 91124, 91126, 91127 [S]), Mariánská Hora, gallery near the Bílá Desná Dam, Jablonec nad Nisou Dist., 24 February 1958, 13 February 1962, 2 December 1964, leg. V. Hanák (cf. Nevrlý 1963, Gaisler & Hanák 1972); – 1 ma (NMP 91128 [S]), Mikulov, Teplice Dist., 13 March 1958, leg. V. Hanák (cf. Gaisler & Hanák 1972); – 1 ma (NMP 91146 [S+B]), Orlické Záhoří, Rychnov nad Kněžnou Dist., 10 February 1977, leg. P. Rybář (cf. Bárta et al. 1981); – 1 fa (NMP 91152 [S]), Pohorská Ves, Žofín, Český Krumlov Dist., 16 June 1973, leg. V. Vohralík (cf. Anděra & Hanák 43 2004); – 1 fa (NMP 91140 [S]), Rokytnice v Orlických horách, Hanička Fortress, Rychnov nad Kněžnou Dist., 22 January 1965, leg. J. Sklenář (cf. Sklenář 1969, Gaisler & Hanák 1972); – 1 ma (NMP 91132 [S+B]), Suchá Rudná, mine, Bruntál Dist., 30 January 1959, leg. V. Hanák (cf. Souček 1970); – 1 ind. a (NMP 91151 [S]), Šumava Mts (SW Bohemia), leg. J. Červený; – 2 fa (NMP 91121, 91122 [S+B]), Vrbno near Blatná, Strakonice Dist., 4 and 5 June 1956, leg. V. Hanák (cf. Hanák 1959); – 1 ma, 1 ms (NMP 91130, 91131 [S]), Zlaté Hory, Poštovní Mine, Bruntál Dist., 29 January 1959, leg. V. Hanák (cf. Gaisler & Hanák 1972). Slovakia (5). 2 fa (NMP 91135 [S+B], 91134 [S]), Demänovská Dolina, Dračia jaskyňa [= Demänovská Ice Cave], Liptovský Mikuláš Dist., 14 February 1961, leg. V. Hanák (cf. Gaisler & Hanák 1972); – 1 ma, 1 ms (NMP 91142, 91143 [S+B]), Dobšiná, Dobšinská Ice Cave, Rožňava Dist., 16 February 1968, leg. V. Hanák (cf. Gaisler & Hanák 1972); – 1 ma (NMP 91145 [S+B]), Tatranská Javorina, Muránska jaskyňa Cave, Poprad Dist., 13 December 1973, leg. J. Gaisler & V. Hanák (cf. Zukal & Gaisler 1991). Eptesicus gobiensis Bobrinskoj, 1926 Kirghizia (6). 6 fa (CUP CT84/24–29 [S+A]), Ala-Arča Nature Reserve, 1990–2000 m a. s. l., ca. 40 km S of Biškek, 30 June 1984, leg. J. Červený & I. Horáček. Eptesicus bobrinskoi Kusjakin, 1935 Iran (10). 3 ma, 2 ms, 2 fa, 3 fs (BMNH 63.1186–1192, NMP 90890–90892 [S+A]), Qutur Su (N slope of the Mt. Sabalan, 2545 m a. s. l.), ca. 20 km ESE of Meshgin Shahr, Ardabil Prov., 21 August 1961, leg. Aberystwyth University Expedition (cf. Harrison 1963), 5 June 2006, leg. P. Benda & A. Reiter. Kazakhstan (11). 1 ma (ZIN 61694 [S+B]), Aryskumy Desert, 27 km SSE of Mustafa, 160 km N of Kzyl-Orda, Kzyl-Ordinskaja Region, 25 July 1974, leg. I. I. Stogov (cf. Strelkov 1980); – 1 fa (ZIN 62247 [S+B]), 10 km NW of Čelkar, Aktjubinskaja Region, 18 June 1975, leg. P. P. Strelkov (cf. Strelkov 1980); – 1 ma, 1 fa (ZIN 65104, 65121 [S+B]), between Džilandy and Kense, Sarysu River, Žetykanur Desert, 120 km SSE of Džezkazgan, Džezkazganskaja Region, 13 June 1977, leg. P. P. Strelkov (cf. Strelkov 1980); – 1 ma (ZIN 68618 [S+B]), Karakum Meteorologic Station, Bokdok Valley, 180 km N of Džusaly, Džezkazganskaja Region, 21 June 1980, leg. P. P. Strelkov (cf. Strelkov & Šajmardanov 1983); – 6 fa (ZIN 62240–62242, 62244–62246 [S+B]), Žetybaj well, 150 km N of Kzyl-Orda, Kzyl-Ordinskaja Region, 5 & 8 June 1975, leg. P. P. Strelkov (cf. Strelkov 1980). Eptesicus nasutus (Dobson, 1877) Afghanistan (1). 1 ma (BMNH 68.475 [S+B]), Bisut, nr. Jalalabad, 34° 26’ N, 70° 25’ E, 7 April 1967, leg. J. Gaisler (cf. Gaisler 1970). Iran (11). 3 m (BMNH 5.10.4.2, 5.10.4.4 [holotype of Vespertilio matschiei pellucens Thomas, 1906], 5.10.4.6 [S+B]), Ahwaz, Karun River, Arabistan [= Khuzestan Prov.], 220 ft, 28 March 1905, leg. R. Wo osnam (cf. Thomas 1906); – 1 ma, 2 fa (NMP 48437, 48438 [S+A], JOC [pb1722] [Sk]), 15 km E of Dehbarez, ca. 22 km W of Manujan, Kerman Prov., 17 April 2000, leg. P. Benda & A. Reiter; – 1 ma, 4 fa (NMP 48404–48408 [S+A]), Pir Sohrab, ca. 60 km NE of Chabahar, Baluchestan-ve-Sistan Prov., 12 April 2000, leg. P. Benda & A. Reiter. Iraq (2). 1 fa (BMNH 19.3.1.2 [S+A]; holotype of Eptesicus walli Thomas, 1919), Basra, leg. F. Wall (cf. Thomas 1919); – 1 m (BMNH 36.4.14.13 [S+B]), Zubier, Mesopotamia, 2 March 1921, leg. R. E. Cheesman (cf. Harrison 1964). Oman (1). 1 fa (BMNH 68.1356 [S+B]; holotype of Eptesicus nasutus batinensis Harrison, 1968), Har mul, 10 mls N of Sohar, 26 March 1967, leg. C. Seton-Browne (cf. Harrison 1968); Saudi Arabia (2). 1 ma (BMNH 48.350 [S+B]), nr. Jedda, 200 ft., 4 July 1948, leg. G. B. Popov (cf. Harrison 1964); – 1 ind. a (BMNH 34.8.8.1 [S+A]), Shanna, Arabia, [22 February], leg. J. Philby (cf. Pocock 1935). Yemen (1). 1 m (BMNH 99.11.6.19. [S+B]; holotype of Vespertilio matschiei Thomas, 1905), Jimel, W. Aden, 850 m, 16 August 1899, leg. W. Dodson (cf. Thomas 1905). 44 Lynx (Praha), n. s., 37: 45–50 (2006). ISSN 0024–7774 Karyotype of the Small-eared dormouse (Graphiurus microtis) from the Nyika Plateau, Malawi (Rodentia: Gliridae) Karyotyp plcha malouchého (Graphiurus microtis) z náhorní plošiny Nyika, Malawi (Rodentia: Gliridae) Hynek Burda1 & Wilbert N. Chitaukali2 1 Department of General Zoology, Institute of Biology, University of Duisburg-Essen, D–45317 Essen, Germany; hynek.burda@uni-due.de 2 Biology Department, Chancellor College, University of Malawi, P. O. Box 280, Zomba, Malawi received on 4 December 2006 Abstract. Karyotype of the African Small-Eared Dormouse (Graphiurus microtis) from Nganda (Nyika Plateau, Malawi) consisted of 2n=52 chromosomes: 11 pairs of metacentrics, 14 pairs of acrocentrics and subtelocentrics, the X-chromosome being a metacentric, Y-chromosome a small acrocentric. It represents a new karyotype reported for African dormice (Graphiurinae). Regarding the scarity of data (thus far ka ryotypes of only four species representing five populations, including the present data, out of currently recognized 13–14 species, distributed throughout Sub-Saharan Africa have been described) it is worth of being published but any efforts to derive conclusions regarding chromosomal speciation in graphiurines would be preliminary and speculative. Introduction Within the Gliridae, the genus Graphiurus Smuts, 1832 is unique in being an African endemic widely distributed from south of the Sahara to the Cape Province in South Africa, and in being the most specious genus of the family (13–14 species are currently recognized, whereas other glirid genera consist of a maximum of three species; Holden 1993, Rossolimo et al. 2001). Graphiurus constitutes one of the three major lineages identified among the Gliridae, justifying its subfamilial rank. The lineage is assumed to have radiated rapidly, following colonization of Africa at ca. 8–10 Myr ago (Montgelard et al. 2003). It is assumed that high rate of speciation was driven by adaptive radiation into diverse habitats ranging from dense forests to rocky areas (Montgelard et al. 2003). Still, the genus appears to be morphologically quite homogeneous (Genest-Villard 1978, Holden 1996), and the question arises whether speciation may have been driven by and/or is reflected in, chromosomal diversification, as for instance is assumed to be the case in the adaptive radiation of blind mole-rats (Spalax Guldenstaedt, 1770 spp.; cf. Nevo et al. 2001). So far karyotypes, ranging from 2n=40 to 2n=70 chromosomes, of only four species, representing five populations have been reported in the literature (Table 1). These results indicate high karyotypic variation within the genus, and its potential to become an interesting model for the study of chromosomal speciation accompanying either adaptive radiation (as exemplified by the Spalax-model, cf. Nevo et al. 2001) or by vicariance and genetic drift (as 45 Table 1. Known karyotypes (diploid numbers of chromosomes) of African dormice (Graphiurus sp.) Tab. 1. Popsané karyotypy (diploidní počty chromosomů) afrických plchů rodu Graphiurus species 2n country authors G. hueti de Rochebrune, 1883 G. murinus (Desmarest, 1822) G. parvus (True, 1893) G. microtis (Noack, 1887) G. murinus (Desmarest, 1822) 40 70 70 52 46 Ivory Coast Ivory Coast Niger Malawi South Africa Trainer & Dosso 1979 Trainer & Dosso 1979 Dobigny et al. 2002 Chitaukali et al. 2001, this paper Kryštufek et al. 2004 exemplified by the African mole-rats of the genus Fukomys Kock, Ingram, Frabotta, Burda et Honeycutt, 2006; cf. van Daele et al. 2004). Within the context of a more general report on results of a faunistic survey of small mammals in the National Park Nyika in Northern Malawi, 1997, we named the diploid number of chro mosomes we found in two dormice captured in the region, yet we did not provide any further information on this aspect (Chitaukali et al. 2001). Regarding the fact that African dormice in general are poorly known karyologically, in contrast to most of the European species, the karyotypes of which have been studied in detail in nany localities (Zima et al. 1994) as well as regarding the potential importance of this kind of information (se above), it is worthy to publish additional data on this finding. Material and Methods Two dormice were captured (in Sherman live traps) in Nganda (S10.26, E 33.51, grid 1033-B4, altitude 2,200–2,300 m a. s. l.), National Park Nyika, Northern Malawi, on alpine meadows in the vicinity of shrubs and rocks in April 1997. One further animals was trapped in the Chipome Valley (S10.20, E33.50, grid 1033-B4, altitude about 1,530 m a. s. l.), surveyed in July 1998. Dominant vegetation type at Chipome is Brachystegia woodland. In 1999, W. N. Chitaukali captured also two dormice in Mzuzu (in the garden of Mr. R. J. Murphy) (1,350 m a. s. l.). Animals were sacrificed, examined using standard mammalogical procedures. Voucher specimens (preserved in ethanol) have been deposited in the Senckenberg Museum Table 2. Measurements of small-eared dormice (Graphiurus microtis) collected in Nyika, Malawi. LC = head and body length, LCd = tail length, LTP = length of the hind foot; under breeding state either the number and distribution of embryos in uterine horns (L = left, R = right) or size of testes are given Tab. 2. Rozměry plchů malouchých (Graphiurus microtis) chycených na náhorní plošině Nyika, Malawi. sex = pohlaví, mass = hmotnost, LC = délka těla, LCd = délka ocasu, LTP = délka zadní tlapky, pinna = délka ucha; breeding state = buď počet a umístění embryí v děložních rozích (L = levý / R = pravý) anebo rozměry varlat ID locality date sex mass (g) LC (mm) LCd (mm) LTP (mm) 69/97Nganda 72/97Nganda 55/98Chipome 61/99Mzuzu 62/99Mzuzu 24.0 16.0 14.0 38.0 46.0 88.6 57.4 61.4 64.5 72.6 93.2 101.0 100.0 15.6 15.0 16.8 12.0 18.0 46 10.04. 1997 10.04. 1997 22.07. 1998 09.08. 1999 09.08. 1999 F M M F F pinna breeding state (mm) 12.0 14.0 13.2 11.5 15.0 L2 / R1 5.6×2.6 in Frankfurt am Main, Germany. Both individuals from Nganda were karyotyped using the standard preparation of bone marrow chromosomes from long bones (Lee 1969, Lee & Elder 1977). The slides were stained by Giemsa. Results and Discussion Basic biometric data are given in Table 2. Dormice from Nyika were preliminarily identified (on the base of morphological features and biogeography) as Graphiurus microtis (Noack, 1887) (determination by Dr. D. Kock, Frankfurt am Main). The specimens from Mzuzu are distinctly larger (Table 2) and may belong to a different species. Finding of a pregnant female (with two embryos) in April in Nganda suggests that the bree ding season involves at least the end of the rainy season. The karyotype of two specimens from Nganda consisted of 2n=52 chromosomes (Fig. 1): 11 pairs of metacentrics, 14 pairs of acrocentrics and subtelocentrics, the X-chromosome being a metacentric, Y-chromosome a small acrocentric. Fig. 1. Karyotype of a male Graphiurus microtis from Nganga, Nyika Plateau, Malawi. Obr. 1. Karyotyp samce plcha malouchého (Graphiurus microtis) z lokality Nganga, náhorní plošina Nyika, Malawi. 47 The karyotypes of dormice are characterized by the prevalence of biarmed autosomes (Zima et al. 1994). This is also the case in G. murinus studied by Kryštufek et al. (2004). In this respect, karyotypes of G. microtis in our study are somewhat unusual being composed of more acrocentric and telocentric chromosomes rather than by clearly biarmed chromosomes. Sex chromosomes (larger metacentric X, and dot like Y) correspond to the pattern known also from other dormouse taxa. In absence of comparative data on karyotypes of other populations of G. microtis, no conclu sion about the taxonomic status of the Nyika population can be made at the present. Regarding the fact that so far, only five populations (representing at least four different species) of Gra phiurus were karyotyped, and in absence of any chromosome banding studies, it would be very speculative to discuss differences between karyotypes from taxonomical or evolutionary point of views. Souhrn Karyotyp afrického plcha malouchého (Graphiurus microtis) z lokality Nganda (Nyika Plateau, Malawi) (obr. 2) sestává z 2n=52 chromozomů: 11 párů metacentrických, 14 párů akrocentrických a subtelocentrických, X-chromozom je metacentrický, Y-chromozom malý akrocentrický. Vzhledem k tomu, že v rámci Fig. 2. Small eared dormouse (Graphiurus microtis) from the Nyika Plateau, Malawi. Obr. 2. Plch malouchý (Graphiurus microtis) z náhorní plošiny Nyika, Malawi. 48 podčeledi Graphiurinae byl karyotyp dosud popsán (včetně této práce) jen u čtyř druhů z pěti oblastí, přičemž je uznáváno 13 až 14 druhů rozšířených v celé subsaharské Africe, představuje každá nová studie cenný příspěvek k poznání ne zcela jasné systematiky a evoluce celé čeledi. Diskuse chromozomové speciace afrických plchů na základě dostupných omezených dat by ale byla předčasná a spekulativní. Acknowledgement The work could not have been done without organisation and logistic support provided by Peter C. Over ton, organiser of the expeditions surveying biodiversity of the National Park Nyika. Assistance in the field by Jana Burda, Marianne J. Overton, Katie Ried, and scouts of the Nyika National Park is highly appreciated. Dieter Kock, Forschungsinstitut und Museum Senckenberg, Frankfurt am Main, Germany, is to be thanked for checking determination of dormice. Last but not least, we thank to the Malawi De partment of National Parks and Wildlife for the permission to work in the Nyika National Park and for all the assistance provided. References Chitaukali W. N., Burda H. & Kock D., 2001: On small mammals of the Nyika Plateau, Malawi. Pp.: 415–425. In: Denys C., Granjon L. & Poulet A. (eds.): African Small Mammals. IRD Editions, Colle ction Colloques et seminaires, Paris, 570 pp. Dobigny G., Nomao A. & Gautun J.-C., 2002: A cytotaxonomic survey of rodents from Niger: implicati ons for systematics, biodiversity and biography. Mammalia, 66: 495–523. Genest-Villard H., 1978: Revision systematique du genre Graphiurus (Rongeurs, Gliridae). Mammalia, 42: 391–426. Holden M. E., 1993: Family Myoxidae. Pp.: 763–770. In: Wilson D. W. & Reeder D. M. (eds.): Mammal Species of the World. A Taxonomic and Geographic Reference. Second Edition. Smithsonian Institution Press, Washington, 1206 pp. Holden M. E., 1996: Systematic revision of sub-Saharan African dormice (Rodentia: Myoxidae: Gra phiurus). I. An introduction to the generic revision, and a revision of Graphiurus surdus. Am. Mus. Novit., 3157: 1–44. Kock D., Ingram C. M., Frabotta L. J., Burda H. & Honeycutt R. L., 2006: On the nomenclature of Bathyergidae and Fukomys n. g. (Mammalia: Rodentia). Zootaxa, 1142: 51–55. Kryštufek B., Haber W., Baxter R. M. & Zima J., 2004: Morphology and karyology of two populations of the woodland dormouse Graphiurus murinus in the Eastern Cape, South Africa. Folia Zool., 53: 339–350. Lee M. R., 1969: A widely applicable technique for direct processing of bone marrow for chromosomes of vertebrates. Stain Technol., 44: 155–158. Lee M. R. & Elder F. F. B., 1977: Karyotypes of eight species of Mexican rodents (Muridae). J. Mammal., 58: 479–487. Montgelard C., Matthee C. A. & Robinson T. J., 2003: Molecular systematics of dormice (Rodentia: Gliridae) and the radiation of Graphiurus in Africa. Proc. R. Soc. Lond. B, 270: 1947–1955. Nevo E., Ivanitskaya E. & Beiles A., 2001: Adaptive Radiation of Blind Subterranean Mole Rats: Naming and Revisiting the Four Sibling Species of the Spalax ehrenbergi Superspecies in Israel: Spalax galili (2n=52), S. golani (2n=54), S. carmeli (2n=58), S. judaei (2n=60). Backhuys Publishers b.v., Leiden, The Netherlands, 204 pp. Rossolimo O. L., Potapova E. G., Pavlinov I. Ya., Kruskop S. V. & Voltzit O. V., 2001: Soni (Myoxi dae) mirovoj fauny [Dormice (Myoxidae) of the World]. Moscow Univ. Publ., Moscow, 233 pp (in Russian). Trainer M. & Dosso H., 1979: Recherches caryotypiques sur les rongeurs de Côte d’Ivoire: résultats préliminaire pour les milieux fermés. Mammalia, 43: 254–256. 49 Van Daele P. A. A. G., Dammann P., Kawalika M., Meier J.-L., Van De Woestijne C. & Burda H., 2004: Chromosomal diversity in Cryptomys mole-rats (Rodentia: Bathyergidae) in Zambia; with the descrip tion of new karyotypes. J. Zool. Lond., 264: 317–326. Zima J., Macholan M. & Filippucci M. G., 1994: Chromosomal variation and systematics of myoxids. Hystrix, n. s., 6(1–2): 63–76. 50 Lynx (Praha), n. s., 37: 51–66 (2006). ISSN 0024–7774 Notes on the genus Ochotona in the Middle East (Lagomorpha: Ochotonidae) Poznámky k rodu Ochotona na Blízkém východě (Lagomorpha: Ochotonidae) Stanislav Čermák1, Ján Obuch2 & Petr Benda3,4 Department of Geology and Paleontology, Charles University, Albertov 6, CZ–128 43 Praha 2, Czech Republic; stanislav.cermak@seznam.cz 2 Botanical Garden, Comenius University, SK–038 15 Blatnica, Slovakia; obuch@rec.uniba.sk 3 Department of Zoology, National Museum (Natural History), Václavské nám. 68, CZ–115 79 Praha 1, Czech Republic; petr.benda@nm.cz 4 Department of Zoology, Charles University, Viničná 7, CZ–128 44 Praha 2, Czech Republic 1 received on 14 September 2006 Abstract. New records of Ochotona from Iran and Turkey are reported. The finding of a bone remain of the Recent age (from the Eagle owl pellet) from Turkey represents the first record of the genus and family in this country and exdends the range of the genus in the Middle East by approximately 600 km to the northwest. All new records from Iran belong to O. rufescens (Gray, 1842), the only known species in the region. Based on the comparison of the Persian material, the pikas from the Zagros and Kopetdag Mts are similar in size and probably do not represent two different subspecies. The newly reported pikas of the Recent age from the Armenian Highlands of Turkey and Iran slightly differ from those inhabiting the rest of Iran and are reported as O. cf. rufescens. INTRODUCTION Only one species of ochotonid lagomorph, the rufescent or collared or Afghan pika, Ochotona rufescens (Gray, 1842), is known to occur in the Middle East (i.e. the region comprising Arabia, Iran, and the Asian part of Turkey). Its distribution range covers the mountainous areas of Afghanistan, north-western Pakistan, Iran and south-western Turkmenistan (Ellerman & Mor rison-Scott 1951, Gromov & Erbaeva 1995). Concerning the Middle East proper, the Afghan pika has been reported to reach the mountains of Iran. Taghizadeh (1964) and Lay (1967) summarised the distribution of the Afghan pika in Iran. Lay (1967: 151) using the data by Blanford (1876), Murray (1884), and Misonne (1956) and his new records, suggested that: “the rufescent pika inhabits all of the mountainous regions of Iran”. However, the westernmost Persian record of pika came from the vicinity of Agbolagh Morched, Kurdistan (Lay 1967), in the central Zagros Mts, and represented also the westernmost Recent record of the species. No records have been reported from the western Elborz Mts, the Talysh Mts or from Armenian Highlands, i.e. from all the mountains of the northwestern part of Iran (see Fig. 1 and/or Taghizadeh 1964: 25, Fig. 12). The last new records of the Afghan pika from Iran were referred by de Roguin (1988) and Obuch & Krištín (2004), including the easternmost Persian record of the species, from Mount Taftan, Baluchestan. 51 Although the talus/steppe-dwelling pikas (sensu Smith et al. 1990) are known to be regu larly distributed in the mountains from Iran and Turkmenistan to the east, several records of Ochotona have been mentioned from Transcaucasia. However, at least some of them actually are the records of fossil or sub-fossil bone fragments found in cave deposits, and their species affiliation is often not fully convincing. Sosnihina (1947, 1948) reported findings of Ochotona from Armenia; a lower jaw of sub-fossil age from a gorge of the Čirahan River and other bones from the vicinity of the Aknalič Lake. She identified these bone remains to come from pellets of the Eagle owl (Bubo bubo) and specifically attributed both records to O. rufescens according to the geographically nearest known species. Another Armenian records of pikas of sub-fossil age were reported by Dal’ (1954, 1957) from caves of the Urcskij Range. However, Dal’ (1957) Fig. 1. Records of Ochotona in the Middle East and Transcaucasia; numbers correspond with the sites numbered in the list of new records (see the text); squares – fossil or sub-fossil records (of the Upper Biharian-Toringian age), circles – Recent records, closed symbols – new records, open symbols – literature data (based on Sosnihina 1947, 1948, Coon 1951, Dal’ 1957, Gadžiev & Aliev 1966, Lay 1967, de Ro guin 1988, Montuire et. al. 1994, and Obuch & Krištín 2004). For the extralimital data on occurrence of Ochotona rufescens in Turkmenistan see Sokolov et al. (1994: 59, Fig. 18), in Afghanistan see Hassinger (1973: 34, Fig. 6), and in Pakistan see Roberts (1997: 310, map 80). Obr. 1. Místa nálezů pišťuchy (Ochotona) na Blízkém východě a v Zakavkazí; čísla odpovídají číslování lokalit v textu (kapitola New Records); čtverce – fossilní, nebo subfossilní nálezy (stáří svrchní bihar až toring), kroužky – recentní nálezy, plné symboly – nové nálezy, prázdné symboly – literární údaje (podle Sosnihiny 1947, 1948, Coona 1951, Dal’a 1957, Gadžieva & Alieva 1966, Laye 1967, de Roguina 1988, Montuireové et. al. 1994 a Obucha & Krištína 2004). Extralimitní rozšíření pišťuchy rezavé (Ochotona rufescens) v Turkmenii viz Sokolov et al. (1994: 59, Fig. 18), v Afghanistanu viz Hassinger (1973: 34, Fig. 6) a v Pakistanu viz Roberts (1997: 310, map 80). 52 placed his own and Sosnihina’s records, vernacularly named as an Armenian pika (‘armenijska ja piščuha’), into morphological proximity of Ochotona eximia (Khomenko, 1914), described from the Upper Miocene site of Taraklia in Moldavia. Vereščagin (1959) summarised these Transcaucasian records of Armenian pikas as of the Holocene age and considered the ‘species’ to be extinct in the Recent. Vekua & Šidlovskij (1958) reported on a record of a large sized ochotonid from the Mous terian, Middle Palaeolithic, of Georgia (Copi Station, Marneuli Dist., E Georgia). Later, based on an additional material from this site, Vekua (1967) described the respective pika as a new species, Ochotonoides transcaucasica (now included in the genus Ochotona, see below). Another pika species from Transcaucasia, Ochotona azerica, is reported from the Acheulean/ Mousterian deposits of the Azyh Cave, Azerbaijan (Aliev 1969, Gadžiev et al. 1979). A second Azerbaijan record of Ochotona was published by Gadžiev & Aliev (1966) from Taglar, also of the Mousterian age. All the above mentioned published records of Ochotona from Transcaucasia are reported to be of fossil to sub-fossil age (the Upper Biharian-Toringian age sensu Fejfar & Heinrich 1983; the Akhalkalaki-Akhstyr faunal units sensu Baryshnikov 2002). However, some Trans caucasian records were mentioned to come from the Eagle owl pellets but without specifying their age, i.e. whether they come from fresh pellets or from deposits of bones from supposed owl pelleting places. Vinogradov & Gromov (1952) suggested possible occurrence of O. rufescens in Armenia, based on findings of pika bones in Eagle owl pellets in the Daralegezskij Range, Mikojan [= Ehednadzor] District. Šidlovskij (1976: 72) mentioned: “remains of lower jaws of pikas have been found in the last years in southern Armenia, in the Eagle owl pellets in a cave near the village of Amagu in the Sarajbulagskij Range and in Georgia, on the eastern slope of the Džavahetskij Range (Dmanisi Dist.)” [translated from Russian]. Probably based on these reports, Hoffmann (1993) and Hoffmann & Smith (2005) mentioned Armenia among the countries inhabited by O. rufescens. On the other hand, Russian authors (i.e., Ognev 1940, Gureev 1963, 1964, Erbaeva 1988, Sokolov et al. 1994, Gromov & Erbaeva 1995, Pavlinov et al. 1995, Pavlinov & Rossolimo 1998) did not mention any Recent representative of the genus Ochotona from Transcaucasia. Moreover, Gureev (1981: 73) wrote in the paragraph concerning O. rufescens: “obviously, this species or a species close to it existed in the Caucasus at the end of the Pleistocene” [translated from Russian]. Thus, the occurrence of pikas in Transcaucasia and adjacent regions of the Middle East in the Recent remains indefinite. Here, we report new findings of Ochotona from several parts of the Middle East in attempt (1) to draw the Recent distribution range of the genus in the region of southwestern Asia, and (2) to evaluate morphologic characteristics and systematic status of particular populations of the Middle East. MATERIAL AND METHODS The most of ochotonid material described in this paper comes from owl pellets (see the list of new records below). The bone remains from owl pellets are fragmented and mostly belong to juvenile or sub-adult individuals. Only non-juvenile specimens (according to Lissovsky 2004) were included in the metric comparison. The following abbreviations are used in the text: P – premolar; M – molar; L – length; W – width; n – sample size; X – arithmetic mean; OR – observed range of the sample; MR – mandible ratio [mandible height at P3 ×100 / alveolar length of P3–M3]; PR – posteroconid ratio [posteroconid length × 100 / total tooth length]. Descriptive dental terms and metrics are used according to López-Martínez (1974), 53 Erbaeva (1988), Sen (1998), and Čermák (2004). Dimensions and drawings were made using binocular microscope. Measurements are given in millimetres and ratios in percentages. For comparative purposes, the morphometric characteristics of O. pusilla (Pallas, 1768), O. pallasi (Gray, 1867), O. rutila Severtzov, 1873, O. macrotis (Günther, 1875), and O. roylei (Ogilby, 1839) provided by Ognev (1940), Gureev (1964), and Erbajeva (1988) were taken into account. The studied material is deposited partly in the collection of the second author (JO), partly in the collection of the Department of Zoology, National Museum, Prague (NMP). NEW Records Turkey 1. 3 km SE of Güzyurdu, small cave in a mountain pass, sub-fossil deposite 2A* (Gümüşhane Prov; 39° 54’ N, 39° 34’ E, ca. 2300 m a. s. l.), 15 September 1995, isolated right M1 and a fragment of left maxilla of Ochotona cf. pusilla collected, leg. I. Horáček. 2. Ishak Paşa Sarayi, 6 km E of Doğu Bayazit, rock outcrop (Ağri Prov.; 39° 31’ N, 44° 08’ E, ca. 1950 m a. s. l.), 1 October 2002, right mandible of Ochotona cf. rufescens (NMP 90157) from a pellet of Bubo bubo collected, leg. J. Obuch. Iran 3. Bastam, 10 km W of Qarah Ziya’oddin, archaeological site (Azerbaijan Gharbi Prov., 38° 53’ N, 44° 58’ E, ca. 1250 m a. s. l.), 30 September 1998, right mandible of Ochotona cf. rufescens (NMP 90156) from a pellet of Bubo bubo collected, leg. J. Obuch. 4. rocks above Gazanak (Mazandaran Prov., 35° 55’ N, 52° 15’ E, ca. 1650 m a. s. l.), 15 May 1997, rema ins of three inds. of Ochotona rufescens from pellets of Bubo bubo (one left and three right mandibles, two maxillar fragments) collected, leg. J. Obuch. 5. 3 km E of Tangeh, 10 km SW of Raz (Khorasan Shamali Prov., 37° 54’ N, 56° 56’ E, ca. 1200 m a. s. l.), 12 May 1997, several colonies of Ochotona rufescens observed, five individuals caught, an adult male (NMP 90153 [S+A]), sub-adult male and a female (NMP 90154, 90155 [A]) collected, leg. P. Benda, J. Sádlová & R. Šumbera. 6. Qarloq, 15 km W of Bojnurd (Khorasan Shamali Prov., 37° 30’ N, 57° 26’ E, ca. 890 m a. s. l.), 12 May 1997, remains of five inds. of Ochotona rufescens from pellets of Bubo bubo (four left and five right mandibles, fragments of five skulls) collected, leg. J. Obuch. 7. two rocky valleys ca. 2 and 5 km E of Emam Qoli, ca. 30 km N of Quchan (Khorasan Razni Prov.; 37° 22’ N, 58° 32’ E, ca. 1710 m a. s. l.), 11 May 1997, remains of 30 inds. of Ochotona rufescens from pellets of Bubo bubo (30 right and 30 left mandibles, remains of 16 skulls), leg. J. Obuch; several colonies of O. rufescens observed, one individual caught, leg. J. Sádlo; 25 May 2006, colonies of O. rufescens observed, leg. A. Reiter (cf. Fig. 2). * Güzyurdu 2A: surface layer of a loamy debris infilling of a narrow karstic fissure in a small rock overhang ca. 3 km SE of the Güzyurdu village which provided the following assemblage (MNI): 1 ind. of Bufo cf. viridis, 1 Rana cf. temporaria, 1 Lacerta sp., 1 Aves indet., 1 Erinaceus sp., 1 Spermophilus cf. xantho prymnus, 1 Apodemus cf. uralensis, 3 Mesocricetus cf. brandtii, 2 Cricetulus migratorius, 1 Allactaga cf. williamsi, 1 Arvicola cf. terrestris, 4 Chionomys nivalis, 7 Microtus cf. obscurus, 2 Microtus cf. sub terraneus, 3 Spalax sp., 1 Ochotona cf. pusilla, 1 Lepus sp. The sample obtained from the lower layer of the infilling was of a similar composition (except for Ochotona, Spalax, and Erinaceus). The bones exhibit signs of fossilisation and undoubtedly are not of the Recent age. Tentatively, the assemblage is considered to be of the Holocene age (I. Horáček in litt.). The accompanying species do occur in a wider surroundings of the site even recently (cf. e.g. Kryštufek & Vohralík 2001). 54 8. dry valley ca. 5 km S of Mina, ca. 25 km SW of Dargaz (Khorasan Razni Prov.; 37° 18’ N, 58° 58’ E, ca. 1075 m a. s. l.), 22–23 May 2006, a colony of Ochotona rufescens observed, leg. A. Reiter. 9. rocks 12 km E of Bazangan, 18 km NNW of Mazdavand (Khorasan Razni Prov., 36° 17’ N, 60° 33’ E, ca. 650 m a. s. l.), 11 May 1997, remains of 16 inds. of Ochotona rufescens in pellets of Bubo bubo (14 right and 16 left mandibles, fragments of 9 skulls) collected, 8 October 2002, remains of 11 inds. of O. rufescens from pellets of Bubo bubo (7 right and 11 left mandibles) collected, leg. J. Obuch. 10. wadi under Emam Sadeh, ca. 12 km W of Kashan (Esfahan Prov., 33° 59’ N, 51° 17’ E, ca. 1130 m a. s. l.), 6 April 2000, a fragment of a left mandible of Ochotona rufescens from a pellet of Bubo bubo collected, leg. J. Obuch. 11. Deh Zireh, 25 km NNW of Natanz (Esfahan Prov., 33° 46’ N, 51° 42’ E, ca. 850 m a. s. l.), 27 April 1996, remains of five inds. of Ochotona rufescens from pellets of Bubo bubo (three left and four right mandibles, remains of one skull) collected, leg. J. Obuch. 12. Qamishlu, ca. 40 km W of Shahreza (Esfahan Prov., 32° 02’ N, 51° 29’ E, ca. 2200 m a. s. l.), 28 April 1996, remains of one skull and two mandibles (right and left) of Ochotona rufescens from a pellet of Bubo bubo collected, leg. J. Obuch. 13. 5 km NE of Deh Bakri, ca. 40 km W of Bam (Kerman Prov., 29° 05’ N, 57° 56’ E, ca. 1930 m a. s. l.), 7–8 April 2000, remains of two inds. of Ochotona rufescens from pellets of Bubo bubo (two left mandibles and one right mandible, fragments of one skull) collected, leg. J. Obuch. Fig. 2. A valley at Emam Qoli, ca. 30 km N of Quchan, Kopetdag Mts, northeastern Iran. Rocky slopes represent a typical habitat abundantly dwelled by Ochotona rufescens (photo by A. Reiter). Obr. 2. Horské údolí nedaleko Emam Koli, asi 30 km na sever od města Kučan, pohoří Kopetdag, severovýchodní Írán. Skalnaté svahy údolí vytvářejí biotop hojně obývaný pišťuchou rezavou (Ochotona rufescens) (foto A. Reiter). 55 NOTES ON BIOGEOGRAPHY New records of Ochotona of the Recent age from the Middle East come from five sub-regions (see Fig. 1); (1) northeastern Iran, the range of the Kopetdag Mts along the border with Turk menistan [records Nos. 5–9]; (2) southeastern Iran, the easternmost extent of the Zagros Mts in the Kerman Province [13]; (3) central Iran, central parts of the Zagros Mts and northern part of the Qohrud Range [10–12]; (4) central Iran, central part of the Elborz Mts [4]; (5) the border region of Iran and Turkey, the eastern part of the Armenian Highlands [2, 3]. Most of the new records come from Iran, where they cover mainly the known distribution range as described by Lay (1967). They come from all main mountainous areas of the Iran Plateau, including the central Persian mountains (Zagros Mts, Elborz Mts, Qohrud Mts) as well as the southeasternmost parts of the Zagros system, the Jebal Barez Mts, bordering the Baluchestani deserts. Records from the latter area were given already by Lay (1967) and de Roguin (1988). Only one record has been published before from the Persian side of the Kopedag Mts (Akhlamad, Khorasan Razni Prov.; Missone 1956). Newly reported data suggest the occurrence of O. rufescens in the whole main range of these mountains, similarly as it is known from the Turkmen part of the region (Sokolov et al. 1994). The new records of pikas from the border area between the northwestern corner of Iran and eastern Turkey [2, 3] are of particular significance because the extant representatives of the family had not been reported from this region before (see e.g. Lay 1967, Kumerloeve 1975). In Turkey, the finding at Ishak Paşa Sarayi represents the first record of the genus and family in this country (Kumerloeve 1975, Kryštufek & Vohralík 2001). The pika jaw was obtained from a pellet of the Eagle owl, at the site where material from pellets had been collected at least three times before, during visits in April 1996, April 1997, and September 1998, and no remnants of Ochotona were found. The Recent age of this finding is therefore undoubted and possible confusion with fossil or sub-fossil material is unlikely. Concerning the Iran side of the region, one pika record comes from Bastam, an archaeological site used as a rest place by the Eagle owl. These records shift the known Recent distribution range of the genus Ochotona some 600 km to the northwest and verify the Recent distribution of this genus in the Caucasus region (see the Introduction chapter). Moreover, they suggest that the records from Transcaucasian countries formerly considered as of rather uncertain age may also be of the Recent age (see Fig. 1). The other Turkish finding of Ochotona [record No. 1] comes from a sub-fossil deposition and its interpretation as an extant occurrence of pika in central Anatolia is irrelevant. Notes on morphology The ochotonid material collected in various parts of the Middle East is quite homogenous in morphology and dimensions. It is represented by a large sized pika (alveolar length P3–M3: X=9.33, n=27; alveolar length P2–M2: X=8.97, n=14) with a relatively long and low mandible (MR: X=59, n=27). The lower incisors extend to below the area between P4 and M1. The posterior mental foramen is located ventrally to M3, or even more posteriorly. The skull is narrow. Frontal bones with well-developed crests form the a narrow interorbital region. The relatively short rostrum possesses a large preorbital foramen of the triangular type. Nasal bones are wider anteriorly than posteriorly. The incisive-palatal foramen is not closed. Third lower premolars (P3; 9 specimens), the most characteristic teeth, are available almost only in non-adult stages; based on the morphology, they belong to not overwintered animals younger than 4 months (sensu Lissovsky 2004). These permanent teeth are worn, nevertheless 56 they still retain their conical structure. So, occlusal dimensions and morphology of the tooth (the main diagnostic features) are not stable during conversion of the crown to its definitive prismatic state, therefore evaluation of demonstrative conclusions is difficult. Differences between occlusal and root morphologies are shown on Fig. 3 (1–4, 1a–4a). Only in one tooth, a well developed lingual fold divides the posteroconid and forms a three-segmented juvenile appearance of the tooth (Fig. 3: 5). Owing to the hypselodont teeth, the alveolus of P3 corresponds in its shape with occlusal outline of P3, therefore the ratios of visible structures of both alveolus and P3 are the same and their comparison possible (see Čermák 2004). According to morphology of the available P3 Fig. 3. Morphology of P3 (non-adult) and P3 teeth in Ochotona rufescens from Iran; all teeth are figured as left specimens (1–2, 5–6, 9–10 are reversed); 1–10 – occlusal views, 1a–4a – root views; scale bar – 1 mm. Obr. 3. Morfologie třenových zubů (P3 – nedospělí jedinci; P3) u pišťuchy rezavé z Íránu; všechny zuby jsou zobrazeny jako levé (1–2, 5–6, 9–10 jsou stranově převráceny); 1–10 – okluzální pohled, 1a–4a – kořenový pohled; měřítko – 1 mm. Legend / legenda: P3: 1–3, 1a–3a, 5 – Emam Qoli, 4, 4a – Bazangan; P3: 6–8 – Emam Qoli, 9 – Qarloq, 10 – Bazangan. 57 Fig. 4. Morphology of P3 alveoli in Ochotona from the Middle East (Iran, Turkey); all alveoli are figured as left specimens (1–4, 8, 9, 21–23, 25 are reversed); scale bar – 1 mm. Obr. 4. Morfologie alveolů třetího třenového zubu (P3) u pišťuchy rezavé z Blízkého východu (Írán, Turecko); všechny alveoly jsou zobrazeny jako levé (1–4, 8, 9, 21–23, 25 jsou stranově převráceny); měřítko – 1 mm. Legend / legenda: 1 – Ishak Paşa Sarayi, 2 – Bastam, 3, 4 – Qarloq, 5 – Gazanak, 6–12 – Emam Qoli, 13–19 – Bazangan, 20–22 – Deh Zireh, 23 – Qamishlu, 24–25 – Deh Bakri. 58 alveoli (L×W = 1.80×2.00, OR = 1.55–2.00 × 1.70–2.30; n=33), the tooth is almost triangular (Fig. 4). The posteroconid is large and long (the relative length is determinable from entoconid protrusion on the lingual side of P3 alveolus – see Fig. 4). The anteroconid is small (PR of P3 alveoli: X=74, n=33), rounded, often with asymmetrical position. The P2 is (according to the shape of alveoli, n=6) oval, in its shape, its lingual side is almost straight. The P3 (Fig. 3: 6–10) is trapezoidal in the occlusal outline (L×W = 1.27×2.61, OR = 1.08–1.44 × 2.04–3.00; n=7). The mesial hypercon of P3 is relatively large (mean of hyperconal width is 1.51); its width is quite variable. There is also often an additional protrusion on the anterior part of the P3 metacon (Fig. 3: 6–8). All the above mentioned features of the compared material are characteristic for Ochotona rufescens and fall clearly within the known variation range of the species (see e.g., Ognev 1940, Erbaeva 1988). At the same time, it differs from the other species inhabiting the surrounding regions. The studied ochotonids from the Middle East, as well as all specimens of O. rufescens, are well distinguishable from O. pusilla by their much larger size, and from all of the other known Fig. 5. Skull and mandible of Ochotona rufescens from the Kopetdag Mts., NE Iran (NMP 90153) (above) and mandible of O. cf. rufescens from Ishak Paşa Sarayi, E Turkey (NMP 90157) (below); scale bar – 5 mm. Obr. 5. Lebka a spodní čelist pišťuchy rezavé (Ochotona rufescens) z pohoří Kopetdag, severovýchodní Írán (NMP 90153) (nahoře) a spodní čelist pišťuchy O. cf. rufescens z letoviska Ishak Paşa Sarayi, vý chodní Turecko (NMP 90157) (dole); měřítko – 5 mm. 59 species by their much smaller P3 anteroconid; the mean PR of P3 alveoli of geographically close species (i.e., O. pallasi, O. rutila, O. macrotis, and O. roylei) falls within the range from 62 to 65. These species differ also from the pikas from Iran in many skull features: e.g., in their closed incisive-palatal foramen (O. pallasi and O. rutila), in their distinctly wider interorbital part (O. rutila, O. macrotis, and O. roylei), or in their presence of foramina in the frontal bones (O. macrotis and O. roylei), etc. (see Ognev 1940, Gureev 1964 and Erbaeva 1988 for details). Based on the above discussed dimensions and morphological characters, the pikas from Ishak Paşa Sarayi (Turkey; record No. 2) and Bastam (NW Iran; No. 3) correspond very closely with O. rufescens and fall clearly into its variation range. Nevertheless, they differ in certain respect. These specimens are somewhat larger than other Persian pikas under study (Figs. 5, 6). However, taking into account the small size of the sample, it is difficult to evaluate the actual meaning of the difference. It seems possible either that (1) Ochotona from the border region of Iran and Turkey falls within the variation range of populations from northeastern and central Iran, while accidentally only the larger individuals were available for comparison, or (2) these western pikas are actually larger than Ochotona from northeastern and central Iran. However, the second possibility does not have to imply the existence of a separate taxon, as there could also be a cline increase in body size in pikas from east to west (the phenomenon known in such geographical arrangement in other mammals, e.g. bats, see Benda & Horáček 1995). In P3 an teroconid proportions, the pikas from Ishak Paşa Sarayi and Bastam lie at the upper margin of variation at the Persian pika samples; PR of P3 alveoli are 83 and 81, respectively. However, as there are no mandibles or additional dentition available (particularly P3), it is quite difficult to make a more conclusive comparison and taxonomical identification of the pikas from the region at the border between Iran and Turkey. Thus, we assign these specimens tentatively to O. cf. rufescens. Using the available material from Iran, the expected differences in body size between popu lations from northeastern Iran (the Kopetdag Mts, an area of occurrence of O. rufescens regina Thomas, 1911) and central Iran (central parts of the Zagros Mts, an area of occurrence of O. r. vizier Thomas, 1911) were not proved (Fig. 6). In any case, based on both morphological and metric characteristics, all the other Persian samples surveyed in this paper can be referred to O. rufescens, with an exception of the material from Emam Sadeh (record No. 10), where the lack of available features does not allow a precise species assignment. However, taking into account the geographical position of the locality, it seems very probable that the mandibular fragment belongs also to that species. NOTES ON PALEONTOLOGY The fossil record of O. rufescens, including the fossils placed in close proximity of this species, is quite poor to date (Erbaeva 1988). The ochotonid remains, almost identical with the extant O. rufescens, known from the Acheulean Palaeolithic site of Sel’ Ungur, Kirghizia, slightly differ from the Recent species by their longer diastema and wider P3 (Erbaeva 1988). Besides the ochotonid from Kirghizia, some large sized pikas referred to O. rufescens have been reported from Transcaucasia (see above). Unfortunately, no detailed characteristics of these specimens are available in the literature, and a relevant comparison is difficult. Dal’ (1957) placed the Armenian findings into proximity of Ochotona eximia (Khomenko, 1914) based on the size and morphological similarities in P3 and mandible. In his opinion, these ochotonids represent an Upper Tertiary relic derived from a heterogeneous group of species from the “eximia-gigas” 60 Fig. 6. Scatter plot of the alveolar length of the lower tooth-row (P3–M3) against the buccal height of the mandible at P3 in the samples of Ochotona coming from the Middle East (Iran, Turkey); only adult specimens were used. Obr. 6. Srovnání alveolarní délky dolní zubní řady (P3–M3) proti bukální výšce spodní čelisti u třetího třenového zubu (P3) jedinců pišťuch z Blízkého východu (Írán, Turecko); použity byly rozměry pouze dospělých jedinců. range (for details see Argyropulo & Pidopličko 1939, Agardžanjan & Erbaeva 1983, Erbaeva 1988, Erbajeva 1994, and Sen 2003). According to Averianov & Baryšnikov (1992), findings from the Čirahan River and Urcskij Range are likely to belong to Ochotona transcaucasica. This species was first described by Vekua (1967) from the Copi Station, Georgia, and due to its large size and general structure of P3 as Ochotonoides transcaucasica, i.e., belonging to a fossil genus, which was known that time from China and Hungary (see Boule & Teilhard de Chardin 1928, Teilhard de Chardin & Young 1931, Teilhard de Chardin 1940, and Kretzoi 1962). With more fossil evidence at hand, Erbaeva (1988) assigned the pika from the Copi Station to the genus Ochotona. In spite of very similar morphology (small and rounded anteroconid in P3, posterior mental foramen in mandible below M3, etc.), this species differs from the ochotonids under study by its larger size; OR of the P3–M3 length is 11.00–12.00 in O. transcaucasica (Vekua 1967) compared to 8.30–9.85, n=27, in the studied ochotonids of the Middle East. Ocho tona transcaucasica was also reported from the Azih cave in Azerbaijan; the available teeth are very close in size and morphology to those of pika from the Copi Station (Markova 1982, Erbaeva 1988). A smaller pika form found at the Azih cave is referred to Ochotona azerica (Aliev 1969, Gadžiev et al. 1979). This pika is of a similar size (OR of P3–M3 length: 8.7–9.5) 61 as O. rufescens presented in this paper, nevertheless it has a more robust mandible and a larger anterokonid of P3 (Erbaeva 1988). Besides the large sized ochotonids from the Middle East, a small sized form was found at the Güzyurdu 2A site in central Turkey (for details see New Records). Poorly preserved remains do not allow a precise species assignment. Nevertheless, in its size (L×W of the available M1 is 1.28×2.44), the pika from Güzyurdu 2A resembles rather O. pusilla than O. rufescens: OR of L×W of M1 is 1.15–1.45 × 2.00–2.40, n=20 in the extant O. pusilla in contrast to 1.50–1.85 × 2.60–3.15, n=11 in the studied O. rufescens. According to the shape of alveolus, the mesial hyperloph of P3 is relatively wide and flat; its width is approximately 75 per cent of the tooth width. Based on body size, we assign tentatively the material from Güzyurdu 2A to O. cf. pusilla. A similarly sized pika, referred to Ochotona sp., was reported by Vekua et al. (1981) from the Toringian site of Bronzovaja in western Georgia; according to its size, it is also placed into systematical proximity of O. pusilla (Vekua et al. 1981, Averianov & Baryšnikov 1992). A pika of an uncertain taxonomical position, referred to Ochotona sp., is known from the Middle Pleistocene (the Lower Toringian) site of Emirkaya-2 from the south of Central Anatolia (Montuire et. al. 1994). Its P3 shares transitional features between O. pusilla and O. rufescens, and therefore this pika is considered to be an early representative of the former species by some authors (e.g., Montuire et. al. 1994, Erbajeva 2001), by others it is referred to the latter species (e.g., Averianov 2001). Because of the limited data on fossil record, phylogenetic relationships of the taxa under study with other extant species are not yet clear. Nevertheless, from the morphological point of view it can be mentioned that O. rufescens as well as the large sized fossil ochotonids from Transcaucasia and O. pusilla share archaic dental features (i.e., P3 with small and rounded anteroconid, wide confluence between anteroconid and posteroconid, shallow proto- and paraf lexid, rounded P2 with a short anteroflexus) and their dentition is much less differentiated than in other extant species. Phylogenetic relationships within the genus Ochotona reconstructed using the mitochondrial gene for cytochrome b (Niu et al. 2004) do not support the current subgeneric classifications (see Allen 1938, Ellerman & Morrison-Scott 1951, Erbaeva 1988, Erbajeva 1994, and others). Niu et al. (2004) recognised five groups; they placed O. rufuscens into the Surrounding Quing hai-Tibet Plateau group, i.e. into phylogenetic proximity of the rock-talus-dwelling species (O. macrotis, O roylei, O. iliensis, O. himalayana, O. rutila, O erythrotis, O. gloveri, O. brookei, and O. muliensis) and intermediate types between the talus- and steppe-dwelling species (O. ladacensis, O. forresti and O. koslowi). The species divergence occurred most probably in the Early Pleistocene and was closely related to the uplifting of the Quinghai-Tibet Plateau and subsequent climatic changes. Due to the relatively stable environment, the differentiation was not so strong within the Central Asian group, including O. pusilla (Niu et al. 2004). Based on fossil remains, the structure of teeth, and molecular data, O. pusilla represents a very distinct and ancient species, which is often considered to be a relic of the Late Pliocene (Erbajeva 1994, Niu et al. 2004). SOUHRN V příspěvku jsou prezentovány nové nálezy pišťuch (12 recentních a 1 subfosilní) z území Turecka a Íránu. Většina recentních nálezů, řazených zde do druhu Ochotona rufescens (Gray, 1842) – pišťucha rezavá, pochází z horských oblastí Íránu; naše nálezy většinou spadají do dosud známého areálu rozšíření tohoto druhu. Nález kosterních pozůstatků pišťuchy ve východním Turecku představuje první recentní nález 62 v této zemi a významně posouvá hranici dosud známého areálu rozšíření rodu (i celé čeledi Ochotoni dae) na Blízkém východě. Nálezy ve východním Turecku a severozápadním Íránu (Arménská vysočina) spadají morfometricky do variační šíře pišťuchy rezavé, přesto se liší od studovaných pištuch z Íránu větší velikostí a trochu odlišnou morfologií třetího třenového zubu. Nicméně vzhledem k nepočetnému materiálu z Arménské vysočiny je věrohodné srovnání problematické a studované pišťuchy jsou provizor ně determinovány jako O. cf. rufescens. Nově nálezený subfosilní jedinec pišťuchy ze střední Anatolie je výrazně menší než předešlé pištuchy a jedná se s největší pravděpodobností o pišťuchu stepní – O. pusilla (Pallas, 1768), rozšířenou v současné době nejblíže v Předkavkazí. Uvedené nálezy byly též morfologicky srovnány s ostatními pišťuchami regionu, a to jak v kontextu neontologickém, tak paleontologickém. ACKNOWLEDGEMENTS We are very grateful to Ivan Horáček and Vladimír Vohralík (Charles University, Prague) for providing us with the material and information related to Güzyurdu (Turkey) and for numerous valuable comments. We thank A. Reiter, J. Mlíkovský, J. Sádlová, J. Sádlo, and R. Šumbera for their help with data and material collection in the field. The research was partly supported by grants GA ČR 206/05/2334 and MK00002327201. References Agadžanjan A. K. & Erbaeva M. A., 1983: Pozdnekajnozojskie gryzuny i zajceobraznye territorii SSSR [Late Cenozoic Rodents and Lagomorphs of the territory of the USSR]. Nauka, Moskva, 189 pp (in Russian). Argyropulo A. I. & Pidopličko I. G., 1939: Predstaviteli Ochotonidae (Duplicidentata, Mammalia) v plio cene SSSR [Representatives of Ochotonidae (Duplicidentata, Mammalia) in the Pliocene of the USSR]. Dokl. Akad. Nauk SSSR, 24(7): 723–728 (in Russian). Aliev S. D., 1969: Fauna Azyhskoj paleolitičeskoj stojanki [Fauna of the Palaeolithic Azyh encampment]. Candidate Dissertation Thesis, unpublished. Institute of Zoology, Baku, 30 pp (in Russian). Allen G. M., 1938: Mammals of China and Mongolia. Natural History of Central Asia. Vol. XI, Part 1. The American Museum of Natural History, New York, xxv+620 pp. Averianov A., 2001: Pleistocene lagomorphs of Eurasia. Deinsea, 8: 1–13 Averianov A. A. & Baryšnikov G. F., 1992: Plejstocenovye zajci (rod Lepus, Lagomorpha) Bol’šogo Kavkaza [Pleistocene hares of the genus Lepus (Lagomorpha) in the Greater Caucasus]. Trudy Zool. Inst. RAN, 246: 4–28 (in Russian). Baryshnikov G. F., 2002: Local biochronology of Middle and Late Pleistocene mammals from the Cau casus. Russ. J. Theriol., 1: 61–67. Benda P. & Horáček I., 1995: Geographic variation in three species of Myotis (Mammalia: Chiroptera) in South of the Western Palearctics. Acta Soc. Zool. Bohem., 59: 17–39. Blanford W. T., 1876: Eastern Persia, an account of the Journeys of the Persian Boundary Commission 1870–71–72. Volume II. The Zoology and Geology. London, vii+516 pp, 28 pls. Boule M. & Teilhard de Chardin P., 1928: Le Paleolithique de la Chine (Paleontologie). Arch. Inst. Pa leontol. Humaine (Paris), 4: 1–102, 20 pls. Coon C. S., 1951: Cave Explorations in Iran 1949. University Museum, Univ. of Pennsylvania, Philadel phia, 125 pp. Čermák S., 2004: A new ochotonid (Lagomorpha) from the Early Pleistocene of Slovakia. N. Jahrb. Geol. Paläontol. Monath., 2004: 662–680. Dal’ S. K., 1954: Paleofauna nazemnych pozvonočnych iz peščer Urcskogo chrebta [Paleofauna of terrestri al vertebrates from caves of the Urcskij range]. Izv. Akad. Nauk ArmSSR, 7(2): 61–71 (in Russian). Dal’ S. K., 1957: Zakavkazskaja piščuha [Transcaucasian pika]. Mater. Izuč. Fauny ArmSSR, 3: 17–26 (in Russian). 63 Ellerman J. R. & Morrison-Scott T. C. S., 1951: Checklist of Palaearctic and Indian Mammals 1758 to 1946. British Museum (Natural History), London, 810 pp. Erbaeva M. A., 1988: Piščuchi kajnozoja (taksonomija, sistematika, filogenija) [Cenozoic Pikas (Taxono my, Systematics, Phylogeny)]. Nauka, Moskva, 224 pp (in Russian). Erbajeva M. A., 1994: Phylogeny and evolution of Ochotonidae with emphasis on Asian ochotonids. In: Tomida Y., Li C. K. & Setoguchi T. (eds.): Rodent and Lagomorph Families of Asian Origins and Di versification. Natl. Sci. Mus. Monogr., Tokyo, 8: 1–13. Erbajeva M. A., Montuire S. & Chaline J., 2001: New ochotonids (Lagomorpha) from Pleistocene of France. Geodiversitas, 23: 395–409. Fejfar O. & Heinrich W.-D., 1983: Arvicoliden-Sukzession und Biostratigraphie des Oberpliozäns und Quartärs in Europa. In: Heinrich W.-D. (ed.): Wirbeltier-Evolution und Faunenwandel im Känozoikum. Schriftenr. Geol. Wissensch., 19–20: 61–109. Gadžiev D. V. & Aliev S. A., 1966: Piščuha (Ochotona sp.) iz mus’terskich otloženij Taglarskoj peščeri [A pika (Ochotona sp.) from the Mousterian deposits of the Taglar cave]. Učen. Zap. Azerb. Gos. Med. Inst., 20: 18–21 (in Russian). Gadžiev D. V., Gusejnov M. M., Mamedov A. V. & Širinov N. Š, 1979: Kratkie rezul’taty kompleksnyh issledovanij Azyhskoj drevněpaleolitičeskoj stojanky [Short results of the complex researches of the Lower Palaeolithic site of Azyh]. Izv. Akad. Nauk AzSSR, 3: 10–16. Gromov I. M. & Erbaeva M. A., 1995: Mlekopitajuščie fauny Rossii i sopredelenyh territorij. Zajceobraznye i gryzuny [The Mammals of Russia and Adjacent Territories. Lagomorphs and Rodents]. Zoologičeskij Institut RAN, Sankt-Peterburg, 522 pp (in Russian). Gureev A. A., 1963: III. Otrjad Lagomorpha – zajceobraznye [III. Order Lagomorpha – lagomorphs]. Pp.: 218–243. In: Gromov I. M., Gureev A. A., Novikov G. A., Sokolov I. I., Strelkov P. P. & Čapskij (eds.): Mlekopitajuščie fauny SSSR. Časť 1 [Mammals of the Fauna of the USSR. Part 1]. Izdatel’stvo Akademii Nauk SSSR, Moskva & Leningrad, 639 pp (in Russian). Gureev A. A., 1964: Fauna SSSR. Mlekopitajuščie. Tom III, vypusk 10. Zajceobraznye (Lagomorpha) [Fauna of the USSR. Mammals. Volume III, Number 10. Lagomorphs (Lagomorpha)]. Nauka, Moskva & Leningrad, 276 pp (in Russian). Gureev A. A., 1981: Otrjad Lagomorpha Brandt, 1855 – zajceobraznye [Order Lagomorpha Brandt, 1855 – lagomorphs]. Pp.: 60–74. In: Gromov I. M. & Baranova G. I. (eds.): Katalog mlekopitajuščih SSSR. Pliocen–sovremennost’ [Catalogue of Mammals of the USSR. Pliocene–Recent]. Nauka, Leningradskoe otdelenie, Leningrad, 456 pp (in Russian). Hassinger J. D., 1973: A survey of the mammals of Afghanistan resulting from the 1965 Street Expedition (excluding bats). Fieldiana Zool., 60: i–xi+1–165. Hoffmann R. S., 1993: Order Lagomorpha. Pp.: 807–827. In: Wilson D. E. & Reeder D. M. (eds.): Mammal Species of the World. A Taxonomic and Geographic Reference. Second Edition. Smithsonian Institution Press, Washington & London, 1206 pp. Hoffmann R. S. & Smith A. T., 2005: Order Lagomorpha. Pp.: 185–211. In: Wilson D. E. & Reeder D. M. (eds.): Mammals Species of the World. A Taxonomic and Geographic Reference. Third Edition. Volume 1. The Johns Hopkins University Press, Baltimore, xxxv+743 pp. Kretzoi M., 1962: Fauna und Faunenhorizont von Csarnóta. Jahresb. Ungar. Geol. Inst., 1959: 297–395. Kryštufek B. & Vohralík V., 2001: Mammals of Turkey and Cyprus. Introduction, Checklist, Insectivora. Knjižnica Annales Majora. Zgodovinsko društvo za južno Primorsko, Koper, 140 pp. Kumerloeve H., 1975: Die Säugetiere (Mammalia) der Türkei. Veröff. Zool. Staatssamm. München, 18: 69–158. Lay D. M., 1967: A study of the mammals of Iran resulting from the Street Expedition of 1962–63. Fi eldiana Zool., 54: 1–282. Lissovsky A. A., 2004: Contribution to age determination of pikas (Lagomorpha, Ochotonidae, Ochotona). Russ. J. Theriol., 3(1): 43–48. 64 López-Martínez N., 1974: Taux taxonomique d’evolution dans l’ordre des Lagomorphes (Mammalia). Bull. Soc. Géol. France, 16: 422–430. Markova A. K., 1982: Mikroteriofauna iz paleolitičeskoj peščernoj stojanky Azyh [Micromammals from the Palaeolithic cave of Azyh]. Paleontol. Sbor., 19: 14–28 (in Russian). Misonne X., 1956: Notes sur les Ochotones de l’Iran. Bull. Inst. Roy. Sci. Natur. Belg., 32(54): 1–7. Montuire S., Sen S. & Michaux J., 1994: The Middle Pleistocene mammalian fauna from Emirkaya-2, Central Anatolia (Turkey): systematics and paleoenvironment. N. Jahrb. Geol. Paläontol. Abhandl., 193(1): 107–144. Murray J. A., 1884: Additions to the present Knowledge of the Vertebrate Zoology of Persia. Ann. Mag. Natur. Hist., ser. 5, 14: 97–106. Niu Y. D., Wei F. W., Li M., Liu X. M. & Feng Z. J., 2004: Phylogeny of pikas (Lagomorpha, Ochotona) inferred from mitochondrial cytochrome b sequences. Folia Zool., 53: 141–155. Obuch J. & Krištín A., 2004: Prey composition of the little owl Athene noctua in an arid zone (Egypt, Syria, Iran). Folia Zool., 53: 65–79. Ognev S. I., 1940: Zveri SSSR i priležaščih stran. Tom 4. Gryzuny [Mammals of the USSR and Adjacent Countries. Part 4. Rodents]. Izdatel’stvo Akademii Nauk SSSR, Moskva & Leningrad, 616 pp (in Russian). Pavlinov I. Ja. & Rossolimo O. L., 1998: Sistematika mlekopitajuščih SSSR. Dopolnenija [Systematics of mammals of the USSR. Additions]. Sbor. Trud. Zool. Muz. MGU, 38: 1–190 (in Russian). Pavlinov I. Ja., Borisenko A. V., Kruskop S. V. & Jahontov E. L., 1995: Mlekopitajuščie Evrazii. II. Non-Rodentia. Sistematiko-geografičeskij spravočnik [Mammals of Eurasia. II. Non-Rodentia. Systematic and geographic review]. Sbor. Trud. Zool. Muz. MGU, 33: 1–333 (in Russian). Roberts T. J., 1997: The Mammals of Pakistan. Oxford University Press, Karachi, 252 pp. de Roguin L., 1988: Notes sur quelques mammifčres du Baluchistan iranien. Rev. Suisse Zool., 95: 595–606. Sen S., 1998: Pliocene vertebrate locality of Çalta, Ankara, Turkey. 4. Rodentia and Lagomorpha. Geo diversitas, 20: 359–378. Sen S., 2003: Lagomorpha. Pp.: 163–178. In: Fortelius M., Kappelman J., Sen S. & Bernor R. L. (eds.): Geology and Paleontology of the Miocene Sinap Formation, Turkey. Columbia University Press, New York, 448 pp. Smith A. T., Formozov N. A., Hoffmann R. S., Zheng C. L. & Erbajeva M. A., 1990: The pikas. Pp.: 14–60. In: Chapman J. A. & Flux J. E. C. (eds): Rabbits, Hares and Pikas. Status Survey and Conservation Action Plan. International Union for the Conservation of Nature, Gland (Switzerland), 168 pp. Sokolov V. E., Ivanickaja E. Ju., Gruzdev V. V. & Geptner V. G. 1994: Mlekopitajuščie Rossii i sopre delenyh regionov. Zajceobaznye [Mammals of Russia and adjacent regions. Lagomorphs]. Nauka, Moskva, 272 pp (in Russian). Sosnihina T. M., 1947: O nahoždenii kostej piščuhi (Ochotona sp.) v Armjanskoj SSR [On the finding of pika’s (Ochotona sp.) bones in Armenian SSR]. Dokl. Akad. Nauk Arm. SSR, 7(2): 77–78 (in Russian). Sosnihina T. M., 1948: Pol’za i vred filina v sel’skom hozjajstve Armjanskoj SSR [Utility and harm functions of the Eagle owl in the agriculure of Armenian SSR]. Dokl. Akad. Nauk Arm. SSR, Biol. Selsko-Hozjajst. Nauky, 1(3): 271–281 (in Russian). Šidlovskij M. V. 1976: Opredelitel’ gryzunov Zakavkaz’ja (izdanie vtoroe) [Key to Identification of Glires of Transcaucasia (Second Edition)]. Mecniereba, Tbilisi, 256 pp (in Russian). Taghizadeh F., 1964: The Harmful Rodents of Iran and their Control. Ministry of Agriculture, Tehran, 109 pp (in Farsi, with a summary in English, paginated separately: 10 pp). Teilhard de Chardin P., 1940: The fossils from Locality 18 near Peking. Paleontol. Sinica, N. S. C, 9: 1–101, 3 pls. Teilhard de Chardin P. & Young C. C., 1931: Fossil mammals from northern China. Paleontol. Sinica, S. C, 9(1): 1–87, 10 pls. Vekua A. K., 1967: Novyj predstavitel’ Lagomyidae iz paleolita Gruzii [A new representative of Lagomyi dae from the Palaeolithic of Georgia]. Soobšč. Akad. Nauk GSSR, 15(1): 139–143 (in Russian). 65 Vekua A. K. & Šidlovskij M. V., 1958: Pervaja nachodka piščuhi (Ochotona) v paleolite Kavkaza [The first finding of pika (Ochotona) from the Palaeolithic of the Caucasus]. Soobšč. Akad. Nauk GSSR, 21(3): 285–288 (in Russian) Vekua A. K., Gabelaja C. D. & Mushelišvili A. T., 1981: Paleolitičeskaja fauna pozvonočnych iz peščer Zapadnoj Gruzii [The Palaeolithic vertebrates from caves of western Georgia]. Peščery Gruzii, 9: 38–50 (in Russian). Vereščagin N. K., 1959: Mlekopitajuščie Kavkaza. Istorija formirovanija fauny [The Mammals of the Caucasus. The History of Forming of the Fauna]. Izdatel’stvo Akademii Nauk SSSR, Moskva & Leningrad, 704 pp (in Russian). Vinogradov B. S. & Gromov I. M., 1952: Gryzuny fauny SSSR [Glires of the Fauna of the USSR]. Izdatel’stvo Akademii Nauk SSSR, Moskva & Leningrad, 298 pp (in Russian). 66 Lynx (Praha), n. s., 37: 67–78 (2006). ISSN 0024–7774 Bats of the Čerchovský les Mts. and the first record of the Greater horseshoe bat (Rhinolophus ferrumequinum) in western Bohemia (Czech Republic) (Chiroptera) Netopýři Čerchovského lesa a první nález vrápence velkého (Rhinolophus ferrumequinum) v západních Čechách (Chiroptera) Jaroslav Červený1,2, Václav Fišr3, Petr Faschingbauer4 & Luděk Bufka5 Institute of Vertebrate Zoology AS CR, Květná 8, CZ–603 65 Brno, Czech Republic; Czech Agriculture University, Faculty of Forestry and Environment, Kamýcká 1176, CZ–165 21 Praha 6, Czech Republic; jardaryscerveny@centrum.cz 3 Domažlice Municipal Forests, Tyršova 611, CZ–344 01 Domažlice, Czech Republic; vaclavfisr@seznam.cz 4 Stará Huť 14, Nemanice, CZ–344 01 Domažlice 1, Czech Republic; P.Fascha@seznam.cz 5 PLA and NP Šumava Administration, Sušická 399, CZ–342 69 Kašperské Hory, Czech Republic; ludek.bufka@npsumava.cz 1 2 received on 8 December 2006 Abstract. The study summarises results of bat survey in the southern part of the Český les Mts. in the period 2004–2006. The following 17 species of bats (73.9% of bat fauna of the Czech Republic) have been confirmed to occur in the study area: Rhinolopus ferrumequinum, Myotis mystacinus, M. brandtii, M. bechsteinii, M. nattereri, M. myotis, M. daubentonii, Vespertilio murinus, Eptesicus serotinus, E. nilssonii, Pipistrellus pipistrellus s. l., P. nathusii, Nyctalus noctula, N. leisleri, Barbastella barbastellus, Plecotus auritus, and P. austriacus. Five species were recorded there for the first time: R. ferrumequinum, E. sero tinus, P. nathusii, N. noctula and N. leisleiri. Numerous records of the “forest species”: M. bechsteinii, V. murinus, E. nilssonii, P. nathusii, N. leisleri, and B. barbastellus are of particular importance. Since the record of R. ferrumequinum in the study area represents the only second record of this species in Bohemia and the seventh record in the Czech Republic after 1950, is seems to be of high importance. INTRODUCTION Although western Bohemia has been subjected to a considerable chiropterological survey (e.g., Hůrka 1973, 1986, 1989, Bufka et al. 2001, Dvořák et al. 2003), the Český les Mts. have been rather neglected. The comparison with the situation in the neighbouring Šumava Mts., which ranks among the best chiropterologically studied regions in the Czech Republic (Anděra & Červený 1994), is particularly striking. The reasons may be not only in the absence of easily controlable winter roosts and in difficult detectability of bats in forest habitats, but especially in very constrained opportunities for zoological research in this area at times, when chiropterology was flourishing in the then Czechoslovak Socialist Republic. Till 1989 much of this geomorphologically very narrow and relatively long stripe at the Czech-German border was closed for political and military regions. As a result, only 12 bat species, representing just 52.2% of the currently known Czech bat fauna were recorded there. However, neither the bordering area of 67 the Oberpfälzer Wald in Bavaria has been satisfactorily studied from the given point of view (Meschede & Rudolph 2004). Regarding these facts we decided to focus on subsequent complementation of our knowledge on the bat occurrence in a broader region of the Český les Mts. In this paper we present results of the bat survey in the southern part of the region, namely in the Čerchovský les Mts. STUDY AREA The studied area of the Čerchovský les Mts. (Fig. 1) encompasses the whole southern third of the Český Les Protected Landscape Area, with the total area of about 250 km2, and is delimited by the state border and the Oberfälzer Wald in the west and south, by Kateřinská kotlina (a basin) and Přimdský les Mts in the north, by Chodská pahorkatina (hills) and Českokubická vrchovina (highlands) in the east. The area is characterized by several elongated mountain ridges, separated by deep valleys. It is composed of three highlands: Haltravská hornatina, Nemanická vrchovina, and Ostrovská vrchovina. The highest point is Mt. Čerchov (1042 m a. s. l.), the average altitude of the area being 633.8 m a. s. l. The climate is mostly mild warm, above 700 m a. s. l. cold. The average annual temperature ranges – according to the altitude – from 8 to 4.5 °C, annual precipitation from 700 to 1000 mm. Most of the area (almost 80%) is covered by submontane or montane forests, with prevailing beech, e.g. acidophilous beechwood and fir-beechwood like Luzulo-Fagion or Eu-Fagetion, locally watered firs (Equiseto-Abietetum) and locally limited also watered spruces (Mastigobryo-Piceetum). Currently, secondary spruce monocultures prevail, yet still many mixed forests (spruce, beech, and fir, with intermixed ash, maple and sycamore), are preserved. The proportion of beech in the whole region makes up about 15%, in the southern part of Čerchovský les, and in the very region of Čerchov Mt. almost 50%. The average age of the forests is about 70 years, except for the area of Čerchov where more than 50% of trees are 111–180 years old. Remark. The locality ‘Černá Řeka’ (gallery) in this study is identical with the locality name ‘Jindřichova Hora’ used by L. Hůrka in his papers. METHODS During the period 2004–2006, bats were mainly observed in their summer and winter roosts. Bats were also caught using mistnets along their flight corridors in the forest or above water. The distribution of each bat species is given using the standard grid map (KFME system), as usual in faunistic surveys (Anděra & Červený 1994, Bufka et al. 2001). The individual findings are specified as follows: square number, site name, altitude, date of finding, and habitat or shelter. For each species, the complete list of localities arranged alphabetically in particular squares is given. Abbreviations. M – male, F – female, F gr – gravid female, F lact – lactating female, ex. – individual(s), s.i. – sex indetermined, S – summer record (15 April – 15 October), W – winter record (16 October – 14 April). RECORDS AND DISCUSSION Greater horseshoe bat, Rhinolophus ferrumequinum (Schreber, 1774) 6542: Černá Řeka (560 m a. s. l.), 4 December 2006, gallery, 2 M. Remarks. The finding of this bat species represents the first record of its occurrence in the study area, but mainly the second record in Bohemia and seventh record in the whole Czech Republic after 1950. At the same time, it represents the highest situated locality of its occurrence in the country. From the Czech Republic, there have been so far only few less reliable 68 41 42 43 63 1 3 2 64 4 CZE 5 GER 65 7 10 8 9 6 11 12 14 13 16 17 19 18 20 27 66 22 21 23 24 28 25 26 29 30 31 Fig. 1. Schematic map of the study area (grey – forested area; dashed line – border of the area under study; bigger closed circle – important settlement; smaller closed circle – locality under study; cross – important hill). Obr.1. Schematická mapa sledované oblasti (šedé pozadí – lesnatá oblast, přerušovaná linie – hranice Čerchovského lesa, větší plný kruh – důležité sídlo, menší plný kruh – sledovaná lokalita; křížek – důležitý vrchol). Sites / lokality: 1 – Železná, 2 – Smolov, 3 – Bělá nad Radbuzou, 4 – Újezd u Svatého Kříže, 5 – Rybník, 6 – Hraničná, 7 – Závist, 8 – Pivoň, 9 – Vranov, 10 – Mnichov, 11 – Lučina, 12 – Nemanice, 13 – Stará Huť, 14 – Díly, 15 – Klenčí pod Čerchov, 16 – Černá Řeka, 17 – Jindřichova Hora, 18 – Capartice, 19 – Výhledy, 20 – Chodov, 21 – Čerchov Mt., 22 – Čerchov, Hánovka (Fig. 2), 23 – Čerchov, Na zlomu, 24 – Čerchov, Zámeček, 25 – Čerchov, Rajská, 26 – Bystřice (Fig. 3), 27 – Pec, 28 – Babylon, 29 – Česká Kubice, Zelená chýše, 30 – Česká Kubice, Česká studánka, 31 – Česká Kubice. 69 data on presence of this species from the 19th and beginning of the 20th centuries; after 1950 singular occurrence is known only from six hibernating sites (Hanák & Anděra 2005). The last finding originates from the Moravian Karst in 1979 (Gaisler 1997). From Bohemia, so far only one record has been published, namely from the Mořina mine near Karlštejn in 1962 (Hanák 1962). In the neighbouring parts of Bavaria, approximately 80 km from the presented locality, in the Danube valley (between Ingolstatd and Regensburg), a stable population of this species occurs and local transitions are known there (Meschede & Rudolph 2004). The size of the Bavarian population has been found to grow (S. Morgenroth ad verb.) and hence, it is well possible that the record described here represents a winter excursion. The ability of long distance wandering in this species has been evidenced out of the Czech Republic by means of ringing (Gaisler et al. 2003). Whiskered bat, Myotis mystacinus (Kuhl, 1817) 6442: Bělá nad Radbuzou (480 m a. s. l.), 18 May 2006, shutter of a hut, 2 M; Rybník (530 m a. s. l.), 17 May 2006, shutter of a cottage, 1 M; 3 October 2006, shutter of a cottage, 2 M; 6542: Černá Řeka (560 m a. s. l.), 27 January 2004, gallery, 4 ex. s.i.; Jindřichova Hora (670 m a. s. l.), 3 October 2006, shutter of a cottage, 1 F M; Stará Huť u Nemanic (560 m a. s. l.), 22 June 2005, shutter of a hut, 3 M; 12 August 2005, shutter of a hut, 1 M; 6642: Čerchov – Na zlomu (875 m a. s. l.), 17 May 2006, wooden panelling of a hut, 1 M; 6643: Babylon – Černý rybník (470 m a. s. l.), 7 May 2006, netting above a stream near the fishpond, 2 M. Previous records. 6542: Černá Řeka (560 m a. s. l.), gallery, W, 1979–2002 (Hůrka 1986, Bufka et al. 2001, Dvořák et al. 2003); Díly (600 m a. s. l.), loft of a barn, S, 1968 (Hůrka 1973); 6642: Čerchov, 1932 (Gaisler 1956). Remarks. The whiskered bat was found at nine summer localities (470–875 m a. s. l.) in the study area. Summer records are often related to human residences at forest edges or close to water bodies, like in the Šumava Mts. (Anděra & Červený 1994). No breeding colony has been found so far. Individual bats were recorded to occupy different fissures of buildings, mainly behind window shutters. The only regular winter roost is in the gallery near Černá Řeka (560 m a. s. l.). Another record of this species is known from the nearby region – the locality Diana (Kůs 1999). The species is considered to be common in western Bohemia (Hůrka 1989, Bufka et al. 2001). Brandt’s bat, Myotis brandtii (Eversmann, 1845) 6442: Rybník (525 m a. s. l.), 17 May 2006, netting above the Radbuza river, 1 M; 6542: Stará Huť u Nemanic (560 m a. s. l.), 22 June 2005, shutter of a hut, 1 M. Previous records. 6542: Černá Řeka (560 m a. s. l.), gallery, W, 2002–2003 (Dvořák et al. 2003). Remarks. Two records of the Brandt’s bat represent the first summer occurrence (525 and 560 m a. s. l.) of this species in the study area. The only winter roost is known in the gallery near Černá Řeka (560 m a. s. l.). Habitats occupied by this species are often identical with those of M. mystacinus. Infrequent records indicate rare occurrence not only in the area under study, but also in western Bohemia in general (Hůrka 1989, Anděra & Červený 1994, Bufka et al. 2001), as well as in the whole Czech Republic (Hanák & Anděra 2006). 70 Bechstein’s bat, Myotis bechsteinii (Kuhl, 1917) 6442: Rybník (525 m a. s. l.), 17 May 2006, netting above the Radbuza river, 1 M; 6542: Černá Řeka (560 m a. s. l.), 27 January 2004, gallery, 5 ex. s.i.; 6 February 2006, 1 ex. s.i., 4 December 2006, 2 ex s.i.; 6642: Bystřice (535 m a. s. l.), 11 August 2005, netting above a small fishpond in the forest near an abandoned village, 1 M; Čerchov – Zámeček (670 m a. s. l), 23 June 2005, netting near a small fishpond in the forest, 1 M; 6643: Česká Kubice – Zelená chýše (642 m a. s. l.), 4 July 2005, netting in the forest, 1 M. Previous records. 6542: Černá Řeka (560 m a. s. l.), gallery, W, 1971–2003 (Hůrka 1973, Bufka et al. 2001, Dvořák et al. 2003). Remarks. Four records of the Bechstein’s bat represent the first summer occurrence of this species in the study area. A low number of records reflects difficult detectability of this species, rather than its relative rarity (as discussed in Hůrka 1989, Anděra & Červený 1994, Bufka et al. 2001, Hanák & Anděra 2006). The records come from middle altitudes (525–670 m a. s. l.), always from habitats with the presence of deciduous forests, mainly beech, or mixed forests. The only regular winter roost is in the gallery near Černá Řeka (560 m a. s. l.). Another record of this species is known from the nearby region – the locality Přimda (Kůs 1999). The species is considered to be rare in western Bohemia (Hůrka 1989, Bufka et al. 2001). Natterer’s bat, Myotis nattereri (Kuhl, 1817) 6442: Rybník (530 m a. s. l.), 17 May 2006, shutter of a cottage, 1 M; 6542: Capartice (760 m a. s. l.), 12 August 2005, shutter of a cottage, 1 M; Černá Řeka (560 m a. s. l.), 27 January 2004, gallery, 5 ex. s.i., 4 December 2006, 1 ex. s.i.; 6 February 2006, 5 ex. s.i.; 6642: Bystřice (555 m a. s. l.), 11 August 2005, netting in front of the cellar of a school in an abandoned village, 2 M; Bystřice (535 m a. s. l.), 11 August 2005, netting above a small fishpond in the forest near an abandoned village, 1 M; 6643: Česká Kubice (505 m a. s. l.), 13 August 2005, netting above a fishpond, 1 F. Previous records. 6542: Černá Řeka (560 m a. s. l.), gallery, W, 1974–2003 (Hůrka 1986, Bufka et al. 2001, Dvořák et al. 2003). Remarks. The Natterer’s bat is typical for lower altitudes with plenty of wetlands (Hůrka 1973, 1989, Anděra & Červený 1994, Bufka et al. 2001, Hanák & Anděra 2006). Six localities of occurrence (505–760 m a. s. l.) were found in the study area, but only some of them were situated near water. No breeding colony has been found so far. Individual bats were hidden in different fissures of buildings, mainly behind window shutters. The only regular winter roost is known in the gallery near Černá Řeka (560 m a. s. l.). The species is considered to be quite common in western Bohemia (Hůrka 1989, Bufka et al. 2001, Hanák & Anděra 2006). Greater mouse-eared bat, Myotis myotis (Borkhausen, 1797) 6442: Bělá nad Radbuzou (440 m a. s. l.), 18 May 2006, town square, 1 M dead; 6542: Černá Řeka (560 m a. s. l.), 27 January 2004, gallery, 11 ex. s.i.; 6 February 2006, 18 ex. s.i., 4 December 2006, 12 ex. s.i.; Pivoň (590 m a. s. l.), 8 November 2006, basement of the monastery, 2 ex. s.i. Previous records. 6542: Černá Řeka (560 m a. s. l.), loft of a school, S, 1967 (Hůrka 1973), gallery, W, 1969–1998 (Hůrka 1973, Hůrka 1986, Bufka et al., 2001), Pivoň (590 m a. s. l.), basement of the monastery, W, 1971–2003 (Hůrka 1973, Bufka et al. 2001, Dvořák et al. 2003). Remarks. The greater mouse-eared bat is one of the most common species of western Bohemia (Hůrka 1973, Anděra & Červený 1994, Bufka et al. 2001, Hanák & Anděra 2006), however, this does not apply for the study area. Here it is known mainly from two winter roosts (560 and 71 590 m a. s. l.). The only two summer records (440 and 560 m a. s. l.) come form the foothills. This species was recorded also in the nearby region – the locality Světce near Tachov (Hůrka 1988). Daubenton’s bat, Myotis daubentonii (Kuhl, 1817) 6442: Bělá nad Radbuzou (485 m a. s. l.), 18 May 2006, netting above a small fishpond, 1 M; Rybník (525 m a. s. l.), 17 May 2006, netting above the river Radbuza, 1 F; 6542: Černá Řeka (560 m a. s. l.), 27 January 2004, gallery, 11 ex. s.i.; 3 September 2005, netting in front of a gallery, 1 M; 6 February 2006, 18 ex. s.i., 4 December 2006, 6 ex s.i.; Lučina-Grafenried (600 m a. s. l.), 7 November 2006, cellar of a destroyed building, 1 M; 6642: Bystřice (540 m a. s. l.), 23 June 2005, netting above a small fishpond in the forest, 1 M; Bystřice (535 m a. s. l.), 11 August 2005, netting above a small fishpond in the forest near an abandoned village, 1 F; Bystřice (545 m a. s. l.), 7 November 2006, the cellar of a building in an abandoned village, 1 ex. s.i.; 6643: Babylon – Černý rybník (470 m a. s. l.), 7 May 2006, netting above a stream near the fishpond, 2 M, 3 F; Česká Kubice (505 m a. s. l.), 13 August 2005, netting above a fishpond, 2 F. Previous records. 6542: Černá Řeka (560 m a. s. l.), gallery, W, 1971–2003 (Hůrka 1973, Bufka et al. 2001, Dvořák et al. 2003). Remarks. A common species, recorded at nine localities (470–600 m a. s. l.). The Daubenton’s bat occurs in various habitats, but forages mainly above water, and its distribution is significantly influenced by the presence of various types of water bodies. No breeding colony or summer shelters have been found so far. The only regular winter roost is known in the gallery near Černá Řeka (560 m a. s. l). The species is also considered to be common in western Bohemia (Hůrka 1973, 1989, Anděra & Červený 1994, Bufka et al. 2001, Hanák & Anděra 2006). Parti-coloured bat, Vespertilio murinus Linnaeus, 1758 6441: Smolov (485 m a. s. l.), 18 May 2006, shutter of a hut, 2 M; 6442: Rybník (530 m a. s. l.), 3 October 2006, shutter of a cottage, 1 M; 6542: Stará Huť u Nemanic (560 m a. s. l.), 12 August 2005, shutter of a hut, 1 M; 3 September 2005, 2 M; 6642: Bystřice – třešňovka (540 m a. s. l.), 9 November 2006, wooden hunting lookout, 1 M dead; 6643: Pec (510 m a. s. l.), 11 June 2005, shutter of a cottage, 1 M. Previous records. 6542: Díly (600 m a. s. l.), room in a building, S, 1967 (Hůrka 1971a). Remarks. The parti-coloured bat was found at six summer localities (485–600 m a. s. l.) in the study area. No breeding or summer male colonies have been found so far, only individual bats hidden in different fissures of buildings (mainly behind window shutters) or fissures in a wooden hunting lookout. This species was formerly considered to be very rare in western Bohemia (Hůrka 1973, 1989, Bufka et al. 2001), but it is relatively common in some habitats of the Šumava Mts. (Anděra & Červený 1994). Serotine Eptesicus serotinus (Schreber, 1774) 6442: Rybník (530 m a. s. l.), 17 May 2006, loft of a house, 1 M; 6542: Nemanice (530 m a. s. l.), 23 June 2005, loft of the church, 1 ex. s.i. Remarks. Two records of the serotine represent the first occurrence of this species in the study area. Both are situated at rather middle altitudes (530 m a. s. l.), in an open landscape and in human settlements in lofts of houses. The species is common throughout western Bohemia (Hůrka 1973, Bufka et al. 2001). 72 Northern bat, Eptesicus nilssonii (Keyserling et Blasius, 1839) 6442: Bělá nad Radbuzou (485 m a. s. l.), 18 May 2006, netting above a small fishpond, 1 F gr; 6542: Černá Řeka (560 m a. s. l.), 27 January 2004, gallery, 1 ex. s.i.; Pivoň (580 m a. s. l.), 8 May 2006, netting above a small fishpond, 1 F; Stará Huť u Nemanic (560 m a. s. l.), 22 June 2005, netting above a small fishpond, 1 F lact.; 6 May 2006, shutter of a hut, 1 M; Vranov (650 m a. s. l.), 7 May 2006, loft of a building, breeding colony of approx. 50 ex. (2 F gr were examined); Závist (590 m a. s. l.), 7 May 2006, wooden panelling of a building, 1 M; 6642: 1 M; Čerchov (1040 m a. s. l.), 17 May 2006, panelling of a building on the top of the hill, 1 M; Čerchov – Zámeček (670 m a. s. l.), 23 June 2005, netting near a small fishpond in the forest, 1 F; 6643: Pec (520 m a. s. l.), 11 June 2005, loft of the building, breeding colony of approx. 80 ex. (1 F gr was examined). Previous records: 6542: Černá Řeka (560 m a. s. l.), gallery, W, 1983–2003 (Hůrka 1986, Bufka et al. 2001,); Díly (600 m a. s. l.), loft of a dwelling house, breeding colony of 70 ex., S, 1976 (Hůrka 1986). Remarks. The northern bat had been considered to be very rare for a long time, but now it seems to be a relatively common species not only in the study area, but also in the rest of western Bohemia (Anděra & Červený 1994, Bufka et al. 2001). Morever, its occurrence was confirmed at 10 localities (485–1040 m a. s. l.), mainly in woodlands. Breeding colonies were found (50–80 individuals) in lofts of dwelling houses or weekend cottages, individual bats were hidden in different fissures of buildings. The only winter roost is in the gallery near Černá Řeka (560 m a. s. l.). Other findings of this species come from the nearby region – the localities Chodovská Huť (Hůrka 1973) and Přimda (Kůs 1999). In the neighbouring Oberpfälzer Wald, a breeding colony is also known (Merkel – Wallner et al.1987) and many records were made using bat detectors (Skiba 1987). Common pipistrelle, Pipistrellus pipistrellus (Schreber, 1774) 6542: Výhledy (680 m a. s. l.), 11 June 2005, 3 October 2006, shutter of a building, 2 M; 6642: Čerchov-Hánovka (900 m a. s. l.), 29 August 2005, netting in the forest, 1 M, 1 F; 6643: Pec (510 m a. s. l.), 11 June 2005, shutter of a building, 3 M. Previous records. 6441: Železná (525 m a. s. l.), loft of the school, S, 1968 (Hůrka 1973); 6542: Klenčí pod Čerchovem (495 m a. s. l.), loft of a house, S, 1965 (Hůrka 1973). Remarks. Probably a common species in the study area, its occurrence was confirmed at five localities (495–900 m a. s. l.). Breeding colonies have not yet been found, individual bats were hidden in fissures of buildings. Although several winter roosts are known from the foothills of the Čerchovský les (Hůrka 1973, Bufka et al. 2001), no hibernating bats were recorded in the study area. The species is considered to be relatively common in western Bohemia (Hůrka 1989, Bufka et al. 2001). However, confusion with a very similar species Pipistrellus pygmeus is possible. Nathusius’ pipistrelle, Pipistrellus nathusii (Keyserling et Blasius, 1839) 6642: Bystřice (535 m a. s. l.), 4 July 2005, netting above a small fishpond in the forest near an abandoned village, 1 F lact; 6643: Česká Kubice (505 m a. s. l.), 13 August 2005, netting above a fishpond, 1 M; Česká Kubice – Zelená chýše (642 m a. s. l.), 4 July 2005, shelter in the roof of a hut, 1 M. Remarks. Three findings of the Nathusius’ pipistrelle represent the first records of this species in the study area. All are situated at rather middle altitudes (505–642 m a. s. l.) in forested areas near water bodies. The species is widespread in some parts of the Czech Republic (Jahelková 73 et al. 2000), however, this does not apply for western Bohemia where it is very rare (Červený & Bufka 1999, Bufka et al. 2001). Noctule, Nyctalus noctula (Schreber, 1774) 6542: Pivoň (580 m a. s. l.), 8 May 2006, netting above a small fishpond, 1 F; 6643: Babylon – Černý rybník (470 m a. s. l.), 7 May 2006, netting above a stream near a fishpond, 1 M. Remarks. Two findings of the noctule represent the first record of this species in the study area. The species is quite common at lower altitudes of western Bohemia (Hůrka 1989, Bufka et al. 2001). Both our records (470 and 580 m a. s. l.) come from forest habitats with water bodies. Similarly as in the lower Šumava Mts. (Červený & Bürger 1989), a sympatric occurrence of N. nocula and N. leisleri was recorded at one locality (Pivoň). Leisler’s bat, Nyctalus leisleri (Kuhl 1817) 6442: Rybník (525 m a. s. l.), 17 May 2006, netting above the river Radbuza, 1 F; 6542: Pivoň (580 m a. s. l.), 8 May 2006, netting above a small fishpond, 1 M, 2 F; 6642: Čerchov-Rajská (830 m a. s. l.), 13 November 2005, wooden hunting lookout, 2 F; 6643: Česká Kubice – Zelená chýše (642 m a. s. l.), 4 July 2005, netting in a forest, 1 F lact. Remarks. Four captures of the Leisler’s bat represent the first record of this species in the study area. The records were made at relatively higher altitudes (525–830 m a. s. l.), always in habitats with deciduous forests, mainly beech, or with mixed forests. Similarly as in the lower Šumava Mts. (Červený & Bürger 1989), a sympatric occurrence of N. nocula and N. leisleri was recorded at one locality (Pivoň). This species is considered to be extremly rare in western Bohemia (Hůrka 1989, Bufka et al. 2001). Barbastelle, Barbastella barbastellus (Schreber, 1774) 6441: Smolov (485 m a. s. l.), 18 May 2006, shutter of a cottage, 1 M; 6542: Černá Řeka (560 m a. s. l.), 27 January 2004, gallery, 7 ex. s.i.; 6 February 2006, 9 ex. s.i.; Stará Huť u Nemanic (560 m a. s. l.), 6 May 2006, shutter of a cottage, 1 M; 6642: Bystřice (555 m a. s. l.), 11 August 2005, netting in front of a cellar of the school in an abandoned village, 1 M. Previous records. 6542: Černá Řeka (560 m a. s. l.), gallery, W, 1976–1980 (Hůrka 1986); Pivoň, (590 m a. s. l.), basement of the monastery, W, 1971–1975 (Hůrka 1973, Bufka et al. 2001). Remarks. The barbastelle was found at three summer localities (485–560 m a. s. l.) in the study area. Summer findings are often related to human residences at forest edges, like in the Šumava Mts. (Anděra & Červený 1994). No breeding colony has been found so far. Individual bats were hidden behind window shutters. The only two winter roosts are situated at the altitudes of 560 and 590 m a. s. l. This species is considered to be relatively common in western Bohemia (Hůrka 1989, Anděra & Červený 1994, Bufka et al. 2001, Hanák & Anděra 2005). Brown long-eared bat, Plecotus auritus (Linnaeus, 1758) 6442: Bělá nad Radbuzou (450 m a. s. l.), 18 May 2006, shutter of a building, 4 F; Rybník (530 m a. s. l.), 23 June 2005, loft of a house, breeding colony of approx. 15 ex.; 6541: Hraničná (685 m a. s. l.), 8 November 2006, cellar of a building, 2 F; 6542: Černá Řeka (560 m a. s. l.), 27 January 2004, gallery, 6 ex. 74 Figs. 2, 3. Sites of bat netting in the Čerchovský les Mts. 2 – Mixed forest on Mount Čerchov, Hánovka (photo by V. Fišr) (above). 3 – Deforested area near the abandoned village Bystřice with Mt. Čerchov in the horizon (photo by J. Červený) (below). Obr. 2, 3. Lokality nettingu netopýrů v Čerchovském lese. 2 – Smíšený lesní porost na lokalitě Čerchov, Hánovka (foto V. Fišr) (nahoře). 3 – Odlesněná oblast u bývalé obce Bystřice s vrcholem Čerchova v pozadí (foto J. Červený) (dole). 75 s.i.; 3 September 2005, netting in front of a gallery, 1 M; 6 February 2006, gallery, 9 ex. s.i, 4 December 2006, 1 ex. s.i.; Nemanice (530 m a. s.l.), 23 June 2005, loft of the church, breeding colony of approx. 8–10 ex.; Pivoň (590 m a. s. l.), 8 November 2006, basement under the monastery, 1 F; Závist (590 m a. s. l.), 7 May 2006, loft of a cottage, breeding colony of approx. 10 ex.; 6642: Bystřice (555 m a. s. l.), 11 August 2005, netting in front of a cellar of the school in an abandoned village, 2 M, 1 F; 6642: Bystřice (545 m a. s. l.), 11 August 2005, netting in front of a cellar of a building in an abandoned village, 1 F; Bystřice (535 m a. s. l.), 11 August 2005, netting above a small fishpond in the forest near an abandoned village, 1 F; Čerchov (1040 m a. s. l.), 17 May 2006, panelling of a building on the top of the hill, 1 M; Čerchov – Zámeček (670 m a. s. l.), 23 June 2005, netting near a small fishpond in the forest, 1 F; 6643: Česká Kubice – Česká Studánka (650 m a. s. l.), 17 July 2005, bird nestbox in the forest, 8 F. Previous records. 6442: Újezd u Svatého Kříže (450 m a. s. l.), loft of the church, S, 1971 (Hůrka 1973); 6542: Černá Řeka (560 m a. s. l.), gallery, W, 1971–2003 (Hůrka 1973, Bufka et al. 2001, Dvořák et al. 2003); Díly (600 m a. s. l.), loft of the chapel, S, 1967 (Hůrka 1971b); Mnichov (725 m a. s. l.), loft of the church, S, 1974 (Hůrka 1986); Pivoň, 590 m a. s. l., basement of the monastery, W, 1971–1972 (Hůrka 1973). Remarks. The brown long-eared bat is the most common species in the study area. It was found at 14 summer localities (450–1040 m a. s. l.) and in three winter roosts (560–685 m a. s. l.). Summer findings are often related to the periphery of human settlements in forested areas, like in the Šumava Mts. (Anděra & Červený 1994). Breeding colonies (8–15 individuals) were found in lofts of dwelling houses or in a bird nestbox, individual bats were hidden in different fissures of buildings. Other findings of this species are known from the nearby parts of the Český les Mts. – the localities Halže and Chodovská Huť (Hůrka 1973), Mílov and Přimda (Kůs 1999). This species is considered to be very common in western Bohemia (Hůrka 1989, Anděra & Červený 1994, Bufka et al. 2001). Grey long-eared bat, Plecotus austriacus (Fischer, 1829) 6542: Pivoň (580 m a. s. l.), 8 May 2006, netting above a small fishpond, 1 M; 8 November 2006, basement of the monastery, 2 M; Stará Huť u Nemanic (560 m a. s. l.), 7 November 2006, cellar of a destroyed building, 1 M; 6643: Česká Kubice (505 m a. s. l.), 13 August 2005, netting above a fishpond, 1 M. Previous records: 6542: Díly (600 m a. s. l.), loft of the chapel, S, 1967 (Hůrka 1973), Chodov (500 m a. s. l.), loft of the school, S, 1966 (Hůrka 1971). Remarks. The grey long-eared bat was found only at four summer localities (500–600 m a. s. l.) and in two winter roosts (560 and 580 m a. s. l.). Summer findings are situated in an open landscape and in human settlements in lofts of houses. Other records of this species come from the nearby region – the locality Halže (Hůrka 1973). This species is considered to be very common throughout western Bohemia (Hůrka 1989, Anděra & Červený 1994, Bufka et al. 2001). SOUHRN Práce shrnuje výsledky výzkumu netopýrů Čerchovského lesa, který zaujímá celou jižní třetinu Chráněné krajinné oblasti Český les o rozloze přibližně 250 km2 a je vymezen státní hranicí (resp. Oberfälzer Wald) na západě a jihu, Kateřinskou kotlinou a Přimdským lesem na severu, Chodskou pahorkatinou a Českokubickou vrchovinou na východě. Území je charakteristické několika podélnými hřbety oddělenými hlubokými údolími. Skládá se ze tří geomorfologických okrsků: Haltravské hornatiny, Nemanické vrchoviny a Ostrovské vrchoviny. Nejvyšší kótou je Čerchov (1042 m n. m.), střední výška území je 633,8 m n. m. Podnebí je převážně mírně teplé (MT 3), nad 700 m n. m. chladné (CH 7). Průměrná roční teplota se podle nadmořské výšky pohybuje od 8 do 4,5 °C, průměrné roční srážky od 700 do 1000 mm. 76 Většinu území pokrývají lesy submontánního až montánního stupně, lesnatost území dosahuje hodnoty téměř 80 %. Rekokonstrukčně převládají acidofilní bučiny a jedlobučiny (Luzulo-Fagion), podmáčené jedliny (Equiseto-Abietetum), místy květnaté bučiny (Eu-Fagetion), omezeně podmáčené smrčiny (Mas tigobryo-Piceetum). V současnosti sice převládají stanoviště sekundárních monokultur smrku, přesto se však zde zachovalo více smíšených porostů (smrk, buk a jedle s vtroušeným jasanem, javorem a klenem), než jinde v České republice. Podíl buku v celé oblasti činí okolo 15 %, v jižní části Čerchovského lesa, a zvláště pak v oblasti samotného Čerchova téměř 50 %. Průměrný věk lesních porostů je přibližně 70 let, v oblasti Čerchova je však více než 50 % ve věku 111–180 let. V letech 2004–2006 byl ve sledované oblasti potvrzen výskyt 17 druhů netopýrů, což představuje 73,9 % netopýří fauny celé České republiky. Zjištěni byli: vrápenec velký (Rhinolophus ferrumequinum), netopýr vousatý (Myotis mystacinus), netopýr Brandtův (Myotis brandtii), netopýr velkouchý (Myotis bechsteinii), netopýr řasnatý (Myotis nattereri), netopýr velký (Myotis myotis), netopýr vodní (Myotis daubentonii), netopýr pestrý (Vespertilio murinus), netopýr pozdní (Eptesicus serotinus), netopýr severní (Eptesicus nilssonii), netopýr hvízdavý (Pipistrellus pipistrellus), netopýr parkový (Pipistrellus nathusii), netopýr rezavý (Nyctalus noctula), netopýr stromový (Nyctalus leisleri), netopýr černý (Barbastella barbastellus), netopýr ušatý (Plecotus auritus) a netopýr dlouhouchý (Plecotus austriacus). Druhy R. ferrumequinum, E. serotinus, P. nathusii, N. noctula a N. leisleri byly zjištěny v oblasti poprvé. Nejhodnotnější údaje představují početné nálezy tzv. “lesních druhů”: M. bechsteinii, V. murinus, E. nilssonii, P. nathusii, N. leisleri a B. barbastellus, které jsou v České republice obecně považovány za druhy vzácné, řídké, nebo alespoň méně běžné. Nález Rh. ferrumequinum je zcela ojedinělý a zárověň velmi významně přispívá k poznání výskytu tohoto druhu u nás. Je teprve druhým prokázáným výskytem v Čechách a sedmým v celé České republice po roce 1950. Zároveň je svou nadmořskou výškou 560 m naší nejvýše položenou lokaltu výskytu. Nález pravděpodobně představuje zálet (cca 80 km) na zimoviště z nejbližší známé letní populace v údolí Dunaje mezi Ingolstatdem a Regensburgem v sousedním Bavorsku. ACKNOWLEDGEMENTS We express our thanks to P. Cehláriková and J. Kadera from the Administration of the Český les Protected Landscape Area and H. Burda who kindly revised the manuscript. Special thanks go to M. Anděra (National Museum, Prague) and L. Dvořák (Šumava National Park) who took part in some field work. REFERENCES Anděra M. & Červený J., 1994: Atlas of the distribution of the mammals of the Šumava Mts. region (SW-Bohemia). Acta Sci. Natur. Brno, 28(2–3): 1–111. Bufka L., Bytel J., Hanzal V. & Vacík R., 2001: The distribution of bats (Chiroptera, Mammalia) in Western Bohemia: a rewiew. Folia Mus. Rer. Natur. Bohem. Occid., Zool., Plzeň, 41: 1–30. Červený J. & Bürger P., 1989: Density and structure of the bat community occupying and old park at Žihobce (Czechoslovakia). Pp.: 475–788. In: Hanák V., Horáček I. & Gaisler J. (eds.): European Bat Research 1987. Charles University Press, Praha, 718 pp. Červený J. & Bufka L., 1999: First records and long-distance migration of the Nathusius’s bat (Pipistrellus nathusii) in western Bohemia (Czech Republic). Lynx, n. s., 30: 121–122. Dvořák L., Bufka L. & Bytel J., 2003: Netopýři na zimovištích západních Čech v letech 1992–2003 a aktualizace jejich rozšíření [Bats on winter roosts in West Bohemia in 1992–2003 and their current distribution]. Erica, Plzeň, 11: 29–73 (in Czech, with a summary in English). Gaisler J., 1956: Faunistický přehled československých netopýrů [Faunistic rewiew of bats in the Czechoslovakia]. Ochr. Přír., 11(6): 161–169 (in Czech, with a summary in German). Gaisler J., 1997: Preliminary data on the distribution of Rhinolophidae in the Czech Republic and variation in numbers of R. hipposideros in S-Moravia. Pp.: 55–57. In: Ohlendorf B. (ed.): Tagungsband. Zur Situation der Hufeisennasen in Europa. IFA Verlag GmbH, Berlin, 184 pp. 77 Gaisler J., Hanák V., Hanzal V. & Jarský V., 2003: Výsledky kroužkování netopýrů v České republice a na Slovensku [Results of bat banding in the Czech and Slovak Republics, 1948–2000]. Vespertilio, 7: 3–61 in Czech, with a summary in English). Hanák V., 1962: Zpráva o nálezu vrápence velkého, Rhinolophus ferrumequinum (Schreber, 1774) a netopýra brvitého, Myotis emarginatus (Geofffroy, 1806) ve středních Čechách [Bericht über das Vorkommen der Grossen Hufeisennase, Rhinolophus ferrumequinum (Schreber 1774) und der Wimperfledermaus, Myotis emarginatus (Geoffroy 1806) in Mittelböhmen]. Lynx, n. s., 1: 21–22 (in Czech, with a summary in German). Hanák V. & Anděra V., 2005: Atlas rozšíření savců v České republice. Předběžná verze, V. Letouni (Chiroptera) – část 1. Vrápencovití (Rhinolophidae), netopýrovití (Vespertilionidae – Barbastella barbastellus, Plecotus auritus, Plecotus austriacus) [Atlas of the Mammals of the Czech Republic. A Provisional Version, V. Bats (Chiroptera) – Part 1. Horseshoe bats (Rhinolophidae), vespertilionid bats (Vespertilionidae – Barbastella barbastellus, Plecotus auritus, Plecotus austriacus)]. Národní museum, Praha, 120 pp (in Czech, with a summary in English). Hanák V. & Anděra V., 2006: Atlas rozšíření savců v České republice. Předběžná verze, V. Letouni (Chiroptera) – část 2. Rod Myotis [Atlas of the Mammals of the Czech Republic. A Provisional Ver sion, V. Bats (Chiroptera) – Part 2. Genus Myotis]. Národní museum, Praha, 187 pp (in Czech, with a summary in English). Hůrka L., 1971a: Nálezy zajímavějších druhů netopýrů v západních Čechách [Discoveries of some interesting species of bats in West Bohemia]. Vertebratol. Zpr., 1: 38–40 (in Czech, a summary in English). Hůrka L., 1971b: Zur Verbreitung und Ökologie der Fledermäuse der Gattung Plecotus (Mammalia, Chiroptera) in Westböhmen. Folia Mus. Rer. Natur. Bohem. Occid., Zool., Plzeň, 1: 1–24. Hůrka L., 1973: Výsledky kroužkování netopýrů v západních Čechách v letech 1959–1972 s poznámkami k jejich rozšíření, ekologii a ektoparazitům [The results of bat banding in Western Bohemia and notes to their distribution, ecology and ectoparasites]. Sborn. Západočes Muz. v Plzni, Přír., 9: 3–84 (in Czech, with a summary in German). Hůrka L., 1986: Doplněk k výskytu netopýrů v západních Čechách [The supplement to the bat occurrence in the Western Bohemia]. Zpr. Mus. Západočes. Kr. Plzeň, Přír., 32–33: 97–104 (in Czech, with a summary in German). Hůrka L., 1988: Zur Verbreitung und Bionomie des Mausohr, Myotis myotis (Mammalia, Chiroptera) in Westböhmen. Folia Mus. Rer. Natur. Bohem. Occid., Zool., Plzeň, 27: 35–55. Hůrka L., 1989: Die Säugetierfauna des westlichen Teils der Tschechischen Sozialistischen Republik. II. Die Fledermäuse (Chiroptera). Folia Mus. Rer. Natur. Bohem. Occid., Zool., Plzeň, 29: 1–61. Jahelková H., Lučan R. & Hanák V., 2000: Nové údaje o netopýru parkovém (Pipistrellus nathusii) v jižních Čechách [New data on the Nathusius’ pipistrelle (Pipistrellus nathusii) in South Bohemia]. Lynx, n. s., 31: 41–51 (in Czech, with a summary in English). Kůs E., 1999: Savci Přimdského lesa [Mammals of the Přimdský les region (Western Bohemia, Czech Republic)]. Lynx, n. s., 30: 77–100 (in Czech, with a summary in English). Merkel-Wallner G., Mühlbauer H.& Heller K.-G., 1987: Ein Wochenstubennachweis der Nordfledermaus, Eptesicus nilssoni (Keyserling et Blasius, 1839) in der Oberpfälz. Myotis, 25: 37–40. Meschede A. & Rudolph B.-U., 2004: Fledermäuse in Bayern. Bayerische Landesamt für Umweltschutz, Eugen Ulmer Verlag, Stutgart, 411 pp. Skiba R., 1987: Zur Vorkommen der Nordfledermaus, Eptesicus nilssoni (Keyserling et Blasius, 1839) im Südosten der Bundesrepublik Deutschland. Myotis, 25: 29–35. 78 Lynx (Praha), n. s., 37: 79–93 (2006). ISSN 0024–7774 Species composition, spatial distribution and population dynamics of bats hibernating in Wisłoujście Fortress, Polish Baltic Sea Coast (Chiroptera: Vespertilionidae) Skład gatunkowy, rozmieszczenie przestrzenne i dynamika populacji nietoperzy zimujących w Twierdzy Wisłoujście na polskim Pobrzeżu Bałtyku (Chiroptera: Vespertilionidae) Mateusz Ciechanowski1, Agnieszka Przesmycka1 & Konrad Sachanowicz2 1 Academic Chiropterological Circle of Polish Society for Nature Protection “Salamandra”, Department of Vertebrate Ecology and Zoology, University of Gdańsk, Legionów 9, PL–80-441 Gdańsk, Poland; matciech@kki.net.pl [MC]; aprzesmycka@o2.pl [AP] 2 Department of Animal Ecology, Nicolaus Copernicus University, Gagarina 9, PL–87-100 Toruń, Poland; chassan@poczta.onet.pl received on 8 June 2006 Abstract. The structure and dynamics of bat assemblage in winter and transitional period was studied in the Wisłoujście Fortress (16th–19th century) located in Gdańsk city, northern Poland. The object was used partially as a commercial store (until 1995), later as a summer tourist attraction, protected in winter from human penetration. Serious threats for existance of this monument forced recently the local officials to the intensive renovation project, leading to deterioration of roosting conditions for bats. In total, nine species were recorded: Myotis myotis, M. nattereri, M. mystacinus, M. brandtii (rare in Pomerania region), M. dasycneme (EN in Poland), M. daubentonii, Eptesicus serotinus, Pipistrellus nathusii (first winter record in Poland) and Plecotus auritus. M. daubentonii predominated among bats counted in autumn (43.4%) and netted in the entrances to the casemates (60%). M. nattereri was the most numerous species during winter (67.3%) and spring censuses (71.0%). M. dasycneme composed 3.5% of all bats counted inside of the Fortress (n=2256 records) but about 18% of individuals captured in mist nets (n=67). Only 8–12 bats were counted during winter 1993/1994. The number of bats increased significantly (r=0.86, p<0.02) until 2005, reaching the total number of 313 individuals in the whole object; however this trend was severely interrupted by restoration works. The adaptation of formerly unprotected roost in the Fortress area (installation of bat grill, walls of air-bricks etc.) compensated this decline only in a limited way. In winter 2002/2003 we studied seasonal dynamics in bats’ numbers. The highest number of M. daubentonii and M. dasycneme was recorded in September. The number of M. nattereri, after a peak in a half of September, declined almost to zero in October and increased again, reaching its maximum value in February. Natura 2000 site was established in the Wisłoujście Fortress, although its future remains uncertain. INTRODUCTION Several species of vespertilionid bats of the temperate zone spend winter in underground roosts maintaining optimal thermal conditions and humidity (Althringam 1996). Such roosts are mainly natural caves, widely distributed in mountain and upland areas. However, almost no shelters of that kind are available in the lowlands of Central Europe, thus many species need to hibernate 79 in their anthropogenic substitutes: cellars (Lesiński et al. 2004), wells (Bernard et al. 1998) and – among the most important – various military structures, often considered as historical monuments (Bogdanowicz 1983, Urbańczyk 1990, Bernard et al. 1991, Fuszara et al. 1996, Hebda & Nowak 2002, Sachanowicz & Zub 2002). The latter ones may serve as crucial sites for taxonomically rich bat assemblages, consisting of unusually large numbers of individuals (e.g. Urbańczyk 1990). Forts, castles, air-raid shelters, bunkers and tunnels of underground factories make an oportunity for long-therm monitoring of bat populations (Urbańczyk 1989), similar to that conducted in caves and abandoned mines of upland areas (for comparision see: Fig. 1. Plan of the Wisłoujście Fortress. Explanations: I – the guard room of the postern tunnel, II – The Południowo-Wschodni (South-Eastern) Bastion, III – The Ostroróg Bastion, IV – The Artyleryjski (Artille ry) Bastion, V – The Furta Wodna Bastion, VI – The Wreath (Arabic numbers refers to the controlled cellars), VII – the powder-magazine of The Eastern Entrenchment. Objects I–VI are parts of the Fort Carré. Rys. 1. Plan Twierdzy Wisłoujście. Objaśnienia: I – wartownia poterny, II – Bastion Południowo-Wschodni, III – Bastion Ostroróg, IV – Bastion Artyleryjski, V – Bastion Furta Wodna, VI –Wieniec (cyfry arabskie odnoszą się do kontrolowanych piwnic), VII – prochownia Szańca Wschodniego. Obiekty I–VI stanowią część Fortu Carré. 80 Kowalski & Lesiński 1991, Weinreich & Oude Voshaar 1992, Řehák & Gaisler 1999). Although selection of roosts by some bat species in military objects is well known recently (Bogdanowicz & Urbańczyk 1983, Lesiński 1986), the factors affecting their population dynamics remained unexplained, even if patterns of this dynamics were described in many localities (Bagrowska -Urbańczyk & Urbańczyk 1983, Fuszara & Kowalski 1995, Fuszara et al. 1996). Several constructions have stopped to serve for military purposes just in the recent times, they became utilized commercially, abandoned or demolished; restoration works are started in some of them in order to save their historical values, however causing serious threats for wintering bats (e.g. Mitchell-Jones 2004). In contrast, unguarded sites are often under a serious pressure of uncon trolled human penetration, intensifying spontaneous bat arousals (Thomas 1995) or even acts of vandalism (Mitchell-Jones 2004). Thus, intensity of human disturbance in bat hibernacula may vary in space and time, however no case studies were published to document any effects of this factor on bat numbers. The Polish Baltic Sea Coast is the area of special biogeographic significance. It is known not only as an important bat migratory path (Jarzembowski 2003), but also as a northern limit of distribution for some European species (Baagøe 2001, Güttinger et al. 2001, Bogdanowicz & Ruprecht 2004). However, the state of knowledge about distribution and size of winter bat colonies in that region remains insufficient, restricted only to a small portion of available underground constructions (Jarzembowski et al. 2000, Wojtaszyn et al. 2001, Dzięgielewska 2002). No detailed informations were published regardind bat hibernation in one of the most valuable monuments of military architecture – 16th-century Wisłoujście Fortress (Gdańsk City). Scattered data from transitional periods revealed significance of the object for some species recognized as endangered in Poland (Myotis dasycneme – Ciechanowski & Przesmycka 2001) or extremely rare in the region (Myotis brandtii – Ciechanowski & Sachanowicz 2003). Single winter records of only four species have been published until recently (Jarzembowski et al. 2000, Ciechanowski & Przesmycka 2001, Ciechanowski & Kokurewicz 2004). The aim of this study was to describe the structure of bat assemblage using the Wisłoujście Fortress during winter and transitional period, as well as long-term and seasonal changes in the number of bats roosting in the monumental military object. The additional goal of the following paper was to document the effect of temporally varying human disturbance on the largest bat hibernaculum of Polish Baltic Sea Cost. Study area and history of the site The Wisłoujście Fortress is located in the Gdańsk city, northern Poland (18° 40’ 43” E, 54° 23’ 39” N). Geographically, it lies in the eastern part of Baltic Sea Coast region – a border between Vistula Spit and Żuławy Wiślane (delta of the Vistula river). The main object – Fort Carré – was built in the 16th century, as a sea-fortress protecting the port of Gdańsk. It is composed of four Bastions: Południowo-Wschodni (South-Eastern), Ostroróg, Artyleryjski (Artillery) and Furta Wodna. They contain large casemates, built of bricks and covered by earth embankements. The lateral walls of the bastions are built of bricks and stones as well. The fortress was built almost on the sea level, thus neither the casamates nor most of the remaining bat hibernacula are the typical underground roosts, however the thick, brick walls made them well insulated, at least before the main renovation works in 2004–2005. Originally, the casemates were opened on the lateral sides, however the secondary walls were built in these openings in the 20th century, making the insides dark and partially free of frost; the effect was increased when wooden doors were installed in the main casemate gates. The remaining bat hibernacula of the Fort Carré are small cellars under the officers’ houses in so called Wreath and small guard-room in the fort’s postern. Outside of the central fort, the belt of external earthwork fortifications (The Eastern Entrenchment) was built in the 17th 81 century, enriched by the 19th century powder-magazine. The latter object, containing the narrow corridor with some ventilation shafts in the ceiling, maintain a significant bat hibernaculum as well. In February, the casemates and cellars were relatively cool (mean=3.7 °C, SD=2.2, range 0.6–9.0 °C, n=33) and moderately humid (mean=66.9%, SD=10.4, range 39.0–81.0%, n=19), although these figures may poorly reflect conditions inside of the crevices or ventilation shafts. Mean external temperature in the same period amounted 2.8 °C (SD=4.3, range –7.0–8.0 °C, n=8) (authors’ unpublished data). Both Fort Carré and The Eastern Entrenchment are surrounded by wide moats, filled by brackish waters from neighbouring Martwa Wisła – the dead branch of the Vistula River. Vegetation on The Eastern En trenchment consists of shrubs, as well as unmanaged tree stand of maples Acer platanoides, old, partially dying black poplars Populus nigra and lane of monumental horse-chestnuts Aesculus hippocastanus. The earthworks of Fort Carré were densly overgrown by Lycium halimifolium, removed completely in 2003. The moat banks are covered by scattered beds of Phragmites australis and Schoenoplectus tanbaermontani. The Wisłoujście Fortress was demilitarised after the First World War and heavily demolished in 1945. After a partial restoration it was used extensively as a store of paints, windows and vegetable preserves. The vertical ventilation shafts were closed by concrete plates or even fulfilled with soil and rubble what increased an insulation of internal parts of the casemates. The monument faced an increasing devastation as an effect of sea waves, penetration of walls by moat waters, precipitation of salt and air-polution by neighbouring “Siarkopol” company, which stored and processed large amounts of sulphur. Deep crevices appeared in the walls and ceilings of all casemates of Fort Carré, causing a serious threat to the buliding, although – both with loose fragments of plaster – maintaining crucial bat shelters during hibernation. After 1995, the Fortress stopped to be used as a commercial store. The monument, managed by The City of Gdańsk History Museum, became utilised only in summer, as a seasonal tourist attraction. The Fort Carré remained closed and guarded in winter, contrary to the powder magazine of the Eastern Entrenchment, facing uncontrolled human penetration and disturbance. The Fortress was included on the list 100 most endangered cultural monuments by World Monument Watch in 2000–2001; the situation of the site resulted in starting of wide-scale restoration projects. First renovation works were performed since September 2003, in the same period, when bats used to aggregate in winter roosts. They included archeological excavations, prospecting drillings and removal of earth em bankements in Bastion Artyleryjski, as well as temporary opening of ventilation shafts in all bastions (the latter was performed in October). In the same period the Fortress became recognized as an important bat site by Polish Ministry of Environment (established as Special Area of Protection PLH 220037 in Natura 2000 European network). All works were stopped during the winter 2003/2004, while the ventilation shafts were closed provisionally (with spreadsheets of plastic foil) after an intervention of environmental officials. Restoration started again in spring 2004, resulting in fulfillment of all crevices in internal walls of Bastion Artyleryjski with cement; in summer 2005 they became completely covered with desalination compress. The small room in its casemates was separated from their remaining parts with a provisional synthetic curtain (only in winter 2004/2005). Additionally, a secondary wall in South-Eastern Bastion was partially removed in 2005, resulting in appearance of ice-cover in one of the corridors. All works were stopped for the two last winter periods. As a partial compensation for negative results of renovation works, the powder-magazine of the Eastern Entrenchment was modified in order to increase its capacity as bat hibernaculum. Its entrance was closed with bat grills (Mitchell-Jones 2004), while walls of air bricks and water reservoir were built inside in summer 2005. Material and methods All four bastions of the Fort Carré and cellar no. 12 of the Wreath were checked every February between 2000 and 2006 (7 checks). In winter season 2002/2003 these objects were checked every month (since September until April), in order to reveal seasonal dynamics in the number of bats (9 checks). Additional objects (cellars no. 2 and 6, guard-room of the postern tunnel) were checked irregularly until 2005, when all potential bat hibernacula in the Fortress area were controlled for the first time. Fort Carré was controlled also 5 times in winter 1993/1994 (December–March), on 27 November 1999, three times in the autumn 82 2003 (29 October, 18 September, 19 December) and on 8 April 2006. The powder-magazine of the Eas tern Entrenchment has been checked during every visit since September 2003. In total, 28 controls of the Fortress were conducted between 1993 and 2006. During each visit all visible bats were counted in the controlled objects, including those hidden in deep crevices and ventilation shafts. Potential shelters were checked using torches and head-lamps. We avoided handling of bats, except for species needing a careful examination of dental features or taking some mea surments (Pipistrellus spp., Myotis mystacinus, Myotis brandtii). Five times (6 October 2000, 21 October 2000, 14 September 2002, 14 September 2003, 15 October 2006) a mist netting was performed at the entrances of bastions of the Fort Carré. Two or four ECOTONE mist nets were set up since dawn until midnight. Captured bats were determined to the species, sexed, aged (in the case of M. daubentonii – Richardson 1994), weighed and their forearms measured prior to release. Results In total, nine bat species were recorded in autumn-spring period: Large mouse-eared bat Myo tis myotis (Borkhausen, 1797), Natterer’s bat Myotis nattereri (Kuhl, 1817), Whiskered bat Myotis mystacinus (Kuhl, 1817), Brandt’s bat Myotis brandtii (Eversmann, 1845), Pond bat Myotis dasycneme (Boie, 1825), Daubenton’s bat Myotis daubentonii (Kuhl, 1817), serotine Eptesicus serotinus (Schreber, 1774), Nathusius’ pipistrelle Pipistrellus nathusii (Keyserling et Blasius, 1839) and brown long-eared bat Plecotus auritus (Linnaeus, 1758). Among them, M. nattereri was the most numerous, with a significant share of M. daubentonii (Table 1). Both species were found during almost every control since the beginning of the study. M. dasyc neme appeared to be the third most numerous species, amounting not more than 5.2% of all bats, but regularly encountered since its first winter observation on 11 February 2002. Single M. myotis appeared irregularly (10 February 2001, 17 April 2003) until 2004, when it became a permanent element of bat assemblage wintering in Fort Carré. The remaining species were noted only sporadically: P. auritus (29 October 2003, 19 December 2003, 24 February 2004, 18 February 2005), E. serotinus (21 November 2002, 21 January 2003, 18 February 2005), M. mystacinus (1 male, 21 November 2002: Bastion Ostroróg), M. brandtii (1 female, 17 April Table 1. The number and percentage of bat records from Wisłoujście Fortress. Autumn: 14 IX – 20 XII, winter: 21 XII – 20 III, spring: 21 III – 17 IV Tab. 1. Liczba stwierdzeń i udział procentowy gatunków nietoperzy w Twierdzy Wisłoujście. Jesień (au tumn): 14 IX – 20 XII, zima (winter): 21 XII – 20 III, wiosna (spring): 21 III – 17 IV species autumn n % winter n % spring n % n total % Myotis myotis Myotis nattereri Myotis mystacinus Myotis brandtii Myotis dasycneme Myotis daubentonii Eptesicus serotinus Pipistrellus nathusii Plecotus auritus indeterminate 2 225 1 – 31 260 1 – 1 78 0.3 37.6 0.2 0.0 5.2 43.4 0.2 0.0 0.2 13.0 10 927 – – 41 239 2 1 2 156 0.7 67.3 0.0 0.0 3.0 17.3 0.1 0.1 0.1 11.3 4 198 – 1 8 50 – – – 18 1.4 71.0 0.0 0.4 2.9 17.9 0.0 0.0 0.0 6.5 16 1350 1 1 80 549 3 1 3 252 0.7 59.8 <0.1 <0.1 3.5 24.3 0.1 <0.1 0.1 11.2 total 599 100.0 1378 100.0 279 100.0 2256 100.0 83 Table 2. The number and species composition of bats hibernating in Wisłoujście Fortress, counted on 18.02.2005. Explanations: I – the guard room of the postern tunnel, II – The Południowo–Wschodni (South-Eastern) Bastion, III – The Ostroróg Bastion, IV – The Artyleryjski (Artillery) Bastion, V – The Furta Wodna Bastion, VI – cellars of the Wreath, VII – the powder-magazine of The Eastern Entrenchment Tab. 2. Liczebność i skład gatunkowy nietoperzy zimujących w Twierdzy Wisłoujście dnia 18.02.2005. Ob jaśnienia: I – wartownia poterny, II – Bastion Południowo-Wschodni, III – Bastion Ostroróg, IV – Bastion Artyleryjski, V – Bastion Furta Wodna, VI – piwnice Wieńca, VII – prochownia Szańca Wschodniego species I II III IV V 12 VI 2 6 VII total % 1.3 78.9 3.5 9.6 0.3 0.3 6.1 M. myotis – M. nattereri 3 M. dasycneme – M. daubentonii – E. serotinus – P. auritus – indeterminate 2 53 10 11 1 1 4 – 8 – 2 – – 4 – 143 – 2 – – 3 – 13 – 4 – – 1 – 3 – – – – 1 – – 2 4 – – – – – – – – 1 2 18 1 11 – – 5 4 247 11 30 1 1 19 total number of species 82 6 14 2 148 2 18 2 4 1 3 1 37 4 313 100.0 6 – 3 1 4 1 2003: Bastion Artyleryjski) and P. nathusii (1 male, 16 February 2003: Bastion Furta Wodna). In total 2256 bat records were noted (Table 1) and 313 bats were counted maximally in the whole Fortress (Table 2). The species composition varied in time. In autumn, M. daubentonii predominated, but in winter and spring M. nattereri was a dominating taxon. The proportion of M. daubentonii compared to M. nattereri was significantly higher in autumn than in winter (Chi-square test with Yates’ correction, χ2=176.50, p<0.0001), in autumn than in spring (χ2=73.85, p<0.0001), but did not differ between winter and spring (χ2=0.00, p>0.9). The total number of hibernating bats in the Fortress has changed significantly since 1994. After closing of the commercial stores it began to increase progressively (r=0.86, p<0.02; y= –41415.7626 + 20.7553957x) until 2005, when it reach the maximum. This trend was interrupted only in winter 2003/2004, when first renovation works, excavations and prospecting drillings were performed in autumn. However, a year later, the number of bats roosting in the Fort Carré dropped about 77% in association with removal of secondary wall in South-Eastern Bastion and covering of the walls with desalination compress in Artillery Bastion. Although the number of bats counted in newly adapted powder-magazine of the Eastern Entrenchment increased almost twice, it did not compensate a population decline in the remaining objects. Hibernating bats were unevenly distributed in the Fortress area, with 85% of the assemblage concentrated in just three objects. The highest species richness was observed in South-Eastern Bastion and powder-magazine of the Eastern Entrenchment, while the highest number of indivi duals appeared in the Artillery Bastion. Only a few bats were noted in the remaining six objects (Table 2). With respect to the species composition, the first two objects appeared to be the most vaulable bat hibernacula, where most individuals of M. dasycneme spend winter and mate in autumn (observations of copulating pond bats in 2002). Progressing restoration works strongly affected the spatial distribution of bats. After their first stage in 2003, performed in the autumn, bats number decreased more than one third in the Artillery Bastion, but increased in the South84 -Eastern Bastion (Fig. 3). However, the fulfillment of crevices in the Artillery Bastion did not cause any decline of bats number in the next winter season (2005). Moreover, they aggregated almost all in one place – a small chamber, separated by a plastic curtain, where three, large clusters of M. nattereri (59, 51 and 17 individuals) hanged directly on its walls and composing a majority (88%) of all bats wintering in this bastion. Such a behaviour was formerly unknown in the Fortress, where most of the animals hibernated hidden in ceiling cracks, dispersed among several, smaller clusters. Only after the second stage of restoration works, bats almost completely abandoned the Artillery Bastion, where not only crevices were filled, but also the temporary curtain was removed (Fig. 4). The number of bats varied strongly through the season as well (Fig. 5). The largest aggregati ons of M. daubentonii appeared in early September. Contrary to the winter period (see above), they did not hide in the crevices, but covered the ceilings and walls of ventilation shafts in dense clusters. The number of M. daubentonii declined visibly in the third decade of September, later increasing again, but not reaching the earlier level. It remained relatively stable between October Fig. 2. Long-therm changes in the number of bats hibernating in the Wisłoujście Fortress in 1994–2006, based on February counts. Explanations: 1. Autumn works in Artillery Bastion (prospecting drillings, archeological excavations, opening of ventilation shafts, removal of earth embankements); 2. Summer renovation works (fulfillment of crevices and laying of desalination compress on walls in Artillery Bastion, partial removal of secondary wall in South-Eastern Bastion). Rys. 2. Długoterminowe zmiany liczebności nietoperzy zimujących w Twierdzy Wisłoujście (lata 1994– 2006), w oparciu o liczenia w lutym. Objaśnienia: 1. Jesienne prace w Bastionie Artyleryjskim (wiercenia rozpoznawcze, wykopaliska archeologiczne, udrożnienie przewodów wentylacyjnych, usunięcie nasypów ziemnych znad sklepienia); 2. Letnie prace remontowe (wypełnienie szczelin i pokrycie kompresem odsalającym wewnętrznych ścian Bastionu Artyleryjskiego, częściowe usunięcie wtórnego zamurowania w Bastionie Południowo-Wschodnim). 85 Fig. 3. Effect of works conducted in Artillery Bastion in autumn 2003 on bats number in particular parts of Fort Carré, expressed as a differrence in bat number between December 2002 and 2003. Rys. 3. Wpływ prac prowadzonych w Bastionie Artyleryjskim jesienią 2003 na liczebność nietoperzy w poszczególnych częściach Fortu Carré, wyrażony jako różnica w liczbie osobników między grudniem 2002 i 2003. Fig. 4. Effect of works conducted in Artillery and South-Eastern Bastions in 2005 on bats number in particular parts of Wisłoujście Fortress, expressed as a differrence in bat number between February 2005 and 2006. Rys. 4. Wpływ prac prowadzonych w Bastionach Artyleryjskim i Południowo-Wschodnim w roku 2005 na liczebność nietoperzy w poszczególnych częściach Twierdzy Wisłoujście, wyrażony jako różnica w liczbie osobników między lutym 2005 i 2006. 86 Table 3. The species and sex composition of bats netted in autumn in the entrances of casemates and pow der-magazine of the Wisłoujście Fortress Tab. 3. Skład gatunkowy i struktura płciowa nietoperzy odławianych w sieci jesienią przy wylotach z ka zamat i prochowni Twierdzy Wisłoujście species males females total % Myotis myotis Myotis nattereri Myotis mystacinus Myotis dasycneme Myotis daubentonii 1 8 1 10 24 2 3 – 2 16 3 11 1 12 40 4 16 1 18 60 total 43 22 67 100 and February, decreasing finally in spring. A similar seasonal dynamics was revealed by M. dasycneme, however basing on a very small sample (n=39 records). The number of M. nattereri – after an early peak in the beginning of season – dropped almost to zero in October. However, it increased about 60 times until January, exceeding the number of M. daubentonii already in the late autumn. The highest number of M. nattereri was recorded in March (declining almost six times in the next month), but the highest total number of bats – in February. Fig. 5. Seasonal dynamics in the number of bats roosting in Wisłoujście Fortress in winter 2002/2003. Rys. 5. Sezonowa dynamika liczebności nietoperzy wykorzystujących Twierdzę Wisłoujście jako kry jówkę zimą 2002/2003. 87 Among bats netted in the autumn (n=67), M. daubentonii predominated, but the high per centage of M. dasycneme (18%) was noticeable (Table 3). Most captured individuals were males, however a sex ratio did not differ significantly from 1:1 (χ2=2.74, p>0.09). Among M. daubentonii 31 juveniles (yearlings) and 9 adults were captured; difference from 1:1 age ratio was statistically significant (χ2=5.41, p=0.02). When the mist netting was conducted, several individuals of small Myotis were observed flying inside of casemates. Discussion The number of bats roosting in the Wisłoujście Fortress during winter period placed it among the largest bat hibernacula in the Polish Baltic Sea Coast, at least in some seasons. Only one site – ruins of amunition factory in Police near Szczecin – appears larger, accumulating more than 1300 hibernating individuals (K. Drabińska, J. Żejmo, M. Dzięgielewska in Kepel et al. 2005). About 200 individuals were counted maximally in the castle of Teutonic Knights in Malbork (Stec & Kasprzyk in Kepel et al. 2005) and 120–150 individuals in bunkers of Kołobrzeg-Stadion and Szczecin-Zdroje (Wojtaszyn et al. 2001, Dzięgielewska 2002). No other bat hibernaculum located in Gdańsk city has ever been used by more than 90 animals (Jarzembowski et al. 2000). Among the species recorded in Wisłoujście, some of them are of special significance, even if only exceptionaly encountered. M. brandtii, although considered as widely distributed in Po land, is extremely rare in the northern part of the country. It is only the second known locality of this species in the Baltic Sea Coast region (Ciechanowski & Sachanowicz 2003). P. nathusii is a long-distance migrant, known to leave Central Europe for winter and hibernating in hollow trees or houses located in milder climate of western Europe (Strelkov 1969). The Wisłoujście Fortress is the first known winter site of P. nathusii in Poland and quite an unusual type of hibernaculum, although the recent winter record of the species from Czech Republic is also associated with a similar object (a castle cellar, Benda & Hotový 2004). Among the regular inhabitants of the Fortress, M. dasycneme should be considered as the most interesting species. Regarded as globally vulnerable (Hutson et al. 2001) and nationally endangered (Wołoszyn 2001), it is included in Annex II of EU Habitat Directive, justifying designation of Natura 2000 site in Wisłoujście Fortress. Polish hibernacula do not concentrate large numbers of M. dasycneme. Its share in winter bat assemblages usually do not exceed 0.5% (Bagrowska-Ur bańczyk & Urbańczyk 1983, Lesiński 1986, Kowalski & Lesiński 1991, Fuszara et al. 1996) and even in some large brick forts no pond bats are encountered (Bogdanowicz 1983, Fuszara & Kowalski 1995, Hebda & Nowak 2002). Wisłoujście Fortress is one of the largest winter sites of M. dasycneme in Poland, along with Fort Osowiec in Biebrza River Valley (at maximum 34 individuals – Lesiński & Kowalski 2002) and four other localities (including three fortification complexes), each concentrating no more than 6–15 individuals (Ciechanowski & Kokurewicz 2004, T. Kokurewicz in litt.). In fact, the real number of pond bats roosting in the Fortress can be even higher, what may be presumed from regular captures of M. dasycneme in autumn and observations of some individuals hiding deeply in narrow crevices. Bogdanowicz (1983) stated that M. dasycneme avoided winter quarters located in heavily urbanized areas, thus being a bioindicator of the changes in a habitat subjected to urbanization pressure. This hypothesis can recently be rejected, as the Wisłoujście Fortress – intensively used by this species both in autumn and winter – is located on the edge of densely populated city, where two large chemical factories have polluted the neighbouring area since 1969. 88 The species composition of bats wintering in Wisłoujście Fortress differed from that recorded in the other large, brick forts of Poland. In central and northeastern part of the country, forts are inhabited mainly by a barbastelle Barbastella barbastellus (Schreber, 1774) composing 50.8% of all bats (Fuszara et al. 1996). Also in the forts of Nysa (Lower Silesia) barbastelle is the most numerous species (Hebda & Nowak 2002), while in the forts of Poznań it composed at least 14.1–20.9% of winter bat colonies (Bogdanowicz 1983). Barbastelle is an oligothermophilous species, hibernating in 0.0–3.0 °C (Bogdanowicz & Urbańczyk 1983) and large forts are considered rather cool roosts with intensive air circulation (due to many entrances and windows), what explains high numbers of B. barbastellus spending winters there. Thus, the lack of any barbastelles hibernating in Wisłoujście Fortress seems astonishing, however the species does not occur anywhere in Gdańsk (Jarzembowski et al. 2000) and it is a very rare bat in the whole Pomerania region (Kowalski & Szkudlarek 2003). Instead, the winter bat assemblage of the studied locality is dominated by moderately thermophilous species, i.e. M. nattereri and M. daubentonii. However, they are known to find shelters in the deep, highly located crevices and ventilation shafts (Bogdanowicz & Urbańczyk 1983, Lesiński 1986), where the temperature may be significantly higher and more stable than in the corridors themsleves. Low number of bats in the period, when object was intensively used as a commercial store, confirms their sensibility to repetitive disturbance (light, noise) during hibernation (Thomas 1995). Later increase in bat numbers appeared similar to that observed in roosts newly opened for these animals, e.g. the old mines, where exploitation came to an end (Lutsar et al. 2000, Bihari 1998 in Güttinger et al. 2001). Also in winter quarters, where bats were intensively ban ded, their numbers increased several times after the end of marking programm (Lesiński 1990, Řehák & Gaisler 1999). Spatio-temporal dynamics of bat numbers in the Wisłoujście Fortress suggest however, that human pressure restricted only to the part of fortification complex forced at least some bats to change roost for the nearest available, not to leave completely the area. Movements among hibernacula in one and the same winter season (apparently associated with flying outside) were reported during some banding programmes (Bogdanowicz & Urbańczyk 1983, Gaisler et al. 2003). Bats may change their shelter not only directly due to the human activity, but its subsequent effects on internal conditions. At least some elements of restoration works (removal of secondary wall, opening of ventilation shafts) might strongly affect microc limate in the casemates, causing bats not only hiding deeper in the crevices (reaction observed frequently in M. daubentonii when ambient temperature decreased – Kokurewicz 2004), but also moving into unaffected parts of the Fortress. The latter behaviour can be compared with that recorded in B. barbastellus wintering in small, concrete bunkers and regularly moving from the upper floor to the dried wells in the coldest months (Sachanowicz & Zub 2002). Our observations showed however, that alternative roosts could only partially compensate the effect of human pressure on hibernaculum, likely due to unpredictable character of this factor. The phenology of bat occurrence in Fortress resembles that observed in the other fortificati ons of lowland Poland. M. daubentonii is a bat appearing earlier in the undergrounds that M. nattereri and the highest numbers of this species are observed in the autumn, decreasing later during winter months (Bagrowska-Urbańczyk & Urbańczyk 1983, Jurczyszyn 1998). However, the mentioned peak in the number of Daubenton’s bats was reported a bit later – from October (Lesiński 1986, Fuszara & Kowalski 1995, Fuszara et al. 1996) or even from November (Jur czyszyn 1998). Thus, the mass occurrence of M. daubentonii in the Fortress in September is worth noticing, possibly representing a short-therm phenomenon shifting in time among years – we observed it on 14 September 2002 but not on 18 September 2003. The species composition 89 of bats netted in autumn resembled that recorded inside of the casemates, although it may not represent the latter ones (they remained in a deep torpor) but the animals visiting the Fortress in order to take part in a swarming behaviour (sensu Parsons et al. 2003). Several individuals of M. daubentonii visited the Modlin Fortress near Warsaw even in August, however they number dropped almost to zero in the first week of September and increased again. Most of the individuals appearing then in Modlin remained active, flying inside of the forts, and only 4% of bats ringed in the autumn (including those found in lethargy) stayed there in winter (Lesiński 1990). Thus it was possible that high numbers of M. daubentonii roosting in the Fortress in September did not refer to the animals counted later in mid-winter but to the bats using the site as a transitional shelter during autumn movements. Contrary to M. daubentonii, the maximum numbers of M. nattereri in hibernacula are usually observed in mid-winter, mostly in January–February (Bagrowska-Urbańczyk & Urbańczyk 1983, Fuszara et al. 1996), sometimes in March (Fuszara & Kowalski 1995) or December (Jurczyszyn 1998). The earlier results are, thus, consistent with our observations, however an early peak in the number of Natterer’s bats, revealed in the half of September, was recorded for the first time. These animals could also use the Fortress as a transitional roost, leaving it after a short time, what may be judged from the very low numbers of M. nattereri in October, confirmed also in the other sites (Lesiński 1986, Fuszara et al. 1996, Jurczyszyn 1998). There are almost no earlier studies to compare with our data on phenology of M. dasycneme. The two series of censuses conducted by Lesiński & Kowalski (2002) in northeastern Poland revealed the numbers of pond bats at least 2–10 times higher in November–December than in February. No such phenomenon was observed in Wisłoujście and the September peak in the number of M. dasycneme seems to reveal the same aspect of annual life cycle as in M. nattereri and M. daubentonii. It should be noted however, that some changes in the number of bats counted in underground hibernacula could be an artifact, unrelated to arrival or departure from roosts, but caused by movements of some individuals into the deep crevices (Jurczyszyn 1998). This effect in M. daubentonii was explained as a reaction to unfavorable thermal conditions (Kokurewicz 2004). The future of the Wisłoujście Fortress as an important bat site remains uncertain. It would be hard to preserve such a large winter colony in the casemates of Fort Carré, because the prospe cting restoration works – necessary to preserve the Fortress itself – will change the microcli mate, as well as the number and quality of micro-shelters. Adaptation of the powder-magazine on Eastern Entrenchment may compensate these losses only in a limited way and there would be a need for a significant delay (about 2–4 years) in renovation of the South-Eastern Bastion, until more bats accept a new roost. The main purpose for the creation of Natura 2000 site on the Fortress’ area – regular occurrence of M. dasycneme – seems possible to be maintained, as the powder-magazine became recently the most important shelter of this species (5 of 7 individuals were noticed there on 8 February 2006). Streszczenie W latach 1994–2006 badaliśmy strukturę i dynamikę zgrupowania nietoperzy wykorzystujących Twierdzę Wisłoujście (Gdańsk, północna Polska) zimą i w okresach przejściowych. Obiekt, budowany w XVI–XIX wieku, do 1995 roku był częściowo użytkowany jako magazyny, w ostatnich latach zaś jako letnia atrakcja turystyczna, zabezpieczona przed ludzką penetracją w okresie zimowym. Poważne zagrożenia dla substancji zabytkowej Twierdzy, skłonily lokalne władze do intensywnej renowacji obiektu, co prowadzi jednak do obniżenia jego wartości jako kryjówki nietoperzy. 90 Łącznie na terenie Twierdzy stwierdzono 9 gatunków nietoperzy: nocek duży Myotis myotis, nocek Natterera M. nattereri, nocek wąsatek M. mystacinus, nocek Brandta M. brandtii (rzadki na Pomorzu), nocek łydkowłosy M. dasycneme (kategoria EN w Polsce), nocek rudy M. daubentonii, mroczek późny Eptesicus serotinus, karlik większy Pipistrellus nathusii (pierwsze w Polsce stwierdzenie zimowe) i gacek brunatny Plecotus auritus. Nocek rudy dominował wśród nietoperzy liczonych jesienią wewnątrz Twierd zy (43,4%) i odławianych w sieci przy wejściach do kazamat (60%). Nocek Natterera był najliczniejszym gatunkiem podczas liczeń zimowych (67,3%) i jesiennych (71,0%). Nocek łydkowłosy stanowił 3,5% wszystkich nietoperzy liczonych wewnątrz Twierdzy (n=2256 stwierdzeń), ale około 18% osobników odławianych w sieci (n=67). Zimą 1993/1994 na terenie Twierdzy znajdowano zaledwie 8–12 nietoperzy. Łączna liczba tych zwierząt istotnie wzrastała do 2005 roku (r=0,86, p<0,02), osiągając wartość 313 osobników w całym obiekcie; jednak trend ten został przerwany na skutek prac renowacyjnych. Adaptacja wcześniej niechronionej Prochowni Szańca Wschodniego (instalacja kraty zabezpieczającej, ścian z cegły-dziurawki) tylko częś ciowo skompensowała spadek liczebności nietoperzy między latami 2005 i 2006. Zimą 2002/2003 badaliśmy sezonową dynamikę liczebności nietoperzy. Najwięcej nocków rudych i nocków łydkowłosych odnotowaliśmy we wrześniu. Liczba nocków Natterera, po osiągnięciu szczytu w połowie września, spadła niemal do zera w październiku, po czym ponownie wzrosła, osiągając mak symalną wartość w lutym. Twierdza Wisłoujście uznana została za Specjalny Obszar Ochrony w sieci Natura 2000, jednak jego przyszłość, jako jednego z największych zimowisk nietoperzy na Pomorzu Gdańskim, pozostaje niepewna. Acknowledgements Authors are cordially thankful to all who have contributed to the following study and conservation of bats in the Wisłoujście Fortress. Members of The City of Gdańsk History Museum staff: Adam Koperkiewicz, Halina Trzebiatowska and Zdzisław Balewski for making access to the study area possible and for their decision to adapt a powder-magazine for bat hibernaculum. Members and co-workers of Academic Chiro pterological Circle of PTOP “Salamandra” in Gdańsk: Urszula Anikowska, Anna Biała, Piotr Chybowski, Tomasz Jarzembowski, Monika Kulas, Anna Miotk, Anna Nalewaja, Justyna Naumowicz, Anna Pawlik, Katarzyna Pazio, Paulina Piasecka, Weronika Rogalska, Aleksandra Rynkiewicz, Katarzyna Wojczulanis, Tomasz Zając and Aneta Zapart for their help in the field studies. References Altringham J. D., 1996: Bats, Biology and Behaviour. Oxford University Press, Oxford, 262 pp. Baagøe H. J., 2001: Eptesicus serotinus (Schreber, 1772) – Breitflügelfledermaus. Pp.: 519–559. In: Krapp F. (ed.): Handbuch der Säugetiere Europas. Band 4: Fledertiere. Teil I. Chiroptera I. Rhinolophidae, Vespertilionidae 1. AULA-Verlag GmbH, Wiebelsheim, x+603 pp. Bagrowska-Urbańczyk E. & Urbańczyk Z., 1983: Structure and dynamics of a winter colony of bats. Acta Theriol., 28: 183–196; Benda P. & Hotový J., 2004: Nález zimujícího netopýra parkového (Pipistrellus nathusii) na jižní Moravě [Hibernation record of Pipistrellus nathusii in southern Moravia (Czech Republic)]. Vespertilio, 8: 137–139 (in Czech, with an abstract in English). Bernard R., Głazaczow A. & Samoląg J., 1991: Overwintering bat colony in Strzaliny (North-Western Poland). Acta Zool. Cracov., 34: 453–461. Bernard R., Gawlak A. & Kepel A., 1998: The importance of village wells for hibernating bats on the example of a village in North-western Poland. Myotis, 36: 25–30. Bihari Z., 1998: Examination of the settlement of Myotis myotis in the abandoned mine. Myotis, 36: 225–228. Bogdanowicz W., 1983: Community structure and interspecific interactions an bats hibernating in Poznań. Acta Theriol., 28: 357–370. 91 Bogdanowicz W. & Ruprecht A. L., 2004: Nyctalus leisleri (Kuhl, 1817) – Kleinabendsegler. Pp.: 717–756. In: Krapp F. (ed.): Handbuch der Säugetiere Europas. Band 4: Fledertiere. Teil II. Chiroptera II. Ve spertilionidae 2, Molossidae, Nycteridae. AULA-Verlag GmbH, Wiebelsheim, x+[605–1186] pp. Bogdanowicz W. & Urbańczyk Z., 1983: Some ecological aspects of bats hibernating in city of Poznań. Acta Theriol., 28: 371–385. Ciechanowski M. & Kokurewicz T. 2004: Myotis dasycneme (Boie, 1825). Nocek łydkowłosy [Myotis dasycneme (Boie, 1825). Pond bat]. Pp.: 368–373. In: Adamski P., Bartel R., Bereszyński A., Kepel A. & Witkowski Z. (eds): Poradniki ochrony siedlisk i gatunków Natura 2000 – podręcznik metodyczny. Tom 6. Gatunki Zwierząt (z wyjątkiem ptaków) [Conservation Handbooks of Natura 2000 habitats and species. Vol. 6. Animal species (except of birds)]. Ministerstwo Środowiska, Warszawa, 500 pp (in Polish). Ciechanowski M. & Przesmycka A., 2001: Stwierdzenie nocka łydkowłosego Myotis dasycneme (Boie, 1825) i nocka wąsatka Myotis mystacinus (Kuhl, 1817) w Gdańsku [Record of a pond bat Myotis da sycneme (Boie, 1825) and whiskered bat Myotis mystacinus (Kuhl, 1817) in Gdańsk]. Nietoperze, 2: 69–73 (in Polish, with a summary in English). Ciechanowski M. & Sachanowicz K., 2003: Pierwsze stwierdzenie nocka Brandta Myotis brandtii (Ever smann, 1845) we wschodniej części polskiego Pobrzeża Bałtyku [First record of Brandt’s bat Myotis brandtii (Eversmann, 1845) in the eastern part of Polish Baltic Sea Coast]. Nietoperze, 4: 178–179 (in Polish, with a summary in English). Dzięgielewska M., 2002: Zimowe liczenia nietoperzy w Szczecinie w latach 1996–1999 [Winter counts of bats in Szczecin in 1986–1999]. Nietoperze, 3: 7–15 (in Polish, with a summary in English). Fuszara E. & Kowalski M., 1995: Bats in underground shelters of Warsaw. Nyctalus (N. F.), 5(6): 545–555. Fuszara E., Kowalski M., Lesiński G. & Cygan J. P., 1996: Hibernation of bats in underground shelters of central and northeastern Poland. Bonn. Zool. Beitr., 46: 349–358. Gaisler J., Hanák V., Hanzal V. & Jarský V., 2003: Výsledky kroužováni netopýrů v České republice a na Slovensku, 1948–2000 [Rusults of bat banding in the Czech and Slovak Republics, 1948–2000]. Vespertilio, 7: 3–61 (in Czech, with a summary in English). Güttinger R., Zahn A., Krapp F. & Schober W., 2001: Myotis myotis (Borkhausen, 1797) – Grosses Mausohr, Grossmausohr. Pp.: 123–207. In: Krapp F. (ed.): Handbuch der Säugetiere Europas. Band 4: Fledertiere. Teil I. Chiroptera I. Rhinolophidae, Vespertilionidae 1. AULA-Verlag GmbH, Wiebel sheim, x+603 pp. Hebda G. & Nowak A., 2002: Winter colonies of bats in old fortifications in Nysa (SW Poland). Przyr. Sudetów Zach., Suppl., 2: 39–48. Hutson A. M., Mickleburgh S. P. & Racey P. A., 2001: Microchiropteran Bats. Global Status Survey and Conservation Action Plan. IUCN/SSC Chiroptera Specialist Group. Gland, Switzerland and Cam bridge, UK, 258 pp. Jarzembowski T., 2003: Migration of the Nathusius’ pipistrelle Pipistrellus nathusii (Vespertilionidae) along the Vistula Split. Acta Theriol., 48: 301–308. Jarzembowski T., Ciechanowski M. & Przesmycka A., 2000: Zimowanie nietoperzy na Pomorzu Gdańskim w latach 1989–1999 [Bats hibernating at the Gdańsk Pomerania in years 1989–1999]. Studia Chirop terol., 1: 29–42 (in Polish, with a summary in English). Jurczyszyn M., 1998: The dynamics of Myotis nattereri and M. daubentonii (Chiroptera) observed during hibernation season as an artefact in some type of hibernacula. Myotis 36: 85–91. Kepel A., Kowalski M., Dzięciołowski R. & Lesiński G., 2005: The Agreement on the Conservation of Population of European Bats (Eurobats). Report on the implementation of the Agreement in Poland 2003–2004. Inf.EUROBATS.AC10.23. http://www.eurobats.org/documents/pdf/National_Reports/ nat_rep_Pol_2005.pdf Kokurewicz T., 2004: Sex and age related habitat selection and mass dynamics of Daubenton’s bats Myotis daubentonii (Kuhl, 1817) hibernating in natural conditions. Acta Chiropterol., 6: 121–144. 92 Kowalski M. & Lesiński G., 1991: Changes in numbers of bats in Szachownica Cave (Central Poland) during 10 years. Myotis, 29: 35–38. Kowalski M. & Szkudlarek R., 2003: Distribution of Barbastella barbastellus in Poland in the years 1980–1998. Nyctalus (N. F.), 8: 599–602. Lesiński G., 1986: Ecology of bats hibernating underground in central Poland. Acta Theriol., 31: 507–521. Lesiński G., 1990: Changes in numbers of Myotis daubentoni (Kuhl, 1819) in autumn shelters and the effect of disturbance. Acta Theriol. 35: 364–368; Lesiński G. & Kowalski M., 2002: Zimowy monitoring nietoperzy w Dolinie Narwi i Biebrzy w latach 1992–1999 [Winter bat censuses in the Narew and Biebrza valleys in 1992–1999]. Nietoperze, 3: 53–60 (in Polish, with a summary in English). Lesiński M., Kowalski M., Domański J., Dzięciołowski R., Laskowska-Dzięciołowska K. & Dzięgielewska M., 2004: The importance of small cellars to bat hibernation in Poland. Mammalia, 68: 345–352. Lutsar L., Masing M. & Poots L., 2000: Changes in the numbers of hibernating bats in the caves of Piusa (Estonia), 1949–1999. Folia Theriol. Eston., 5: 111–117. Mitchell-Jones A. J., 2004: Conserving and creating bat roosts. Pp.: 111–134. In: Mitchell-Jones A. J. & McLeish A. P. (eds): Bat Workers’ Manual. Joint Nature Conservation Comitee, Peterborough, 178 pp. Parsons K. N., Jones G., Davidson-Watts I. F & Greenway F., 2003: Swarming of bats at underground sites in Britain – implications for conservation. Biol. Conserv., 111: 63–70. Řehák Z. & Gaisler J., 1999: Long-term changes in the number of bats in the largest man-made hiberna culum of the Czech Republic. Acta Chiropterol., 1: 113–123 Richardson P. W., 1994: A new method of distinguishing Daubenton’s bats (Myotis daubentonii) up to one year old from adults. J. Zool., London, 233: 307–344. Sachanowicz K. & Zub K., 2002: Numbers of hibernating Barbastella barbastellus (Schreber, 1774) (Chi roptera, Vespertilionidae) and thermal conditions in military bunkers. Mammal. Biol., 67: 179–184. Strelkov P. P., 1969: Migratory and stationary bats (Chiroptera) of the European part of the Soviet Union. Acta Zool. Cracov., 14: 369–439. Thomas D. W., 1995: Hibernating bats are sensitive to non-tactile human disturbance. J. Mammal., 76: 940–946. Urbańczyk Z., 1989: Changes in the population size of bats in the “Nietoperek” Bat Reserve in 1975–1987 (Preliminary report). Pp.: 507–510. In: Hanák V., Horáček I. & Gaisler J. (eds): European Bat Research 1987. Charles University Press, Praha, 718 pp. Urbańczyk Z., 1990: Northern Europe’s most important bat hibernation site. Oryx, 24: 30–34. Weinreich J. H. & Oude Voshaar J., 1992: Population trends hibernating in marl caves in the Netherlands. Myotis, 30: 75–83. Wojtaszyn G., Stephan W. & Rutkowski T., 2001: Nietoperze zimujące w schronach w Kołobrzegu (1998–2001) [Bats hibernating at shelters in Kołobrzeg (1998–2000)]. Studia Chiropterol., 2: 75–79 (in Polish, with a summary in English). Wołoszyn B. W., 2001: Myotis dasycneme (Boie, 1825). Nocek łydkowłosy [Myotis dasycneme (Boie, 1825). Pond bat]. Pp.: 51–52. In: Głowaciński Z. (ed.). Polska czerwona księga zwierząt. Kręgowce. [Polish Red Data Book of Animals. Vertebrates]. Państwowe Wydawnictwo Rolnicze i Leśne, Warsza wa, 449 pp (in Polish). 93 Lynx (Praha), n. s., 37: 95–105 (2006). ISSN 0024–7774 The Early Miocene mammalian assemblages in Jebel Zelten, Libya Spodnomiocénní společenstva savců z Džebelu Zelten, Libye Oldřich Fejfar1 & Ivan Horáček2 Department of Geology and Palaeontology, Charles University, Albertov 6, CZ–128 43 Praha 2, Czech Republic; fejfar@natur.cuni.cz 2 Department of Zoology, Charles University, Viničná 7, CZ–128 44 Praha 2, Czech Republic; horacek@natur.cuni.cz 1 received on 8 December 2006 Abstract. The small mammal assemblage obtained from six sites of the Miocene deposits in Jebel Zelten, Libya, in 1982, 1983 and 1997, is composed of 10 species of rodents (seven different clades), 2 spp. of ochotonid Lagomorphs and one vespertilionid bat. A special attention is given to the forms characterising the uppermost part of the series: they belong (mostly as new species) to the genera Potwarmus (Muridae), Mellalomys (Myocricetodontinae), Heterosminthus (Lophiocricetinae), Sayimys (Ctenodactylidae) and a new genus of Vespertilionidae. The most parsimonious dating for them is the early Middle Miocene (ca. 15 My) as the cricetids and ctenodactylids are apparently less advanced than their congeneric forms in Beni Mellal (14 My) while Potwarmus, representing the first wave of murid expansion from Asia to Africa, is more advanced than the congeneric forms from Banda daud Shah in Pakistan (16 My). All these forms reveal close relations both to the clades known from several other African localities and to those found in the Early or Middle Miocene of Pakistan and illustrate the intesive s. c. “southern” faunal interchange between north Africa and Asia in time of the Early and Middle Miocene. Introduction Small mammal remains from seven localities were collected during two geological and paleontological field campaigns (1982, 1983 and 1997). The assemblages are small, but the twelve species recognized and described represent seven rodent families, one lagomorph and one bat family. The Jebel Zelten large mammal fauna was considered in most literature to represent one time-slice, although the interpretation of its age has been diverse. However, on basis of the evolutionary stage of the small mammal species, the faunal compositions and the stratigraphic sequence the Jebel Zelten assemblages represent three periods in time and cover approximately 4 million years. Three assemblages can be assigned to the Middle Early Miocene (18–19 Ma), one to the Late Early Miocene (16–17) and two to the Middle Miocene (14–15 Ma) (Wessels et al. 2003). From sites with a high concentration of vertebrate remains at the surface, the fine cross-bedded (estuarine – fluviatile) sands were extensively dry sieved by O. Fejfar in 1982 and 1983. On two sites, the “Measured Section 2” (MS 2) in the middle part of the escarpment and Wadi Umm Ash Shatirat (WS) in the most southern part of the escarpment (Fig.1), isolated molars of several taxa of rodents were collected. Site MS 2 corresponds with the vertebrate site “H – area 6409” and the site of Wadi Shatirat (WS) corresponds with the Vertebrate site “LP – areas 641295 16” of Savage & Hamilton (1973). Each assemblage is derived from a different stratigraphical level, e.g., MS2 belongs to a stratigraphically lower level than Wadi Shatirat. A geological and paleontological campaign El Arnauti-Daams in 1997, resulted both in collection of large mammals and further small collection of rodents and lagomorphs. In many localities of the Jebel Zelten escarpment three to four fossilliferous units of sandstones were recognised. The lowermost fossiliferous unit consists of shallow channel deposits containing rust-coulored sands, small clay lenses, reworked clay pebbles, remnants of bioturbation, wood (stumps) and large mammal bones. The second unit is a channel deposit also, consisting of coarse green sands and large bones. The third unit consists mostly of white (bleached) sands intersected by small pebble layers. Bioturbation and large bones are common. The fourth unit is composed of coarse sands with large bones. These units, however, are not continuous and the correlation of the localities in different sections is therefore mainly based on fossil content. Large mammal remains were recoverd from many localities. After wet screening of sediment (with water from an oil well) rodent and lagomorph remains were found in only five localities. SYSTEMATIC LIST Vespertilionidae Gray, 1821 Vespertilionidae gen. nov. et sp. nov. Despite respresented by a single specimen only (found in the washed residuum of site MS2 by Fejfar in 1983), the chiropteran record from Jebel Zelten is of a considerable significance. It is a right mandible of a robust vespertilionid bat (CM3 alv. 7.2 mm) with well preserved myotodont molars (M1 length 1.76 mm, width 1.06/1.20 mm, M2 length 1.85 mm, width 1.20/1.22 mm). Preliminary we reported it as Scotophilus sp. (Wessels et al. 2003) as it corresponds to Recent Table 1. Species list and sequence of localities, dry sieved in the field in 1983 at sites A, B, and washed at the other sites in 1997 (from Wessels et al. 2002) Tab. 1. Soupis druhů/forem a přehled lokalit, ze kterých byl materiál nasucho prosíván roku 1983 a vymýván roku 1997 (převzato z Wesselse et al. 2002) Sites / lokality: 1 – ATH7A2 (2), 2 – ATH7A3 (2), 3 – ATH5A1 (1), 4 – ATH4B (3), 5 – QAB1C (4), 6 – MS2 (A), 7 – Wadi Shatirat (B) (sub)family Cricetidae Cricetidae Murinae Myocricetodontinae Myocricetodontinae Rhizomyinae Lophocricetinae Ctenodactylidae Thryonomyidae Ochotonidae Ochotonidae Vespertilionidae 96 sites: ?Cricetidae gen. et sp. indet. gen. et sp. indet. Potwarmus sp. nov. Mellalomys sp. cf. Myocricetodon sp. Prokanisamys sp. Heterosminthus sp. indet. Sayimys nov. sp. gen. nov. et sp. nov. Alloptox sp. ?Kenyalagomys sp. gen. nov. et sp. nov. 1 2 3 4 5 6 7 – – – – – + – – + – – – – – – – – – – – + – + – – + – – – + – – – – – – – – – + + + – + + – – – – – – + – – – – – – – – + – + + – – + + – + – + – – + – – – – + – – – – Scotophilus kuhlii Leach, 1821 or S. viridis (Peters, 1852) in degree of reduction of unisuspid teeth and robust construction of mandibular body and molars i.e. the characters by which it differs from other extant genera like Eptesicus Rafinesque, 1820, Otonycteris Peters, 1859 or Scotomanes Dobson, 1875. Of course, as demonstrated elsewhere (Horáček et al. 2006) it cannot be directly coidentified with extant Scotophilus Leach, 1821 because of less advanced form of molars and with the combination of these characters it does not respond to diagnostic criteria of any other genus, both extanct and extinct, except for partial agreement with the Paleogene African clade Philisidae Sigé, 1985 (Philisis sphingis Sigé, 1985 from Oligocen of Fayum, Dizzya exultans Sigé, 1991 from early Eocene of Chambi, Tunisia, Phylisis sevketi Sigé, 1994 from Oligocene of Taqah, Oman) from which it differs by more advanced degree of unicuspid reduction. Thus, it cannot be excluded that the Jebel Zelten bat represents a transitional stage between Phylisinae and the extant genus Scotophilus. The latter genus is now distributed throughout Africa and Madagascar (with 14 spp.) and in southern Asia. The Asiatic Scotophilus kuhlii is considered the most primitive both in morphological and molecular respects, its divergence time from the African clades can be estimated (based on molecular clock) to Middle or Late Miocene. For more details see Horáček et al. (2006). Fig. 1. Geographical map of the middle part of northern Libya with localities of fossil mammals in the Jebel Zelten escarpment (modified after Savage & Hamilton 1973). Obr. 1. Geografická mapka střední části severní Libye s nalezišti fosilních savců v Džebelu Zelten (upraveno podle Savage & Hamiltona 1973). Legend / legenda: sites / lokality 1983: A – Measured Section 2 / zaměřený profil 2 (MS2), B – Wadi Umm Ash Shatirat (WS); sites / lokality 1997: 1 – ATH5A, 2 – ATH7A, 3 – ATH4B, 4 – QAB1C. 97 Murinae Gray, 1821 Potwarmus Linsay, 1988 Potwarmus sp. nov. (Fig. 3: 1–4) The material of Potwarmus sp. nov. was recorded at the sites and layers: 1. locality 1 (MS 2), in lower part of the Qarat Jahannam member of Maradah Formation, middle part of the SW Jebel Zelten escarpment, and 2. locality at Section of Wádí Shatírát. The record of Potwarmus from Jebel Zelten is characterized with inflated median cusps (t5, t8), absence of anterior and posterior mure, enterostyle (t4) at the level of t6, and long posterior cingulum on M1. It shows some light differences in size and morphology of the cusps from Potwarmus primitivus Wessels et al., 1982 (Chinji, Pakistan), P. thailandicus Jaeger et al., 1985 (Li, Thailand) and P. minimus Wessels et al., 1987 (Pakistan). Potwarmus sp. nov. from Jebel Zelten seems to be more advanced than P. primitivus and P. thailandicus. Generally the molars of the north African form are less bunodont than other species of Potwarmus which could represent a more advanced stage of evolution in comparison with the asiatic forms. The occurrence of the genus Potwarmus in North Africa indicates a migration of this genus from southern Asia to Africa. Its migration route is unknown since primitive murines are not known from Asia minor or the Arabian peninsula. Potwarmus sp. nov. is slightly more evolved than Potwarmus from Banda daud Shah in Pakistan (Wessels et al. 1982; dated ca. 16 Ma), excluding a migration during the Early Miocene times (18 Ma) (Wessels et al. 2003). Considering the uncertainties of its relationship(s) to African muroid subfamilies, Potwarmus is regarded as a primitive murid, as is Antemus Jacobs, 1978, although both genera lack the (for true murids) characteristc chevrons of three cusps is the first upper molar. Myocricetodontinae Lavocat, 1962 Mellalomys Jaeger, 1977 Mellalomys sp. (nov. ?) (Fig. 3: 7–9) The material from Jebel Zelten is clearly distinct from Mellalomys lavocati Wessels, 1996 from Sind, Pakistan, Lower-Middle Miocene and is except for the poorly divided anterocone, clearly more evolved. The presence of a double anterocone, the short ‘normal’ longitudinal crest, an Fig. 2. Representants of selected species of rodents from the Jebel Zelten sites MS2 and Wadi Shatirat. Occlusal views of the molars, figured as left. 1–4 – Potwarmus sp. nov., M1 sin. (1); M1 (2, 3); M1 and M2 from one individual (4, inv.); 5, 6 – Heterosminthus sp. indet., M2 dext. (inv.); 7, 8 – Mellalomys sp. nov.; 7 – M1 dext. (inv.); 8 – M2 sin.; 9, 10 – ?Cricetidae gen. et sp. indet., 9 – M2 dext.; 10 – M2 dext.; 11–14 – Sayimys sp. nov., 1a – M2 dex; 1b – anterior view; 2a – M3 (inv.); 13, 14 – posterior views. Scale A to the Figs. 1–10, scale B to the Figs. 11, 12, scale C to the Figs. 13, 14. Obr. 2. Zástupci vybraných druhů hlodavců z nalezišť v Džebelu Zelten: MS2 a Wadi Shatirat. Okluzální plochy molárů jsou vyobrazeny jako levé. 1–4 – Potwarmus sp. nov., M1 sin. (1); M1 (2, 3); M1 a M2 z jediného kusu (4, inv.); 5, 6 – Heterosminthus sp. indet., M2 dext. (inv.); 7, 8 – Mellalomys sp. nov.; 7 – M1 dext. (inv.); 8 – M2 sin.; 9, 10 – ?Cricetidae gen. et sp. indet., 9 – M2 dext.; 10 – M2 dext.; 11–14 – Sayimys sp. nov., 1a – M2 dex; 1b – pohled zpředu; 2a – M3 (inv.); 13, 14 – pohledy odzadu. Měřítko A k obr. 1–10, měřítko B k obr. 11, 12, měřítko C k obr. 13, 14. 98 99 elongated anteroconid are characteristic for Mellalomys Jaeger, 1977. A short mesolophid (M1) is known to occur in primitive Mellalomys species from Pakistan (Wessels 1996). Mellalomys sp. (nov. ?) is smaller than Mellalomys atlasi (Lavocat, 1961), has lower cusps and ridges, the anterocone less well divided, the anterior ledge on the anterocone smaller (or absent) and the longitudinal crest is not oblique. Mellalomys from Jebel Zelten could represent a new species, related to and more primitive than Mellalomys atlasi from Beni Mellal (14 Ma; Jaeger 1977). Dipodidae Fischer von Waltheim, 1817 Lophocricetinae Savinov, 1970 Heterosminthus Schaub, 1930 Heterosminthus sp. indet. (Fig. 3: 5) This specimen shows similarity with Heterosminthus which has four roots on the M1 and M2, a prominent cusp on the postero-lingual edge of the protocone and lacks the lingual branch of the anteroloph. It differs from Heterosminthus in lacking the lingual branch of the posteroloph and having the metaloph connected to the posteroloph (Qiu 1996). It differs from the more progressive genus Arabosminthus Whybrow et al., 1982 by its elongate shape, the less robust cusps and anterior arm of the hypocone, the wide first labial syncline, the presence of a low connection between protocone and paracone and the strong connection between metacone and posteroloph. Heterosminthus is known from the Late Oligocene and the Miocene of Asia (Daxner-Höck 2001). Ctenodactylidae Gervais, 1853 Sayimys Wood, 1937 Sayimys sp. nov. The upper molars are similar to Sayimys intermedius De Bruijn et al., 1989 from the Middle Miocene of Pakistan (De Bruijn et al. 1989), and differ from an African miocene Ctenodactylid Africanomys pulcher Lavocat, 1961 (in Jaeger 1971) in having a transverse metalophule (the metacone is connected to the labial part of the posteroloph in A. pulcher) while the posterior lobe of the M3 is more reduced in A. pulcher. The Sayimys sp. nov. from the sites MS2 and Wadi Shatirat can be regarded as the predecessor of Africanomys pulcher. Ctenodactylids, known from the Lower Miocene of Turkey, Lower and Middle Miocene of Pakistan and Middle Miocene of North Africa (Africanomys pulcher, Beni Mellal) and Israel (Metasayimys) occur in the same Jebel Zelten localities as the Myocricetodontinae. The Jebel Zelten Ctenodactylidae are more primitive than those from Beni Mellal, they seem to have entered Africa at about the same time as the Myocricetodontinae or earlier. Myocricetodontinae Lavocat, 1962 Myocricetodon Lavocat, 1952 cf. Myocricetodon sp. In Myocricetodon cherifiensis Lavocat, 1952 and Myocricetodon parvus (Lavocat, 1961) the cusps are more voluminous and the anterior arm of the hypocone is in most M1 and M2 obli100 Fig. 3. Important remains of the Miocene rodents from Jebel Zelten, Libya (occlusal views of molars). Obr. 3. Významné nálezy miocénních hlodavců z Džebelu Zelten. Okluzální pohledy molárů. Legend / legenda: 1–3 – Potwarmus sp. (nov. ?), 1 – M1; 2, 3 – M1; 4 – Mellalomys sp., M1; 5 – Sayimys sp., M2. 101 quely directed towards the paracone, with a ‘new’ longitudinal crest formed between hypocone and paracone in M. parvus (Wessels 1996). Our specimens seem to be more primitive in these characters. Several primitive Myocricetodon species appear in the middle Miocene of Pakistan (Wessels 1996). The specimens from Jebel Zelten are similar to Myocricetodon cf. M. parvus from HGSP 8224 (Wessels et al. 1987) which shows a weakly developed anterior arm of the hypocone in the M2. Our specimens seems to be more evolved. – The origin and migration pattern of the Myocricetodontinae is not yet fully understood, but primitive Myocricetodontinae are known from the Lower Miocene of Turkey (Wessels et al. 2003; MN3) and other, more derived, Myocricetodontinae are known from Pakistan (18–13.7 Ma), Turkey (Yeni Eskihisar) and Saudi Arabia (16 Ma). The origin and initial development of the Myocricetodontinae may have been on the Arabian Peninsula. Mellalomys sp. is more primitive than Mellalomys atlasi from Beni Mellal (14 Ma) and is thus considered to be older. The Myocricetodon and Mella lomys from Jebel Zelten are more primitive than those of Beni Mellal and Berg Aukas. These Jebel Zelten localities are therefore considered to be older than Beni Mellal (14 Ma) and Berg Aukas (13 Ma). Rhizomyinae Winge, 1887 Prokanisamys De Bruijn, Hussain et Leinders, 1981 Prokanisamys sp. Prokanisamys cheek-teeth are characterised by their small size, low crowns, the cuspidate cheekteeth. The short mesolophid and short or absent mesoloph are regarded as primitive in the Rhizomyinae. The teeth from Libya are similar to the rhizomyids from the Lower Miocene of Pakistan. The oldest known rhizomyid comes from Pakistan (20 Ma; Lindsay, 1996), either derived from a (yet unknown) Pakistani cricetodontine or migrated into Pakistan from an unknown area. Prokanisamys sp. from the Jebel Zelten faunas is close to Prokanisamys major, known from Pakistani assemblages dated between 19.5 and 16.4 Ma. The Rhizomyinae from Jebel Zelten are similar to the Early Miocene taxa from Pakistan (Wessels & De Bruijn 2001) and not to the Middle Miocene forms, therefore the immigration of the Rhizomyinae into North Africa must have taken place in Early Miocene times. Prokanisamys sp. is considered by us to be ancestral to Pronakalimys from Fort Ternan (14 Ma; Tong & Jaeger 1993). Thryonomyidae Pocock, 1922 Thryonomyidae gen. nov. et sp. nov. The morphology of six specimens from localities of 1997 collection, exclude our material from the Thryonomyid-like genera: Paraulacodus Hinton, 1933; Neosciuromys Stromer, 1926; Paraphiomys Andrews, 1914; Apodecter Hopwood, 1929 and Kochalia De Bruijn & Hussain, 1985. The M3 of Rodentia indet. (only one M3 and one M3) from the Middle Miocene site from the Hadrukh Formation of eastern Saudi Arabia (Whybrow et al. 1982) is very similar to the M3 of our material. This specimen seems to represent a more evolved species of the Jebel Zelten thryonomyid. The Thryonomyidae from Jebel Zelten are considered to be more closely related to Late Eocene Phiomyids from Algeria, and less closely to the Oligocene forms of Libya and Egypt and the Miocene Phiomyids and Thryonomyidae from Eastern Africa (Lavocat 1973, Denys 1992, Winkler 1992). The Phiomyidae become extinct after the Early Miocene, the Thryonomyidae are known from the Middle Miocene of Africa, Saudi Arabia, Pakistan and India. 102 Conclusions based on the record of Rodents (Fig. 4) The presence of Potwarmus sp. and Mellalomys sp. in the assemblages of MS2 and WS places these assemblages betweeen 16 and 14 Ma. In conclusion, the fauna od small mammals of Jebel Zelten localities spans approximately 4 Millions years, from 19 Ma to 15 Ma. The differences between the particular assemblages suggest that the Jebel Zelten mammal associations represent at least three different time periods. Fig. 4. Paleogeographic map of Europe, North Africa and the Middle East during the Early Miocene around 17 Ma (late Burdigalian, Ottnangian): (1) the Eastern Mediterranean seaway is closed producing a landbridge which enabled the faunal (mammalian) interchange between Africa and Asia (B). (2) in the West the Atantic Ocean communicated with the Tethys-Mediterranean and Paratethys (A). The northern faunal (mammalian) interchange (C) was of different character and apparently separated from the southern one in the region of the today Balkan peninsula (B). The symbol of the star indicates the position of Jebel Zelten. Obr. 4. Paleogeografická mapa Evropy, severní Afriky a Blízkého východu během spodního miocénu před cca 17 mil. let (svrchní burdigal, otnang): (1) moře ve východním Středomoří se uzavřelo a tím se vytvořil pevninský most umožňující migrační výměnu savců mezi Afrikou a Asií (B). (2) na západě byl Atlantský oceán spojen se Středozemním mořem (Tethydou) a dále na severovýchod s Paratethydou (A). Severní migrační výměna savčích faun (C) se svým složením lišila od jižní (B), a byla od ní zřejmě v prostoru dnešního Balkánského poloostrova oddělena. Hvezdičkou je označena poloha Džebelu Zelten. 103 In any case, the particularly characteristic feature of Jebel Zelten small mammal assemblages is that nearly all forms composing them reveal apparent relations both to the clades known from other African or Arabian Paleogene or Neogene sites and to those found in early or middle Miocene of Pakistan. This fact corresponds well to the extensive rearrangements of the paleogeographic situation in the zone of the eastern Tethyan seaway during the Early Miocene which promoted multiple faunal interchanges between Eurasia and Africa. Two main migration waves have been recognized until now. The first, dated approximately to 18–19 Ma, and the second, dated to around 16–17 Ma (Thomas 1985, Rögl 1998). Ochotonidae, primitive cricetids, sciurids and rhizomyines invaded Africa during the first period of faunal interchange while the anthracothere Brachyodus dispersed into Europe and Pakistan. The range expansion of Myocricetodontinae and Ctenodactylidae fall most probably in the stage of the second migration wave but it was limited onto North Africa. If the age determination of Potwarmus is correct (younger than 16 Ma), then Potwarmus migrated into Africa during the Middle Miocene, perhaps during the period when Griphopothecus, Alloptox and Heterosminthus migrated into Anatolia and Central Europe (Rögl 1998). Unfortunately, the available fossil record is still too scarce to enable a detailed paleobiogeographic reconstructions of the Early to Middle Miocene Afro-Asiatic faunal interchange. In any case, the situation found in Jebel Zelten demonstrate as well the vivid dispersal dynamics in the respective taxa as the fact that the range expansions were most probably followed by well pronounced subsequent divergences in marginal populations. In result, not only the geographically distant populations diverged but apparent anagenetic shifts can be observed during the Early and Middle Miocene also on a local scale. Worth mentioning is that such a dynamics most probably characterized not only the Asiatic invaders in Africa but, as the above discussed situations in Myocricetodontidae, Phiomyidae-Thryonomyidae, and Phylisinae-Scotophilus suggest, the clades whose ranges were centered in the North Africa and/or in Arabia in the late Paleogene and which might contributed the respective interchange in an essential way too. Souhrn Práce podává přehled společenstev drobných savců získaných v průběhu let 1982, 1983 a 1997 ze spodněmiocenních uloženin Džebelu Zelten v Libyi. Celkem zde bylo nalezeno 10 druhů hlodavců, dva druhy pišťuchovitých zajícovců a jeden druh netopýra. Ve všech případech vykazují nalezené formy zřetelné vztahy jak k pozdně paleogenním či miocenním formám africkým tak k formám doloženým se spodního či středního miocenu Pakistanu. Zvláště důležitým je nález myšovitého hlodavce rodu Potwarmus, který dokládá jeden z prvních výskytů této čeledi v Africe a současně datuje nejmladší ze zkouamných faun do úseku 15–16 miliónů let. Zkoumané fauny dokládají výmluvně velmi dynamickou tzv. “jižní” faunovou výměnu mezi Afrikou a Asií související se změnami paleogeografické situace ve spodním miocenu. Současná “severní” výměna faun mezi Evropou a Asií měla odlišný charakter. References De Bruijn H. & Hussain S. T., 1985: Thryonomyidae from the Lower Man-char Formation of Sind, Pakistan. Proc. Konink. Nederl. Akad. Wetensch. B, 88: 155–166. De Bruijn H., Hussain S. T. & Leinders J. J. M., 1981: Fossil Rodents from the Murree Formation near Banda daud Shah, Kohat, Pakistan. Proc. Konink. Nederl. Akad. Wetensch. B, 84: 71–99. Horáček I., Fejfar O. & Hulva P., 2006: A new genus of vespertilionid bat from Early Miocene of Jebel Zelten, Libya, with comments on Scotophilus and early history of vespertilionid bats (Chiroptera). Lynx, n. s., 37: 131–150. 104 Jacobs L. L., 1978: Fossil rodents (Rhizomyidae and Muridae) from Neogene Siwalik deposits, Pakistan. Mus. North Arizona Press, 52: 1–95. Jaeger J.-J., 1971: Un cténodactiylidé (Mammalia, Rodentia) nouveau, Irhoudia bohlini n. g. n. sp. du Pléistocène inférieur du Maroc, rapports avec les formes actuelles et fossiles. Notes Serv. Géol. Maroc., 31(237): 113–140. Jaeger J.-J., 1977. Rongeurs (Mammalia, Rodentia) du Miocene de Beni-Mellal. Palaeovertebrata, 7(4): 91–125. Lavocat R., 1952: Sur une faune de mammifères Miocènes découverte à Beni-Mellal (Atlas marocain). Compt. Rend. Acad. Sci. Paris, 235: 189–191. Lavocat R., 1961: Le gisement de vertébrés Miocènes de Beni-Mellal (Maroc). Etude systématique de la faune de mammifères et conclusions générales. Notes Mém. Serv. Géol. Maroc, 155: 29–94; 109–144. Lavocat R., 1973: Les Rongeurs du Miocène d’Afrique Orientale. Ecole Pract. Hautes Etudes (3ème Sect.) Mém. Trav. Inst. Montpellier, 1: 1–248. Lindsay E. H., 1988: Cricetid rodents from Siwalik deposits near Chinji Village. Part I: Megacricetodontinae, Myocricetodontinae and Dendromurinae. Palaeovertebrata, 18: 95–154. Qiu Z. D., 1996: Middle Miocene Micromammalian Fauna from Tunggur, Nei Mongol. Science Press, Beijing, 216 pp. Rögl F., 1998: Palaeogeographic Considerations for Mediterranean and Paratethys Seaways (Oligocene to Miocene). Ann. Naturhist. Mus. Wien 99A: 279–310. Savage R. J. G. & Hamilton W. R., 1973: Introduction to the Miocene mammal faunas of Gebel Zelten, Libya. Bull. Brit. Mus. Natur. Hist., Geol., 22: 515–527. Thomas H., 1985: The Early and Middle Miocene land connection of the Afro-Arabian plateau and Asia: a major event of hominoid dispersal? Pp.: 42–50. In: Delson E. (ed.): Ancestors: the Hard Evidence. Alan R. Liss, Inc., New York, x–xii+1–366 pp. Tong H. & Jaeger J.-J., 1993: Muroid rodents from the middle Miocene Fort Ternan locality (Kenya) and their contribution to the phylogeny of muroids. Palaeontographica A, 229: 51–73. Wessels W., 1996: Myocricetodontinae from the Miocene of Pakistan. Proc. Konink. Nederl. Akad. Wetensch, 99: 253–312. Wessels W. & De Bruijn H., 2001: Rhizomyidae from the lower Manchar Formation (Miocene, Pakistan). Ann. Carnegie Mus., 70: 143–168. Wessels W., De Bruijn H., Hussain S. T. & Leinders J. J. M., 1982: Fossil rodents from the Chinji Formation, Banda Daud Shah, Kohat, Pakistan. Proc. Konink. Nederl. Akad. Wetensch. B, 85: 337–364. Wessels W., Fejfar O., Peláez-Campomanes P., van der Meulen A. & De Bruijn H., 2003: Miocene small mammals from Djebel Zelten, Libya. Pp.: 699–715. In: López-Martínez N., Peláez-Campomanes P. & Hernández Fernández M. (eds.): Coloquios de Paleontología. En Honor al Dr. Remmert Daams. Volumen Extraordinario 1. Universitad Complutense de Madrid, 715 pp. Whybrow P. J., Collinson M. E., Daams R., Gentry A. W. & McClure H. A., 1982: Geology, fauna (Bovidae, Rodentia) and flora of the early Miocene of eastern Saudi Arabia. Tertiary Research, 4(3): 105–120. 105 Lynx (Praha), n. s., 37: 107–110 (2006). ISSN 0024–7774 Pipistrellus pygmaeus and two more species of bats recorded on the Island of Kefalonia, Greece (Chiroptera: Vespertilionidae) Zjištění Pipistrellus pygmaeus a dalších dvou druhů netopýrů na ostrově Kefalonie, Řecko (Chiroptera: Vespertilionidae) Jiří GAISLER Institute of Botany and Zoology, Faculty of Science, Masaryk University, Kotlářská 2, CZ–611 37 Brno, Czech Republic; gaisler@sci.muni.cz received on 21 July 2006 Abstract. At a small harbour town of Poros, SE Kefalonia, numerous foraging Pipistrellus kuhlii and P. pygmaeus were acoustically recorded (Petersson D200 bat detector) on 22 and 24 June, 2006. On 22 June, a high flying noctule, probably Nyctalus lasiopterus, was also heard. These records complete the list of bats, known to occur on the island, to a total of five. INTRODUCTION AND METHODS Kefalonia (also Cephalonia, Kefaloniá, Kefallinia) is the largest of Ionian Islands. Contrary to the more northerly Corfu (Kérkira), the bat fauna of the island has been little studied. In Argostoli, the capital of the island, 10 individuals of Pipistrellus kuhlii (Kuhl, 1817) (3 males and 1 female identified) were recorded in 1908. Fitídi Cave, Karavómilos, was the only other known locality where bats were observed. In 1970, one male of Rhinolophus hipposideros (Bechstein, 1800) and in 1971, two males and one female of R. euryale Blasius, 1853 were collected in the cave. For the localities and details of the records, see Hanák et al. (2001). During a touristic sojourn at a small town of Poros in June 2006, the author and his wife recorded the echolocation signals of bats with the aid of a Petersson D200 heterodyne bat detector. Foraging bats were seen and heard under a canopy of trees in a dry bed of a brook. In addition to very numerous P. kuhlii (peak energy frequency 40 kHz), foraging individuals of P. pygmaeus (Leach, 1825) were also detected (53–55 kHz). The main goal of the paper is to contribute to the knowledge of the distribution of the latter species. The raison d’etre of this taxon, originally separated from P. pipistrellus on the basis of its relatively high echolocation signals, has been generally accepted (Simmons 2005). Its occurrence, however, is far from being sufficiently known. 107 RESULTS AND DISCUSSION A locality where bats were expected to fly was discovered in the centre of Poros at a bridge over a brook which was dry at this time of year. The only water existing during our observation of bats was a small pool ca. 5 by 0.5–1.0 m, at the left bank upstream of the brook. The locality is situated only about 100 m away from the estuary, its elevation therefore is roughly 0 m. While the bed downstream of the bridge is bare, the banks of that lying upstream are grown with de ciduous trees forming a dense canopy. On 26 June, we found that the bed was bulldozed and the pool destroyed, probably to prevent the development of mosquito larvae. This situation is shown in Fig. 1. Acoustic bat detectoring was performed on 22 June from 21:20 to 21:45 hours and on 24 June from 21:00 to 21:30 hours local (eastern European summer) time. In both cases, the greatest activity was recorded at the beginning when the bats were also detected visually. No bats were detected during the last five minutes of each observation. The bats were foraging and their feeding buzzes were frequently heard. The greatest density of foraging bats was recorded above and close to the pool but commuting and foraging bats were also recorded upstream to about 50 m from the bridge. Within the same time span no activity was recorded downstream between Fig. 1. Dry bed of a brook at Poros, Kefalonia, where foraging Pipistrellus kuhlii and P. pygmaeus were recorded by a bat detector on 22 and 24 June 2006. Photo by the author. Obr. 1. Vyschlé řečiště potoka v městečku Poros na Kefalonii, kde byli 22. a 24. 6. 2006 detektorem ultra zvuku zjištěni lovící netopýři druhů Pipistrellus kuhlii a P. pygmaeus. Snímek autora. 108 the bridge and the estuary. It was difficult to estimate the number of flying bats but there were certainly more than 10 individuals flying at the pool at one time. Judging from the density of 40 kHz and 55 kHz signals, there were ca. three times more P. kuhlii than P. pygmaeus. After the end of the first observation in the dry bed of the brook, we continued the bat detectoring walking slowly through Poros. At 21:50 h we recorded the typical loud, low frequency signals of a noctule. The bat (bats?) was heard for five seconds but was not seen, probably performing a commuting flight at a high elevation. Kefalonia is a green island and the slopes bordering Poros are grown by shrubs and some trees. Noctules, therefore, can be expected to occur in that habitat. Since the signals we recorded had the peak frequency of 18 kHz, Nyctalus leisleri (Kuhl, 1817) can be excluded and, of the two remaining species, N. lasiopterus (Schreber, 1780) is more likely than N. noctula (Schreber, 1774). According to the present evidence, P. pygmaeus is distributed from British Isles, S Scandina via, Ukraine and W Russia south to the Mediterranean (Mayer & von Helversen 2001, Hulva et al. 2004, Simmons 2005). Molecular and phylogeographic analysis demonstrated that the divergence of P. pipistrellus and P. pygmaeus probably took place in the Mediterranean region (Hulva et al. 2004). Mediterranean islands, therefore, can be expected to harbour one of the two or both Pipistrellus species, depending on habitats. While P. pipistrellus is rather a generalist, P. pygmaeus prefers lowland, in particular riparian habitats (Mayer & von Helversen 2001). In addition to Kefalonia, we recorded P. pygmaeus on Rhodes, Mallorca and Cyprus, of which the Rhodes and Cypriot records were published (Hanák et al. 2001). On Mallorca (Balearic Isles), P. pygmaeus was acoustically detected at Playa d’Alcúdia, between Port d’Alcúdia, the sea and the hills Puig de Sant Martí, on 3 and 5 September, 2002. As in Kefalonia, P. kuhlii were more frequently detected than P. pygmaeus. In the same habitat, Eptesicus serotinus was recorded on 4 September. The bats were only flying where there were trees growing (Gaisler, unpublished). According to Hanák et al. (2001), the presence of P. pygmaeus has been demonstrated by genetical analyses from four localities in the mainland Greece and with the aid of bat detectors from another 11 localities in both the mainland and two Greek islands. Very likely, the species is more common than shown by the hitherto evidence. Concerning Kefalonia, our records increase the number of bat species known to live there to five but the species identity of the noctule remains uncertain. Considering that 12 species of bats have been known from Corfu (Hanák et al. 2001), the list of the Kefalonian species is certainly incomplete. There are various habitats, from lowlands up to 1620 m a. s. l., and the island is also rich in natural caves. The situation is promising for a more systematic study the Kefalonian bat fauna. SOUHRN Početně létající a lovící netopýři druhů Pipistrellus kuhlii a P. pygmaeus byli zjištěni ultrazvukovým de tektorem (Petersson D200) ve dnech 22. a 24. 6. 2006 v přístavním městečku Poros na ostrově Kefalonia, Řecko. Dne 22. 6. byly zachyceny také signály vysoko letícího netopýra rodu Nyctalus, pravděpodobně N. lasiopterus. Včetně námi zjištěných je v současnosti z ostrova Kefalonia známo pět druhů netopýrů. LITERATURA Hanák V., Benda P., Ruedi M., Horáček I. & Sofianidou S., 2001: Bats (Mammalia: Chiroptera) of the Eastern Mediterranean. Part 2. New records and review of distribution of bats in Greece. Acta Soc. Zool. Bohem., 53: 279–346. 109 Hulva P., Horacek I., Strelkov P. P. & Benda P., 2004: Molecular architecture of Pipistrellus pipistrellus / Pipistrellus pygmaeus complex (Chiroptera: Vespertilionidae): further cryptic species and Mediterra nean origin of the divergence. Mol. Phylogenet. Evol., 32: 1023–1035. Mayer F. & von Helversen O., 2001: Sympatric distribution of two cryptic bat species across Europe. Biol. J. Linn. Soc., 74: 365–374. Simmons N. B., 2005: Order Chiroptera. Pp.: 312–529. In: Wilson D. E. & Reeder D. M. (eds.): Mammal Species of the World. Third Edition. The John Hopkins University Press, Baltimore, xxxviii+2142 pp. 110 Lynx (Praha), n. s., 37: 111–121 (2006). ISSN 0024–7774 On some “forgotten” samples of small mammals from the Western Tatra Mts – Roháče Mts (Slovakia) (Insectivora, Rodentia, Carnivora) O několika “zapomenutých” vzorcích drobných savců ze Západních Tater – Roháčů, Slovensko (Insectivora, Rodentia, Carnivora) Jiří GAISLER1 & Jan ZEJDA2 Institute of Botany and Zoology, Masaryk University, Kotlářská 2, CZ–611 37 Brno, Czech Republic; gaisler@sci.muni.cz 2 State Plant Protection Administration, Trnkova 103, CZ–628 00 Brno, Czech Republic; jan.zejda@srs.cz 1 received on 22 February 2006 Abstract. In 1964 to 1974, 511 specimens of 14 mammal species were collected in the Roháče Mts, northern Slovakia, mostly at elevations of 1,000–2,000 m. The following mammalian families and species are represented in the samples, viz, Soricidae: Sorex araneus, S. alpinus, S. minutus, Neomys fodiens; Talpidae: Talpa europaea; Arvicolidae: Clethrionomys glareolus, Chionomys nivalis, Microtus subterraneus, M. tatricus, M. agrestis; Muridae: Apodemus flavicollis; Gliridae: Muscardinus avellanarius, Dryomys nite dula; Mustelidae: Mustela nivalis. In addition, Eliomys quercinus was observed on 22 May 1964 in the valley Spálená dolina, 1,520 m a. s. l. Besides its description, an exact drawing of the animal supports the record. Details are given on the sample of 170 specimens obtained in September 1964 by 4,730 trap-nights at elevations of 1,500–2,100 m. Mammals were trapped at the forest upper edge (ecotone) and at higher situated habitats up to the mountain ridge. Alpine vole species, M. tatricus and C. nivalis, clearly dominate this sample (Table 1). Additional eight samples were obtained mostly within the forest zone and their species diversity and numbers of records per species corroborate previous data on the development of small mammal communities with respect to the ecosystem succession (Kratochvíl & Gaisler 1967). Cranial measurements were recorded of all adult and grown-up individuals of M. tatricus and C. nivalis trapped in 1964–1974 in the Roháče Mts and available as museum specimens with undamaged skulls. The data are given in Tabs. 2 and 3, together with external measurements and body weights of the same animals as recorded in the field. Although certain biometrics of the specimens may have been included in earlier statistical evaluations (Kratochvíl 1970, 1981), individual measurements of particular specimens have never been published. The purpose of their presentation in this paper is to submit data on rare and not readily obtainable material for future examination. INTRODUCTION In the 1960’s, Professor J. Kratochvíl, director of the then Institute of Vertebrate Research, CSAS, Brno, built a team to study small terrestrial mammals on the territory of the valley Roháčská dolina in the Western Tatra Mts. In addition to problems of sampling methods dealt with in two short papers (Kratochvíl & Gaisler 1964, Gaisler & Kratochvíl 1966), the main goal of the study was to follow natural succession of small mammal communities. Kratochvíl was inspired by the paper of Grodziński (1959), who investigated the succession by comparing mammal communities on an overgrown clearing and subsequent stages of a mountain forest 111 existing simultaneously and close to each other. Accordingly, six plots of a Sorbeto-Piceetum forest were selected in the Roháče Mts, representing stands of various ages from a clearing grown with trees one to 10 years old up to a forest 100–110 years of age, considered a climax stage. In 1963, small mammal communities inhabiting the plots were sampled by a standardized method and the results were published by Kratochvíl & Gaisler (1967). In addition to the material obtained in 1963, results of a preliminary sampling in 1959–1962 and a sample obtained in 1964 were included as well. The latter sample, however, was only used to complete the knowledge of the spectrum of small mammal species living on the territory in question. The original goal of the present paper was to publish both qualitative and quantitative data on a sample of small terrestrial mammals obtained in September 1964 in the Roháče Mts by trapping above the upper edge of a continuous forest, at the elevation of ca. 1,500–2,100 m. Kratochvíl himself did not participate in this trapping campaign but delegated the first author of this paper (J. Gaisler) to form a team and organize the field work. Six persons, including the second author (J. Zejda), participated in the work. The results obtained were entered into a field notebook which is still deposited with the collection of the present Institute of Vertebra Fig. 1. Professor Kratochvíl and his team in front of a log cabin near Zverovka, Roháče, September 1963. From left to right: Z. Machař (technician), F. Tenora (helminthologist), M. Lichard (microbiologist), M. Klíma (zoologist), J. Kratochvíl (zoologist), J. Šmarda (botanist), I. Toušková (zoologist), and M. Hejnolová (technician). Obr. 1. Profesor Kratochvíl a jeho tým před srubem nedaleko turistické chaty Zverovka v Roháčích, září 1963. Zleva doprava (bez titulů): Z. Machař (preparátor), F. Tenora (helmintolog), M. Lichard (mikro biolog), M. Klíma (zoolog), J. Kratochvíl (zoolog), J. Šmarda (botanik), I. Toušková (zooložka) a M. Hejnolová (technička). 112 te Biology, AS CR, Brno. After having analysed this material, we realized that further samples of small mammals collected in the Roháče Mts in 1964–1974 were also worth considering, although their documentation in field notebooks was less complete. This was probably due to other than faunistical or ecological aims of the field works, such as obtaining material for anatomical, embryological, or caryological studies. In this paper, we describe those samples briefly. Identification of the respective notebooks and names of the organizers of research are given in the next chapter (Material etc.). All persons who took part in the fieldwork, as far as recorded in the notebooks and excepting ourselves, are mentioned in Acknowledgments. In addition to the basic data of the 1964–1974 samples, we aimed at the biometrics of alpine species Chionomys nivalis and Microtus tatricus, which can still be regarded as relatively little known. Kratochvíl published three monographs focussed on these vole species, yet the first one could not concern that material because of the publication date (Kratochvíl 1956). The material collected in 1964–1974, or a portion of it, could have been included in further two monographs (Kratochvíl 1970, 1981) but only within the statistical evaluation of samples, not as measurements of particular individuals. In contrast, we give biometrical data of each complete specimen to enable their future use in both ecological and taxonomical studies. Numerous papers by Slovak mammalogists (Kocian et al. 1985, Kocian & Kocianová 2002, Kocianová et al. 2002, Martínková et al. 2004, Žiak et al. 2004, Adamcová 2004, Žiak 2005) are based on recent research of C. nivalis, M. tatricus and other small mammals in the Western Tatra Mts. Our Slovak colleagues suggested that historical data be presented on small mammal populations comparable with the recent data (Kocian in litt.). MATERIAL, LOCALITIES, METHODS The samples comprise 15 mammal species the full generic names of which are given when mentioned for the first time, henceforward by their first letter only. All taxa follow the present nomenclature, regardless of how they were entered into the respective field notebook, e.g. Microtus tatricus, not Pitymys tatricus. The notebook KR contains data on specimens collected in 1964–1966, other notebooks are specified below. The first sampling on 16 to 24 June 1964 was done by J. Kratochvíl and his collaborators in spruce forest and dwarf pine habitats in the valley Roháčská dolina at elevations ca. 1,000–1,700 m. Both standard (small) and large snap traps were used, 6,400 trap nights in total, and an unspecified number of live traps. The traps were baited alternatively with a piece of wick soaked in fat and root vegetables, and this method was applied in the second sampling as well. The total number of snap-trapped mammals was 123 (1.8% of trap nights), that of live-trapped 10, with the following distribution among the species: Micro tus subterraneus 39, Microtus tatricus 31, Clethrionomys glareolus 21, Sorex araneus 18, Chionomys nivalis 11, Microtus agrestis 8, Dryomys nitedula 2, Muscardinus avellanarius 1, Apodemus flavicollis 1 and Neomys fodiens 1. Two species, D. nitedula and N. fodiens, were recorded by only this sampling. One D. nitedula was trapped in a building, probably in the log cabin near Zverovka where the researchers were accommodated, the other in a forest 1,000 m a. s. l. N. fodiens was trapped at a brook in the valley Salatínská dolina, 1,400 m a. s. l. The next sample, most important from the point of view of this study (see Introduction), was obtained by J. Gaisler and J. Zejda plus four researchers on 14 to 23 September 1964. Line trapping with standard snap traps (4,736 trap nights in total) yielded the species Sorex minutus, Sorex alpinus, S. araneus, C. nivalis, M. tatricus, M. subterraneus, M. agrestis, C. glareolus, A. flavicollis and M. avellanarius (see Table 1 for details). Except inside a hunting lodge just beneath the forest upper edge, probably in the valley Spálený žlab (Kocian in litt.), all trappings were carried out in various habitats above the tree line in the valleys Spálená dolina and Zadná Spálená dolina, around the tarn Štvrté Roháčské pleso and on the slopes and ridges of Mts Salatín, Rákoň, Zelené and Tri Kopy. The highest situated trap line (75 traps) was set at an elevation of ca. 2050–2100 m but there was a heavy snow-fall the next night. Owing 113 to that, most traps were lost, yet a female C. nivalis was excavated from under the snow close to the first summit of Mt Tri Kopy. The third sample was obtained on 30 August to 3 September 1966 by J. Kratochvíl and his collabo rators, incl. J. Gaisler. Unspecified numbers of live traps were set in the valleys Roháčská and Spálená dolina ca. 1000–1500 m a. s. l. Only specimens that died in traps were entered into the field notebook: S. araneus 16, S. minutus 1, C. glareolus 8, C. nivalis 3, M. tatricus 2, M. subterraneus 1, A. flavicollis 2, and M. avellanarius 2. The next sampling was focussed on the beginning of the growing season and was done by J. Krato chvíl, O. Štěrba and V. Hrabě (assisted by technicians?). Trapping, probably using snap traps, was carried out on 24 to 27 April 1968. The results concerning reproduction of small mammals were published by Kratochvíl (1968), other data were either not utilized or incorporated in the data sets for the monographs on Pitymys species (Kratochvíl 1970) and C. nivalis (Kratochvíl 1981). The bag was obtained in the valley Roháčská dolina in plots II and IV (see Results), ca. 1200–1300 m a. s. l., and consisted of S. araneus 9, S. alpinus 1, Talpa europaea l, C. glareolus 21, C. nivalis 2, M. tatricus 16, M. subterraneus 11 and M. agrestis 1 (field notebook R). The following sampling was done by B. Král and V. Králová in October 1968, the exact date not spe cified. According to the field notebook RO, S. araneus 21, C. glareolus 20, C. nivalis 1, M. subterraneus 1, and A. flavicollis 4 were snap or live trapped in plot VI in the valley Roháčská dolina and on the bank of the river Orava near Podbiel. Only the name of B. Král is connected with the next three samples from the Roháče Mts (notebook CH). The first trapping was done on 28 April to 5 May 1969, the bag consisted of C. nivalis 2, M. subterraneus 2 and M. agrestis 2, and neither traps nor localities were specified. The second trapping, 9 to 16 June1969, concerned plot VI and a locality on the river Orava near Podbiel. The bag of 24 specimens included M. subterraneus 19, M. agrestis 3 and M . avellanarius 2. The third sampling by B. Král, with both snap and live traps (numbers unspecified), was done in the valley Roháčská dolina and near Oravský Podzámok on 9 to 15 June 1971. In the field notebook, the following localities are specified: environs of the chalet Ťatliakova chata, plots II, III and VI, at a brook (probably Studený potok) and above a waterfall. The bag of 30 specimens consisted of C. glareolus 5, C. nivalis 2, M. sub terraneus 8, M. tatricus 7, M. agrestis 3, A. flavicollis 4. A weasel, Mustela nivalis, was live trapped. The last sampling in the Roháče Mts was carried out by O. Štěrba, R. Obrtel and V. Hrabě in September 1974. We found neither an exact date nor information about the localities in the respective notebook (CH). The bag consisted of C. glareolus 3, M. tatricus 1, M. agrestis l and Microtus arvalis 1. Since the latter species was not recorded in any other sample and the respective specimen was not preserved, we consider the record of M. arvalis questionable and exclude it from the evaluation. Museum specimens of M. tatricus and C. nivalis collected in 1963–1974 in the Roháče Mts and deposi ted in the collection of the Institute of Vertebrate Biology were examined. Only adult and fully grown-up individuals were selected and their skulls checked for completeness. Undamaged or nearly undamaged skulls were then measured under a stereo microscope with a vernier calliper (J. Zejda). Data on external measurements and weights of the same specimens were assumed from field notebooks. All measurements are specified in Table 2. To determine male maturity and sexual activity, a testovesicular index (TVI) was calculated as a multiple of testicle and vesicular gland lengths (Hrabě 1972). RESULTS AND DISCUSSION During eleven years 1964 to 1974, 87 insectivores, 423 rodents and 1 carnivore, totalling 511 small mammals, were trapped in the Roháče Mts by Brno zoologists. The species are represented by the following number of individuals: S. araneus 81, S. alpinus 3, S. minutus 2, N. fodiens 1, T. europaea 1, C. glareolus 107, C. nivalis 58, M. subterraneus 114, M. tatricus 102, M. agrestis 20, A. flavicollis 14, M. avellanarius 6, D. nitedula 2, and M. nivalis 1. The results of the September 1964 sampling are summarized in Table 1. The total number of trapped individuals represents 3.6% of the number of trap nights. Relatively highest (18.8%) 114 Tab. 1. The main sample according to species and habitats, TN = trap nights, n = individuals, % = individuals in percent of trap nights. Habitats: A – hunting lodge; B – upper forest edge; C – dwarf pine stands; D – brook; E – tarns; F – meadows up to 2,000 m a. s. l.; G – block fields up to 2,000 m; H – ridge above 2,000 m Tab. 1. Hlavní vzorek podle druhů a prostředí, TN = počet pastí krát nocí, n = počet jedinců, % = n jako % TN. Prostředí: A – lovecká chata; B – horní hranice lesa; C – kosodřevina; D – potok; E – plesa; F – louky do 2000 m n. m.; G – balvanitá pole do 2000 m; H – hřeben nad 2000 m habitat / prostředí TN A 16 B 1200 C 925 D 620 E 300 F 680 G 920 H 75 S. araneus n % S. minutus n % S. alpinus n % C. nivalis n % M. tatricus n % M. subterraneus n % M. agrestis n % C. glareolus n % A. flavicollis n % M. avellanarius n % 0 0 0 0 0 0 0 0 0 0 0 0 0 0 2 12.5 1 6.3 0 0 11 0.9 1 0.1 1 0.1 8 0.7 14 1.2 4 0.3 1 0.1 9 0.8 0 0 1 0.1 1 0.1 0 0 0 0 8 0.9 10 1.1 1 0.1 0 0 6 0.7 0 0 0 0 2 0.3 0 0 1 0.2 2 0.3 7 1.1 11 1.8 1 0.2 6 1.0 2 0.3 0 0 1 0.3 0 0 0 0 4 1.3 8 2.7 0 0 0 0 3 1.0 0 0 0 0 1 0.2 0 0 0 0 0 0 2 0.3 16 2.4 0 0 0 0 0 0 0 0 1 0.1 0 0 0 0 14 1.5 4 0.4 1 0.1 0 0 3 0.3 0 0 0 0 0 0 0 0 0 0 1 1.3 0 0 0 0 0 0 0 0 0 0 0 0 total / celkem 3 18.8 50 4.2 26 2.8 32 5.1 16 5.3 19 2.8 23 2.5 1 1.3 n % was the bag from inside a hunting lodge where two common forest species, C. glareolus and A. flavicollis, were captured. In contrast, relatively lowest (1.3%) was the bag from a mountain ridge above 2,000 m where only one C. nivalis was found in a trap due to unfavourable weather (see Material). For various reasons the two samples A and H are incomparable with the remaining samples and are excluded from further evaluation. Among the samples B to G, that from the environs of the tarn Štvrté Roháčské pleso (E, 5.3%) and from the bank of an unspecified brook (D, 5.1%) show relatively high percentages of trapped mammals. The sample from the upper border of the forest stand (ecotone) in the valley Zadná Spálená dolina (B, 4.2%) ranks third, comprising nine species and thus showing the highest species diversity. In samples B to G, M. tatricus is represented by 45 specimens, thus being the most often trapped species. Its relative abundance ranges from 0.3% (meadows) to 2.7% (tarn environs). The second most common species, C. nivalis, is represented by 36 specimens with relative abundance ranging from 0.3% (brook) to 1.5% (rock fields), the species was not recorded in alpine meadows. In contrast, M. subterraneus, represented by the total of 33 specimens, was most often trapped in meadows 115 (2.4%). Next comes C. glareolus with 27 specimens evenly distributed among various habitats (0.3–1.0%), missing in the habitat meadows only. S. araneus was the most common shrew species. The 17 specimens of this shrew are evenly distributed over all habitats though in low numbers (0.1–0.9%). The remaining species are only represented by 1–2 specimens each. The qualitative (species) and quantitative composition of further eight samples has already been specified under Material etc. We are unable to evaluate this material according to habitats Fig. 2. Habitat of the snow vole (Chionomys nivalis) in the Spálená dolina Valley, with Mt. Tri Kopy in the background, September 1966. Both photos by J. Gaisler. Obr. 2. Stanoviště hraboše sněžného (Chionomys nivalis) na konci Spálené doliny, v pozadí v mracích vrcholky Trech kop, září 1966. Oba snímky J. Gaislera. 116 Tab. 2. External and cranial measurements of Microtus tatricus. Explanations: No. – logo of the notebook and excursion number; G – weight; H+B – head and body length; T – tail length; HF – hind foot length; E – ear length; M – male; TVI – testovesicular index (Hrabě 1972); F – female; gr – pregnant; MC – placental scars; Cb – condylobasal length; LSp – length of splanchnocranium (measured from prosthion to the aboral edge of M3); DMx – length of the maxillar diastema; Zyg – zygomatic breadth; LMd – maximum length of mandible (measured to the aboral edge of processus angularis); DMd – length of the mandibular diastema. Weight in g, measurements in mm, cranial ones to the nearest 0.1 mm Tab. 2. Tělesné a lebeční rozměry Microtus tatricus. Vysvětlivky: No. – označení protokolu a exkursní číslo; G – hmotnost; H+B – délka těla; T – délka ocasu; HF – délka zadní tlapky; E – výška ušního boltce; M – samec; TVI – testo-vesikulární index (Hrabě 1972); F – samice; gr – březí; MC – děložní skvrny; Cb – kondylobazální délka lebky; LSp – délka splanchnokrania (měřená od prosthionu k zadnímu okraji M3), DMx – délka horní diastemy; Zyg – zygomatická šířka; LMd – největší délka mandibuly (měřená po zadní okraj processus angularis); DMd – délka dolní diastemy. Hmotnost v gramech, rozměry v mm, u lebečních s přesností 0,1 mm No. KR 13 KR 38 KR 46 KR 47 KR 48 KR 63 KR 92 KR 97 KR 98 KR 100 KR 130 KR 134 KR 162 KR 214 KR 234 KR 258 RY 31 RY 33 G 31.5 25.5 31.5 28.0 48.0 29.5 26.0 24.0 30.5 29.5 26.0 29.5 26.0 23.5 25.0 24.0 31.0 25.0 H+B T HF 113 40 114 41 111 44 109 41 104 43 116 48 106 103 37 101 112 41 103 43 107 43 112 40 109 39 102 42 100 40 111 44 104 40 17.5 17.0 18.0 18.0 17.5 18.0 17.0 17.0 17.5 18.0 18.0 17.0 18.0 17.0 17.0 17.0 18.5 18.0 E M TVI F Cb 11.5 168 12.0 80 24.6 11.0 grMC 25.1 10.5 121 24.8 10.5 MC 24.1 12.0 121 25.3 12.0 110 24.7 13.0 grMC 24.1 11.5 gr 23.8 11.5 108 24.6 12.0 gr 24.2 11.0 72 R 24.6 11.0 gr 24.1 11.0 MC 24.2 11.0 MC 24.3 11.2 70 24.3 11.5 120 25.1 11.5 110 LSp DMx Zyg LMd DMd 15.3 15.1 15.3 14.9 14.7 15.4 14.8 14.8 14.9 15.5 15.2 14.9 15.3 14.8 15.1 15.1 15.4 14.3 7.7 14.5 7.7 14.4 7.8 14.8 7.7 14.6 7.5 14.3 8.1 14.7 7.5 14.4 7.2 13.8 7.8 13.7 7.5 14.5 7.4 14.1 7.9 7.5 14.5 7.4 14.2 7.5 7.8 14.2 8.1 15.2 7.5 14.3 14.5 14.4 14.3 14.3 14.2 15.2 14.2 14.1 14.6 14.2 14.5 14.5 14.2 13.9 14.5 13.8 14.7 14.1 4.2 4.1 4.2 3.6 4.1 4.7 3.8 4.1 4.3 4.1 4.1 3.9 3.8 3.8 4.0 4.1 4.3 4.2 due to the lack of pertaining data contained in the respective field notebooks. In these notebooks, trapping localities are often denoted by Roman numbers of study plots. They correspond to the plot labelling in Kratochvíl & Gaisler (1967) where plots I to VI are described in detail and entered in a map. Most often, plot VI was mentioned as the locality trapped in 1964 and later. It was situated at the elevation of 1,150–1,250 m and grown with a stand resembling an old growth forest. The large number of species and the relatively large number of trapped animals correspond to the data in Kratochvíl & Gaisler (l. c.) who recorded the largest number of species and the second largest number of individuals on plot VI. The same number of species (nine) and a still larger number of individuals was recorded on plot I, but that plot was not trapped in 1964 and later. Instead, two other plots were selected. Plot II was grown by patches of P. abies 10–20 years old at the elevation of 1,950–1,130 m and there was a permanent brook. Plot III was grown by a dense spruce forest 20–30 years old at the same elevation as plot II. Again, the samples taken in 1964 and later on correspond to the earlier data on the large number of species 117 (13 on plot II and 11 on plot III) and on the average or low number of individuals (the lowest on plot III from all plots studied) (Kratochvíl & Gaisler l. c.). Considering all samples obtained in 1964 and later on in the Roháče Mts, the high represen tation of alpine vole species M. tatricus a C. nivalis is obvious. This can be due to the frequent trappings at high elevations. In the total material, only C. glareolus and M. subterraneus were still more numerous. The former species was recorded in a variety of forest or forest-like habi tats, which corresponds to its nature as a forest generalist. The latter was most often recorded in habitats corresponding to early stages of forest succession, as far as it can be deduced from the data in the field notebooks. In general, the data underlying this paper correspond to the earlier ones presented by Kratochvíl & Gaisler (1967) and Kratochvíl (1956, 1970, 1981) and to the recent ones obtained by Slovak authors (Kocian et al. 1985, Kocianová et al. 2002, Martínková et al. 2004, Žiak et al. 2004, Adamcová 2004, Žiak 2005). The only major difference seems to be in the extremely low representation of A. flavicollis in the 1964–1974 material, compared to both earlier and later samples from the Roháče Mts, the causes of which are unclear. In addition to specimens collected, the field notebook KR contains an observation and drawing concerning a species not recorded by trapping. On 22 May 1964 at 11:30 a.m., M. Klíma (Brno) and M. C. Saint-Girons (Paris) observed a dormouse, Eliomys quercinus, in the valley Spálená dolina near a small tarn at the elevation of 1,520 m, where the yellow and blue marked tourist pathways meet (or did so at that time). That tarn was probably Zelené pleso below Mt Predné Zelené (Kocian in litt.). The dormouse was observed from a distance of two metres as it climbed down a shrub (dwarf pine?), ran slowly across an open space and climbed up another shrub. Its distinct species-specific characters were recorded, namely the grey brownish upper parts, large eye, black stripe from eye to ear, and the enlarged black and white tail tip. Its tail appeared to be longer than head and body. We consider the description and species determination by two experienced zoologists reliable. Furthermore, M. Klíma, now professor emeritus of human anatomy, is a well-known artist and illustrator. To our knowledge, this observation has not been published so far. According to Anděra & Horáček (1982), the species was mentioned “in earlier reports from Orava“. The authors do not quote the source of that information. Very likely, however, it had been adopted from Kocyan, a well known naturalist who worked in the region of Orava in the second half of the 19th century. The following is the statement by Kocyan (1888) about the garden dormouse, referred to as Myoxus quercinus: “Sehr selten. Ich habe während 24 Jahren nur zwei Exemplare aus dem Hochgebirge bekommen”. Although no particular locality can be inferred from the term “Hochgebirge” (high mountains), it implies occurrence at high elevations (Kocian in litt. – by the way, Kocian is the grandson of Kocyan). Therefore, the observation by Klíma and Saint-Girons is not as unlikely as it could appear at first sight. The external and cranial measurements of adult and grown up individuals of M. tatricus and C. nivalis available as museum specimens are given in Tables 2 and 3. In two M. tatricus and three C. nivalis, the zygomatic breadth could not be reliably determined and in one C. nivalis the maximum length of mandible was undeterminable. The skulls of other specimens were complete. Of the total number of animals trapped, only a small percentage could be used in this respect, viz., 17.6% of M. tatricus and 37.9% of C. nivalis. On the one hand, this reflects the low representation of fully adult individuals in the sample and, on the other, the unsuitability of commercial snap traps that damage skulls of captured animals. All measurements presented in Tabs. 2 and 3 lie within the minimum-maximum range known in adults of the two species (cf. Kratochvíl 1970, 1981) and they corroborate the species-specific status of the two taxa. The data are designed for future analyses and the specification of each specimen has to assist 118 Tab. 3. External and cranial measurements of Chionomys nivalis. For explanations see Tab. 2 Tab. 3. Tělesné a lebeční rozměry Chionomys nivalis. Vysvětlivky u tabulky 2 No. G H+B T HF E M TVI F Cb KR 13 31.5 113 40 17.5 11.5 168 KR 119 52.0 123 61 20.5 18.0 grMC KR 120 115 61 21.0 MC 28.8 KR 121 53.0 123 56 21.0 16.0 gr 28.6 KR 122 58.5 126 54 21.0 16.0 MC 28.7 KR 124 49.5 125 54 21.5 16.5 255 28.7 KR 125 58.5 130 61 21.0 16.0 224 29.1 KR 132 49.0 126 64 21.0 16.0 182 29.1 KR 145 53.5 125 61 21.0 16.0 75 R 30.2 KR 157 43.0 119 53 21.0 16.0 MC 27.9 KR 173 42.0 117 55 21.0 17.0 MC 28.2 KR 190 46.0 124 62 21.0 16.5 MC 29.1 KR 199 48.0 123 60 20.4 15.8 MC 29.1 KR 217 41.5 119 54 21.0 18.0 6 R 27.9 KR 262 46.5 115 62 20.0 16.0 MC 28.1 KR 299 56.5 122 57 21.0 19.0 MC 30.5 CH 286 42.5 123.5 58 20.5 16.8 MC 29.1 CH 293 43.0 128.5 59,5 20.8 16.8 gr 29.2 RY 89 54.0 115 54 19.0 18.0 120 RY 122 58.0 126 19.0 15.0 162 30.1 RY 125 54.0 116 57 21.0 15.0 224 28.5 RY 167 54.0 117 60 20.0 16.0 MC 28.2 RY 174 60.0 121 67 20.0 17.5 gr 30.1 LSp DMx Zyg LMd DMd 15.3 17.7 17.3 17.4 16.9 17.1 17.2 17.2 17.7 16.8 16.9 17.3 17.5 16.4 17.2 18.4 17.1 17.5 17.1 17.7 16.8 16.9 17.5 7.7 14.5 14.5 9.5 17.1 17.9 9.0 17.1 18.1 9.1 16.7 16.5 9.1 17.1 8.3 17.1 17.1 9.2 17.2 17.1 9.3 16.5 16.7 9.3 16.9 17.6 9.1 16.1 17.2 9.1 16.6 16.8 9.1 16.7 16.5 9.3 17.1 8.4 16.1 16.2 8.8 16.2 16.8 9.3 17.5 17.7 9.4 16.4 17.8 9.4 16.8 16.6 9.1 16.7 9.8 16.7 16.7 9.3 16.7 16.7 8.5 16.4 9.4 16.7 17.1 4.2 5.7 5.7 4.5 5.2 4.7 4.9 4.5 4.9 4.7 4.7 4.8 5.1 4.7 4.8 4.7 5.1 5.2 4.7 5.7 4.7 3.5 5.1 researchers who wish to use more sophisticated methods (epigenetic, molecular, etc.) to locate the material in the collection. SOUHRN V letech 1964 až 1974 bylo v Roháčích na severním Slovensku, většinou v nadmořských výškách 1000–2000 m, uloveno 511 jedinců 14 druhů savců. Ve vzorcích jsou zastoupeny tyto čeledě a druhy: Soricidae: Sorex araneus, S. alpinus, S. minutus, Neomys fodiens; Talpidae: Talpa europaea; Arvicolidae: Clethrionomys glareolus, Chionomys nivalis, Microtus subterraneus, M. tatricus, M. agrestis; Muridae: Apodemus flavicollis; Gliridae: Muscardinus avellanarius, Dryomys nitedula; Mustelidae: Mustela nivalis. Kromě toho byl 22. 5. 1964 pozorován ve Spálené dolině, 1520 m n. m., živý jedinec Eliomys quercinus. Pro věrohodnost pozorování svědčí nejen popis, ale i zdařilá kresba zvířete. Podrobně je zhodnocen vzorek 170 jedinců drobných savců získaný v září 1964 v nadmořských výškách 1500–2100 m odchytem do sklapovacích pastí (4730 pastí × nocí). Pasti byly kladeny při horním okraji smrkového lesa (ekotonu) a ve vyšších polohách až po hřebeny. Ve vzorku zřetelně dominují vysokohorské druhy hrabošů M. tatricus a C. nivalis (tab. 1). Dále jsou podány základní informace o složení jiných osmi vzorků z Roháčů. Tyto odchyty do sklapovacích nebo živolovných pastí byly většinou provedeny v pásmu lesa Sorbeto-Picee tum a jejich složení potvrzuje dřívější poznatky o vývoji populací drobných savců během sukcese tohoto ekosystému (Kratochvíl & Gaisler 1967). Ze sbírkového materiál druhů M. tatricus a C. nivalis z téže oblasti a období byly vybrány nepoškozené lebky dospělých a plně dorostlých kusů. Zjištěná kraniomet rická data spolu s tělesnými rozměry a údaji o váze přejatými z terénních protokolů jsou uvedena v tab. 2 119 a 3. Tento materiál mohl být částečně či zcela zahrnut v dřívějších publikacích (Kratochvíl 1970, 1981), ale jen v rámci statistického hodnocení souborů, rozměry jednotlivých kusů publikovány nebyly. Cílem jejich zveřejnění je poskytnout data o vzácném a nesnadno získatelném materiálu pro pozdější analýzu. ACKNOWLEDGMENTS The authors wish to thank Ľ. Kocian (Bratislava) for stimulus to compile the paper, various advices concer ning literature and geographic names, and readiness to consult any problems connected with the Western Tatra Mts and their small mammals. The authors are also indebted to all who participated in field trips to obtain small mammal specimens in the Roháče Mts in 1964–1974, some of whom are no more among us. They were, in alphabetic order: M. Hejnolová, V. Hrabě, J. Kocian, B. Král, V. Králová, J. Kratochvíl, L. Kratochvílová, R. Obrtel, O. Štěrba, F. Tenora, and M. Zapletal (Brno). Finally, the authors wish to thank J. Zima and P. Koubek for loan of field notebooks and authorization to study museum specimens in the collection of the Institute of Vertebrate Biology, AS CR in Brno. REFERENCES Anděra M. & Horáček I., 1982: Poznáváme naše savce [We Get to Know Our Mammals]. Mladá Fronta, Praha, 254 pp. Adamcová M., 2004: Zalezność arealów osobniczych nornika śnieznego (Chionomys nivalis mirhanrei ni) w Tatrach Zachodnich od wieku i struktury socjalnej populacji [Individual range of the snow vole (Chionomys nivalis mirhanreini) in relation to age and social structure of the population in the Western Tatras]. Rozprawa doktorska, Kraków, 93 pp (in Polish). Gaisler J. & Kratochvíl J., 1966: Zur Frage der Verwendbarkeit von Klappfallen nach Reinwaldt beim Fangen kleiner Säugetiere. Säugetierk. Mitt., 14: 42–45. Grodziński W., 1959: The succession of small mammal communities on an overgrown clearing and landslip mountain in the Beskin Średni (Western Carpathians). Ekologia Polska, A 7 (4): 83–143. Hrabě V., 1972. Contribution to the analysis of the sexual cycle of male Clethrionomys glareolus Schreb. Zool. Listy, 21: 309–318. Kocian Ľ. & Kocianová M., 2002: History of the discovery of snow vole (Chionomys nivalis) in the Car pathians. Lynx, n. s., 33: 123–131. Kocian Ľ., Kocian A., Kocianová E. & Halák K., 1985: Príspevok k poznaniu stavovcov subalpínskeho a alpínskeho stupňa Západných Tatier – Roháčov a ich bioindikačný význam [Contribution to a knowledge of vertebrates of sub-alpine and alpine zones in the West Tatra – Roháče and their bioindicative meanings]. Oravské múzeum, 2: 123–131 (in Slovak). Kocianová M., Žiak D. & Kocian Ľ., 2002: Charakteristiky individuálnych okrskov hraboša snežného tatranského (Chionomys nivalis mirhanreini Schaefer, 1935) v Západných Tatrách – Roháčoch [Home range characteristics of Tatra snow vole (Chionomys nivalis mirhanreini Schaefer, 1935) in the West Tatra mountains-Roháče]. Pp.: 121–130. In: Urban P. (ed.): Výskum a ochrana civcavcov na Slovensku V. Zborník referátov z konferencie (Zvolen 12.–13. 10. 2001) [Research and Protection of Mammals in Slovakia V. Proceedings from the conference (Zvolen 12–13 October 2001)]. Štátna ochrana prírody Slovenskej republiky, Banská Bystrica, 174 pp (in Slovak, with an abstract in English). Kocyan A., 1888: Die Säugethiere der Nord Tatra. Termeszetrajzi fuzetek, 11 (1): 41–50. Kratochvíl J., 1956: Hraboš sněžný tatranský Microtus (Chionomys) nivalis mirhanreini Schaefer, 1935 [Tatra snow vole Microtus (Chionomys) nivalis mirhanreini Schaefer, 1935]. Acta Acad. Sci. Čechoslov. Brno, 28(1): 1–39 (in Czech). Kratochvíl J., 1968: Der Antritt des Vermehrungsprozesses der kleinen Erdsäugetiere in der Hohen Tatra. Zool. Listy, 17: 299–310. Kratochvíl J., 1970: Pitymys-Arten aus der Hohen Tatra (Mam., Rodentia). Acta Sci. Natur. Brno, 4(12): 1–63. Kratochvíl J., 1981: Chionomys nivalis (Arvicolidae, Rodentia). Acta Sci. Natur. Brno, 15(11): 1–62. 120 Kratochvíl J. & Gaisler J., 1964: Der Einfluß des Köders auf die Zusammensetzung der Populationsproben von Kleinsäugern bei ökologischen und populationsdynamischen Arbeiten. Zool. Listy, 13: 289–294. Kratochvíl J. & Gaisler J., 1967: Die Sukzession der kleinen Erdsäugetiere in einem Bergwald Sorbeto -Piceetum. Zool. Listy, 16: 301–324. Martínková N., Žiak D. & Kocian Ľ., 2004: Habitatová selekcia drobných cicavcov v heterogénnom prostredí subalpínského stupňa Západných Tatier [Habitat selection of small mammals in heterogeneous landscape of subalpine zone in the Western Tatra Mountains]. Pp.: 167–181. In: Adamec M. & Urban P. (ed.): Výskum a ochrana cicavcov na Slovensku VI. Zborník referátov z konferencie (Zvolen 10.–11. 10. 2003) [Research and Protection of Mammals in Slovakia VI. Proceedings of the Conference (Zvolen 10–11 October 2003)]. Štátna ochrana prírody SR, Banská Bystrica, 189 pp (in Slovak, with an abstract in English). Žiak D., 2005: Charakteristika synúzie drobných zemných cicavcov a populacie hraboša sněžného v NPR Roháčske plesa [Characterisctics of the synusy of small terrestrial mammals and the snow vole population in the Roháčske plesa National Nature Reserve]. Doktorandská práca, Bratislava, 64+34 pp. Žiak D., Kocianová-Adamcová M., Kocian Ľ. & Martínková N., 2004: Vysoká diverzita drobných zemných cicavcov v subalpínskom stupni Západných Tatier [High diversity of small mammals in the subalpine zone of the Western Tatra Mts]. Pp.: 45–57. In: Adamec M. & Urban P. (ed.): Výskum a ochrana cicavcov na Slovensku VI. Zborník referátov z konferencie (Zvolen 10.–11. 10. 2003) [Research and Protection of Mammals in Slovakia VI. Proceedings of the Conference (Zvolen 10–11 October 2003)]. Štátna ochrana prírody SR, Banská Bystrica, 189 pp (in Slovak, with an abstract in English). 121 Lynx (Praha), n. s., 37: 123–130 (2006). ISSN 0024–7774 Brown-nosed coati (Nasua nasua vittata) on the Roraima tepui (Carnivora: Procyonidae) Výskyt nosála červeného (Nasua nasua vittata) na stolové hoře Roraima (Carnivora: Procyonidae) Pavla HAVELKOVÁ 1, Jan ROBOVSKÝ 1, Marek AUDY2 & Amelia DÍAZ DE PASCUAL3 1 Department of Zoology, Faculty of Biological Sciences, University of South Bohemia, Branišovská 31, CZ–370 05 České Budějovice, Czech Republic; pavlusa@bf.jcu.cz; JRobovsky@seznam.cz 2 Tyršova 332, CZ–679 06 Jedovnice, Czech Republic; Audy@iol.cz 3 Department of Biology, Faculty of Sciences, University of Los Andes, 5101 Mérida, Venezuela; adiaz@ula.ve received on 12 June 2006 Abstract. Tepuis are geomorphologically (physically) isolated mountains in the northeastern area of South America. Their long-time isolation has promoted the evolution of a highly endemic fauna at the genus and species level. In general, the rare incidence is typical for all vertebrates, especially for mammals (there are no fishes on the tops of the tepuis). Herein we report of two independent observations of coatis (genus Nasua) in March 2002 and January 2003 on Roraima tepui (on the borders of Venezuela, Guyana and Brazil). The observed animals possess evident distinctive black and olive-brown coloration. Based on its distribution and published reports we suppose that these individuals belong to the subspecies of Brown-nosed coati (Nasua nasua vittata). Its distribution on Roraima tepui could be interpreted as a case of habitat and food source opportunism in coatis. The observation of January 2003 includes aggressive interaction between two individuals. Our observations could be added to other examples of mammal distribution on tepuis. The exact number of coatis on Roraima, their detailed feeding strategy and the potential dependence on human activity in this area remain unclear. INTRODUCTION The Guayana highlands are a complex of igneous and sedimentary formations in the northeastern part of South America. The sedimentary formations take the form of many isolated sandstone and quartzite mesas (tepuis). These tepuis are of archean and proterozoic age, making them one of the oldest massifs on Earth (e. g. Rosales & Huber 1996, Givnish et al. 2000, Kelloff & Funk 2004). Tepuis are typical with orthogonal hillsides that caused the long isolation of their plateaus from the surrounding landscape. Consequently, very specific habitats developed on top of tepuis with a high percentage of plant and animal endemism (e. g. Rosales & Hu ber 1996, Pérez-Zapata et al. 1992, Rull 2005). In spite of the high endemism, tepuis do not house some “Lost World fauna” sensu Sir A. I. Conan Doyle. In contrast, the Roraima tepui is disproportionately much more barren than the belletristic authors supposed. All tepuis are, in fact, a paradise for geologists (and recently for speleologists too) (e. g. Rosales & Huber 1996, Audy 2003), botanists and algologists (e. g. Rull 1991, Pokorný 1996, Givnish et al. 2000, 123 Kelloff & Funk 2004, Rull 2004a, Rull 2005), and entomologists, but vertebrate zoologists are partly unsatisfied (e. g. Rosales & Huber 1996). Vertebrates are poorly distributed on tepuis – represented by some endemic frogs (e. g. Rivero 1961, Zweifel 1986, Myers & Donnelly 2001), reptiles (e. g. Myers & Donnelly 2001, Macculloch & Lathrop 2004), birds (e. g. Bar rowclough et al. 1997, Barber & Robbins 2002, Braun et al. 2003, Hilty 2003) and mammals (e. g. Pérez-Zapata et al. 1992). Mammals on tepuis are scarce and probably represent stray animals (e.g., Panthera onca in Auyán tepui) or extremely rare endemic taxa (e.g. Podoxymys roraimae) (Pérez-Zapata et al. 1992, Pokorný 1996). For a long time, the tepuis were considered as refugia for archaic forms (“living fossils”) (the Lost World hypothesis – for details see Rull 2004a, b), but, in spite of high endemism new research shows that tepui biotas have many relatively young connections with other biogeographical areas (e.g., Andean region, Amazonia etc.) (e. g. Rivero 1964, Pérez-Zapata et al. 1992, Da Silva & Patton 1998, Kelloff & Funk 2004, Steiner & Catzeflis 2004) with vertical movements of biotas during the Pleistocene glacial cycles (the Vertical Displacement hypothesis – for details see Rull 2004a, b). Nowadays, the dispersal scenario (i.e., repeated dispersion and local speciation) in the combination with vicariant scenario is more considered than simply vicariant (refuge) scenario (for details see e. g., Rull 2004a, b, c, 2005) – in this context, many so-called living fossils (e. g., the toad Oreophrynella) are now unsupported illusions. Mount Roraima (5° 12’ N, 60° 44’ W, 2810 m a. s. l.) represents one of the highest tepuis and is situated on the secondary deforested savanna (called Gran Sabana) on the borders of Venezu ela, Brazil and Guyana. Roraima possesses a typical flora with high percentage of endemism (Pokorný 1996). It is also occupied by some typical birds (Barber & Robbins 2002, Braun et al. 2003, Hilty 2003), and other endemic herpetological fauna like e.g., Roraima Bush Toad (Oreophrynella quelchii) (Rivero 1961), Colostethus roraima (Barrio 2004) and the rattlesnake Crotalus durissus ruruima (Hoge 1966) and very rare example of mammalian endemism of tepuis – akodontine cricetid Podoxymys roraimae (Pérez-Zapata et al. 1992). We report here on the occurrence of Brown-nosed coati (Nasua nasua) on the Roraima tepui. Our observations are interesting for two main reasons: first, the occurrence of coatis has not been reported from the top of tepuis yet (Mondolfi 1997) – coatis inhabit mainly woodland areas (Nowak 1991). Second, the observed coatis were of a relatively unusual appearance in comparison with indi viduals of normal-colored Nasua nasua with reddish brown to black general coloration and yellowish to dark brown below (Nowak 1991). RESULTS AND DISCUSSION The first author (PH) observed and took a picture of one unusually colored adult specimen of Brown-nosed coati, Nasua nasua (Linnaeus, 1766) on the top of Roraima tepui in March 2002 (Fig. 1). In January 2003, M.A. repeatedly observed and photographed two individuals (Figs. 2, 3) of the same unusual coloration. During both these occasions (2002 and 2003), the observed individuals were not too shy. Good quality photographs were taken in January 2003, due to repeated visits of the camp by one (?) male of coati. All observations were made during daytime, which is congruent with the known diurnal activity of coatis (Nowak 1991). The appearance of these coatis is relatively unusual in comparisons with the typical colora tion of Brown-nosed coati with orange or reddish to dark brown pelage coloration and head with white spots around the eyes (for more details see Gompper & Decker 1998). However there is extensive variation known throughout the distribution range of Nasua nasua (Gompper 124 & Decker 1998). The individual on the Figs. 2 and 3 is probably an adult male with relatively slender snout and a yellow-black pattern of pelage. Its head is black without distinct facial light markings, only one small yellow spot is situated on the right side of left eye (for left lateral view see Fig. 2). The nose is suspiciously gray. Dorsal side of the ears and fur behind them is orange or orange-brown. The black color of the head continues over the dorsal side of the neck to the level of shoulders. Feet and distal parts of limbs are black. Dorsal side of the tail Fig. 1. Our first observation of the distinct colored coati on the Roraima tepui (2002). Photo by Pavla Havelková. Obr. 1. Naše první pozorování odlišně zbarveného nosála na Roraimě (2002). Snímek Pavly Havelkové. 125 Fig. 2. The male of coati near the speleologists’ camp (2003). Photo by Marek Audy. Obr. 2. Samec nosála při návštěvě speleologického tábora (2003). Snímek Marka Audyho. is blackish melting to brown at the tip, ventral side is yellow. Approximately six black rings, which are about twice as wide as the light ones, and a black tip are well recognizable on the tail. The rest of the body (throat, trunk, proximal parts of limbs) are light – yellow or beige, but the underfur is probably black (Fig. 3). This individual was determined as Nasua nasua vittata Tschudi, 1844, based on published data (Allen 1904, Gompper & Decker 1998, Linares 1998) and consultations with the fourth author. Observed individuals correspond in almost all aspects with the description by J. A. Allen of Nasua phaeocephala (junior synonym of N. n. vittata). This form was originally described by J. J. von Tschudi as Nasua vittata from the region near Roraima tepui. In Venezuela, two subspecies of Nasua nasua occur – N. n. dorsalis Gray, 1866 in the Andean region and N. n. vittata that is distributed in the region south of the Orinoco River, but it is minimally reported from tepuis, although it was recorded from Auyán tepui (Linares 1998). It would be interesting to investigate using molecular methods how old and how much differentiated are the subspecies of Nasua nasua throughout their range. The distribution of such a relatively big carnivore is quite unexpected in so barren habitat. The third author (M.A.) had the opportunity of repeated and long-lasting observations of this coati. His observations include an aggressive interaction (a tag) between two individuals. The stronger individual drove its rival by hardly defined bark to the edge of the tepui. These repea ted observations were possible due to several visits of coatis to the expedition camp. It could 126 imply some exploitation of the human touristic activity by coatis at the top of Roraima tepui. Diet of the Brown-nosed coati consists mainly of forest floor invertebrates, fruits and carrions with a minor amount of vertebrates (fishes, snakes, infrequently chickens in urban environment, small mammals or eggs of caiman) (Bisbal 1986, Gompper & Decker 1998). Its ecology (e.g., the presence in secondary forests, etc.) implies ecological plasticity (Eisenberg 1989, Gompper & Decker 1998). In this point of view, the coati as an omnivorous opportunist is well predisposed for its distribution on tepuis. We can only estimate the diet of the Roraima coati – it may consist of rare fruits, vertebrates (e.g., the endemic frogs and akodontine rodents), invertebrates (e. g., endemic beetles and many other insects and spiders etc.), and theoretically birds or their eggs, plus finally some human foodrests. Recently Beisiegel & Mantovani (2006) showed relatively robust home range shifts of Nasua nasua in a period of three years in a pluvial tropical Atlantic forest area. These shifts could be connected to resource availability. The same authors also described coatis foraging for food (invertebrates and vertebrates) inside bromeliads as part of Fig. 3. Portrait of the coati from Roraima tepui with its remarcable distinct coloration (2003). Photo by Marek Audy. Obr. 3. Portrét neobvykle zbarveného nosála z Roraimy (2003). Snímek Marka Audyho. 127 their typical behavior. The bromeliads are relatively abundant on top of the Roraima tepui, and this behavior could thus facilitate the survival of coatis in this relatively inhospitable habitat. There are two different points of view for this coati’s distribution. It could either be caused by a stray of a few individuals, or be a regular distribution, either seasonal or permanent (stable). The following conditions could be of importance: First, the slopes of Roraima and Kukenán (the tepui in near neighborhood of Roraima) are covered by tropical rain forest which could represent a suitable and stable habitat for coatis, in contrast to the secondary deforested dry and grassy savanna. Second, the top of Roraima is relatively well accessible – a touristic activity could facilitate the access for coatis and moreover, the climb on the top should be no problem for such agile animals. It could be interesting that both our observations happened in the end of the dry season. At least four other expeditions in different seasons did not observe any coatis. It could imply some seasonal distribution of coatis on the top of Roraima (related to possible seasonal specific behavior – e.g., higher male’s exploration activity for females, higher density of individuals in forest, decrease of food sources in surrounding areas etc.). In conclusion, we have reported a rare observation of the omnivorous opportunistic Brown-nosed coati on Roraima tepui. The exact number of coatis on Roraima, their detailed feeding strategy and potential dependence on human visiting activity in such area remain unclear. We will be deeply indebted for any additional information about coatis (or some other mammals) on whichever tepui. SOUHRN Stolové hory venezuelské Guayany (tzv. tepui) jsou izolovanými horami či pohořími, u kterých jejich dlouhodobá samostatnost snížila výskyt řady živočichů a rostlin, ale na druhou stranu přispěla k vysokému podílu místního endemismu. K velmi vzácným zvířatům vrcholových partií stolových hor patří savci. Náš příspěvek zmiňuje opakované pozorování poddruhu nosála červeného (Nasua nasua vittata) na stolové hoře Roraima (hranice Venezuely, Guyany a Brazílie) v letech 2002 a 2003. Výskyt této středně velké šelmy v tomto relativně nehostinném prostředí spojujeme s všeobecným oportunismem nosálů (ve vztahu k potravě i biotopu). V roce 2003 se dokonce podařilo pozorovat agresivní střet dvou jedinců. Vzhledem k tomuto sledování se zdá, že nejde pouze o zatoulané jednotlivce, ale možná o výskyt více zvířat trvalejšího rázu. Nelze také vyloučit, že přežívání nosála červeného na Roraimě je v určité míře umožněno také přiživováním na turistech. ACKNOWLEDGEMENT Our thanks are due to Václav Mikeš (Faculty of Biological Sciences, University of South Bohemia, České Budějovice, Czech Republic) for some ornithological information from Roraima tepui and Oldřich Říčan (Faculty of Biological Sciences, University of South Bohemia) for valuable comments. This work was partly supported by the grant No. MSMT 6007665801. REFERENCES Allen J. A., 1904: New mammals from Venezuela and Colombia. Bull. Am. Mus. Natur. Hist., 20: 327–335. Audy M., 2003: Křemencový kras venezuelské Guyany [Quarcite karst of Venezuelan Guyana]. Vesmír (Praha), 82(5): 263–265 (in Czech). Barber B. R. & Robbins M. B., 2002: Nest and eggs of the Tepui antpitta (Myrmothera simplex). Wilson Bull., 114(3): 287–288. 128 Barrio-Amorós C. L., 2004: Amphibians of Venezuela, Systematic List, Distribution and References, An Update. Rev. Ecol. Lat. Am., 9(3):1–48. Barrowclough G. F., Lentino R. M. & Sweet P. R., 1997: New records of birds from Auyán-tepui, Estado Bolívar, Venezuela. Bull. B. O. C., 117(3): 194–198. Beisiegel B. M. & Mantovani W., 2006: Habitat use, home range and foraging preferences of the coati Nasua nasua in a pluvial tropical Atlantic forest area. J. Zool., London, 269: 77–87. Bisbal E. F. J., 1986: Food habits of some neotropical carnivores in Venezuela (Mammalia, Carnivora). Mammalia, 50: 329–339. Braun M. J., Robbins M. B., Milensky C. M., O’Shea B. J., Barber B. R., Hinds W. & Prince W. S., 2003: New birds for Guyana from Mts Roraima and Ayanganna. Bull. B. O. C., 123(1): 24–33. Da Silva M. N. F. & Patton J. L., 1998: Molecular phylogeography and the evolution and conservation of Amazonian mammals. Mol. Ecol., 7: 475–486. Eisenberg J. F., 1989: Mammals of the Neotropics – The Northern Neotropics. Vol. 1. The University of Chicago Press, Chicago and London, 449 pp. Givnish T. J., Evans T. M., Zjhra M. L., Patterson T. B., Berry P. E. & Sytsma K. J., 2000: Molecular evolution, adaptive radiation, and geographic diversification in the amphiatlantic family Rapateaceae: evidence from ndhF sequences and morphology. Evolution, 54(6): 1915–1937. Gompper M. E. & Decker D. M., 1998: Nasua nasua. Mammal. Species, 580: 1–9. Hilty S. L., 2003: Birds of Venezuela. 2nd Edition. Princeton University Press, Princeton & Oxford, 878 pp. Hoge A. R., 1966: Preliminary account on Neotropical Crotalinae (Serpentes, Viperidae). Mem. Inst. Butantan, 32: 109–184. Kelloff C. L. & Funk V. A., 2004: Phytogeography of the Kaieteur Falls, Potaro Plateau, Guyana: floral distributions and affinities. J. Biogeogr., 31: 501–513. Linares O. J., 1998: Mamíferos de Venezuela. Sociedad Conservacionista Audubon de Venezuela & British Petroleum (Eds.), Caracas, 691 pp. Macculloch R. D. & Lathrop A., 2004: A new species of Dipsas (Squamata: Colubridae) from Guyana. Rev. Biol. Trop., 52(1): 239–247. Mondolfi E., 1997: Lista provisional anotada de los mamíferos de la Cuenca del Río Caura, Venezuela. Pp.: 11–63. Huber O. & Rosales J. (eds.): Ecología de la Cuenca del Río Caura, Venezuela. II. Estudios especiales. Sci. Guaianae, 7: 1–473. Myers C. W. & Donnelly M. A., 2001: Herpetofauna of the Yutajé-Corocoro massif, Venezuela: second report from the Robert G. Goelet American Museum-Terramar expedition to the northwestern tepuis. Bull. Am. Mus. Natur. Hist., 261: 1–85. Nowak R. M., 1991: Walker’s Mammals of the World. Volume II. Fifth Edition. The Johns Hopkins Uni versity Press, Baltimore and London, xii+643–1629 pp. Pérez-Zapata A., Lew D., Aguilera M. & Reig O. A., 1992: New data on the systematics and karyology of Podoxymys roraimae (Rodentia, Cricetidae). Ztschr. Säugetierk., 57: 216–224. Pokorný P., 1996: Tepui [Tepui]. Vesmír (Praha), 75(10): 557–564 (in Czech). Rivero J. A., 1961: Salientia of Venezuela. Bull. Mus. Comp. Zool., 126(1): 1–207. Rivero J. A., 1964: The distribution of Venezuelan frogs V, The Venezuelan Guayana. Carib. J. Sci., 4: 411–420. Rosales J. & Huber O. (eds.), 1996: Ecología de la Cuenca del Río Caura, Venezuela. I. Caracterización general. Sci. Guaianae, 6: xx+1–131. Rull V., 1991: Contribución a la paleoecología de Pantepui y la Gran Sabana (Guayana Venezolana: clima, biogeografía y ecología. Sci. Guaianae, 2: xxii+133 pp. Rull V., 2004a: An evaluation of the Lost World and Vertical Displacement hypotheses in the Chimantá Massif, Venezuelan Guayana. Global Ecol. Biogeogr., 13: 141–148. Rull V., 2004b: Biogeography of the “Lost World“: a palaeoecological perspective. Earth-Sci. Rev., 67: 125–137. Rull V., 2004c: Is the “Lost World“ really lost? Palaeoecological insights into the origin of the peculiar flora of the Guayana Highlands. Naturwissenschaften, 91: 139–142. 129 Rull V., 2005: Biotic diversification in the Guayana Highlands: a proposal. J. Biogeogr., 32: 921–927. Steiner C. & Catzeflis F. M., 2004: Genetic variation and geographical structure of five mouse-sized opossums (Marsupialia, Didelphidae) throughout the Guiana region. J. Biogeogr., 31: 959–973. Zweifel R. G., 1986: A new genus and species of microhylid frog from the Cerro de la Neblina region of Venezuela and a discussion of relationships among New World microhylid genera. Am. Mus. Novit., 2863: 1–24. 130 Lynx (Praha), n. s., 37: 131–150 (2006). ISSN 0024–7774 A new genus of vespertilionid bat from Early Miocene of Jebel Zelten, Libya, with comments on Scotophilus and early history of vespertilionid bats (Chiroptera) Nový rod netopýra z basálního miocenu Džebelu Zelten, Libye, s poznámkami k rodu Scotophilus a rané historii vespertilionidních netopýrů (Chiroptera) Ivan Horáček1, Oldřich Fejfar2 & Pavel Hulva1 1 Department of Zoology, Charles University, Viničná 7, CZ–128 44 Praha, Czech Republic; horacek@natur.cuni.cz, hulva@natur.cuni.cz 2 Department of Geology and Palaeontology, Charles University, Albertov 6, CZ–128 43 Praha, Czech Republic; fejfar@natur.cuni.cz received on 8 December 2006 Abstract. A well preserved mandible of a vespertilionid bat is described from the MN4–5 site Jebel Zelten MS2, Libya. The bat shows a greatly derived state in most of dental characters, but it differs from the Recent genera with corresponding degree of dental reduction (Eptesicus, Scotomanes, Hesperoptenus), in shape of molars and symphyseal region. In certain respects it reminds the Recent Scotophilus and the Late Paleogene African genus Philisis. A possibility that Philisis and the Jebel Zelten bat, described here as Scotophilisis libycus gen. nov et sp. nov, form a stem line of Scotophilus is discussed in context with recent molecular data on position of the genus. Introduction The fossil records from a series of sedimentary series in Jebel Zelten in northern Libya (Savage & Hammilton 1973) represent quite important source of information on development of mammal fauna of the eastern part of North Africa in the Early and Middle Pleistocene, i.e. after the stage well documented in classical Oligocene sites in Fayum and prior the aridisation during the Messinian Crisis and the extensive rearrangements related to it. Structure of the mammalian assemblages from Jebel Zelten and details on particular finding sites were recently surveyed by Wessels et al. (2003), further data and their palaeobiogeographic implications are in another paper of this volume (Fejfar & Horáček 2006). The present contribution is confined to just one group that is particularly rare in the North African fossil records: bats. Unfortunately it is represented with just a single specimen in Jebel Zelten collections. Nevertheless, it is for more respect quite curious and intersting. The specimen belongs undoubtedly to a vespertilionid bat, which, at first sight, must be closely related to extant genera bearing a greatly advanced dental pattern (such as Scotophilus Leach, 1821 to which it was tentatively assigned in previous reports by Horáček 2001 and Wessels et al. 2003). Its actual species and even generic identity is in no way a simple and transparent, of course. We will demonstrate that in fine details the specimen differs markedly from any yet known genus 131 both extant and fossil. This fact and its possible consequences present a non-trivial problem that is worth a separate analysis. A bat of Jebel Zelten: results of a study The specimen. The specimen was found in site MS2 of Jebel Zelten, northern Libya (see Wessels et al. 2003, Fejfar & Horáček 2006 for details) which fauna dates it to MN4–5, i.e. ca. 14–16 My. The specimen is a well preserved right mandible with M1 and M2 and alveoli of all remaing teeth. The mandible is figured in Fig. 1, dimensions are (in mm, alveolar in parantheses): (I1–M3): 7.8, (C–M3): 7.2, (P4–M3): 5.8, (M1–M3): 4.6, M1–M3): 5.2, M2–M3): 3.5, M1–M2: 3.55, (I1–I3): 0.81, (I1–P4): 3.1, (C–P4): 2.4, heigth of mandible below M1: 2.4, symphysis 2.7×1.5, M1: length 1.76, trigonid length 0.85, talonid length 0.91, trigonid width 1.06, talonid width 1.20, M2: length 1.85, trigonid length 0.88, talonid length 0.97, trigonid width 1.20, talonid width 1.22. Description. (1) Dental formula is 3123, (2) unicuspid section is compressed to about 37% of tooth row and 60% of the molar row length: (3) 3 incisor alveoli minute, all of roughly the same size and their position suggest considerable overlap between crowns of the respective teeth (type C in sense of Menu 1985), (4) C alveolus large, nearly rounded (0.86 mm), not axially compressed, also (5) P3 alveolus is rounded and not compressed or displaced of the row, (6) two P4 alveoli are of the same size as that of P3 but compressed mesiodistally, (7) alveolar lenght of P4 (0.93) is almost the same as that of C. (8) molars are myotodont in sense of Menu & Sigé (1971), i.e. postcristids connect hypoconids and entoconids, hypoconulids stay appart, (9) both molars are robust with well developed talonids, (10) conspicuously thick wall of trigonids and shallow trigonid fossids (protofossids), (11) high standing lingual base of protofossid that is at the same level as the deepest inflexion of entoconid crest, (12) longitudinal distance between para- and metaconid is equal to the distance between meta- and entoconid or slightly larger (on M1), (13) hypoconulid is low standing without connection with the cingular band, (14) paraconid of M1 is somewhat tappered mesially and relatively low in comparison with metaconids, (15) there is an indistinct but clear undulation of crown base at level of metaconid, (16) alveoli of M3 are roughly of the same size as in M1 and M2, including the distance between the alveoli, the distal alveolus of M3 is not reduced, (17) the mandible body is robust and haevily build, (18) symphysis is conspicuously broad and symetrical, oval-shaped, without a chin thickenning or mesial tappering, (19) inflexion point of the line of symphysal distal margin is situated below the centre of P3 and at nearly at a half of height of the mandible body, (20) foramen mentale is situated below P3, (21) angulus mandibulae is indistinct, the base of ramal section diverges from the line of of the mandible base by 13° only. Comparisons. Most of the above surveyed characters are just the apomorphies of vespertilionid dental organisation, hence, the specimen under study belongs apparently to a clade characterized by quite advanced degree of dental specialisation. The considerable degree of premolar reduction excludes the genera bearing the less advanced pattern, first of all Myotis Kaup, 1829 or Keri voula Gray, 1842 and ancient vespertilionine genera like Karstala Czaplewski et Morgan, 2000 Fig. 1. The bat of Jebel Zelten: Scotophilisis libycus gen. nov. et sp. nov. – right mandible, lateral, lingual and occlusal view. Obr. 1. Netopýr z Džebelu Zelten: Scotophilisis libycus gen. nov. et sp. nov. – pravá mandibula, laterální, linguální a oklusální pohled. 132 133 or Hanakia Horáček, 2001 that in Miocene may come in account, eventually. Also the genera bearing the plesiomorphic nyctalodont molar pattern (cf. Menu & Sigé 1971) can be excluded: this concerns among other those to which the specimen in study approach in dental formula and general morphology, e.g. Scotoecus Thomas, 1901, Scotozous Dobson, 1875, Nyctalus Bowditch, 1825. What remains are the genera characterized by myotodont molars and considerbale degree of premolar reduction. Excluding those which differ in their small body sizes and/or in their sleder or less specialized dentitions (Hypsugo Kolenati, 1856, Neoromicia Roberts, 1926, Nycticeinops Hill et Harrison, 1987, Laephotis Thomas, 1901, Glauconycteris Dobson, 1875, Arielulus Hill et Harrison, 1987), or those improbable for geographic reasons (Australian Scotorepens Troughton, 1943, Nyctophilus Leach, 1821, Falsistrellus Troughton, 1943, Vespadellus Troughton, 1943, Neotropic Rhogeessa Allen, 1866, Baeodon Miller, 1906), the list of the extant genera which come in acount is further reduced onto: Eptesicus Rafinesque, 1820, Scotomanes Dobson, 1875, Hesperoptenus Peters, 1868, Vespertilio Linnaeus, 1758, Otonycteris Peters, 1859, Antrozous Allen, 1862, and Scotophilus Leach, 1821. None of them, fit in all the characters, of course. The fossil form exceeds the limits of variation in most genera particularly in the characters 2, 10, 11, 18, 19, 20, and 22. Considerable differences are in architecture of molars which are distinctly haevily build. Spacious and deep fossids, narrow and vertically oriented para- and metaconids in Eptesicus, Hesperoptenus, Scotomanes, Vespertilio, Antrozous, or Otonycteris, clearly contrast with high massive lingual wall and shallow bases of fossids in M1 and M2 of the fossil form. Even the most robust forms of these genera differ also in shape and position of symphysis and degree of reduction of incisor row. We stress these facts especially because just these genera, especially Eptesicus, were the first hot candidates for defaut identification of the fossil specimen at least for clear resemblance in structure of dentition, shape of mandible and last but not least for biogeographic reasons. Eptesicus is the genus recently wide-spred and taxonomically diversified in the respective region. We compared directly the fossil specimen with Eptesicus serotinus (Schreber, 1774), E. (s.) turcomanus (Eversmann, 1840), E. isabellinus (Temminck, 1840), E. anatolicus Felten, 1971, E. bottae (Peters, 1869), E. nasutus (Dobson, 1877) and E. nilssonii (Keyserling et Blasius, 1839) (besides of further taxa) and constantly confirmed the above mentioned differences. Vespertilio exhibits correspondingly high degree of reduction in incisor row but markedly differ in molar design and the same hold true for Scotomanes ornatus (Blyth, 1851) and Hesperoptenus tickelli (Blyth, 1851) that well approach the fossil specimen also in overall size. Otonycteris hemprichii (Peters, 1859) and Samonycteris majori Revilliod, 1922 which would come in account also for biogeographic reasons differ in molar design even more (comp. Figs. 3, 4). Compared to the primitive condition in vespertilionids (e.g. in Stehlinia Revilliod, 1919, Myotis, Hanakia) the fossil form exhibits an apparent increase in relative mass of trigonids. In contrast to the above mentioned extant genera, all these trends are particularly pronounced in Fig. 2. Right mandibles of Scotophilisis libycus gen. nov. et sp. nov., Jebel Zelten, leg. O. Fejfar (1ab), Scotophilus cf. viridis, Khartoum, Sudan, leg. P. Štys (2ab), Scotophilus kuhlii, CAM 40, Pakse, Laos, leg. I. Horáček (3ab), Eptesicus serotinus, Q2 Chlum4, Bohemia, leg. I. Horáček (4a), and Eptesicus (serotinus) turcomanus, CT84/19 Frunze, Kirghizia, leg. I. Horáček (5b). a – lateral view, b – occlusal view. Obr. 2. Pravé mandibuly Scotophilisis libycus gen. nov. et sp. nov., Džebel Zelten, leg. O. Fejfar (1ab), Scotophilus cf. viridis, Chartúm, Sudan, leg. P. Štys (2ab), Scotophilus kuhlii, CAM 40, Pakse, Laos, leg. I. Horáček (3ab), Eptesicus serotinus, Q2 Chlum4, Čechy, leg. I. Horáček (4a), a Eptesicus (serotinus) turcomanus, CT84/19 Frunze, Kirgisie, leg. I. Horáček (5b). a – laterální pohled, b – oklusální pohled. 134 135 Scotophilus, the genus which bears the most advanced state of the respective dental characters of all vespertilionid bats. There is a broad measure of agreement between the Jebel Zelten bat and smaller species of Scotophilus [viridis (Peters, 1852), kuhlii Leach, 1821, and partly dingani (Smith, 1833) and leucogaster (Cretzschmar, 1830)] in overal size and proportions of mandible (including relative height of mandible body, shape and form of symphysis and/or in indistinct angulus mandibulae) as well as in proportions of molar to unicuspid row or a degree of reduction of incisors and premolars contrasting to quite a large canine. In general dimensions, the fossil form falls in variation range of S. viridis. Nevertheless, all the examined extant species of Scotophilus clearly differ in the diagnostic character of the genus: considerable degree of reduction of talonids, which are compressed mesio-distally and low staying in respect to the lingual basis of a tooth. The larger forms of the genus, e.g. S. heathi (Horsfield, 1831), reach even higher degree of that trend. These differences seem to exclude a direct coidentification of the Jebel Zelten bat with Scotophilus, though it is the extant genus which shows the most similarities with it in comparison to other extant genera. Genus Scotophilus is also reported from middle Miocene of Central Europe by Engesser (1972): a well preserved mandible from MN7 Steinheim and 8 isolated teeth from MN8 Anwil (including 3 M1 with length: 2.08–2.12, and 3 M2: 2.20–2.24) were found to correspond both in size and in shape of the particular teeth (C1, M1, M2) to the situation in extant Scotophilus temminckii (Horsfield, 1824) (= S. kuhlii Leach, 1821 – comp. Simmons 2005). Unfortunately, the respective items were not figured in details and until now, we did not succeed either to examine them or to compare them and the specimen in question directly. In any case, similarly as with extant Scotophilus kuhlii, their measurements are clearly larger than those in the Jebel Zelten bat. In comparison to the extant species, the Steinheim specimen (judging from Engesser 1972: 129, Fig. 37) exhibits a lesser degree of canine and premolar mesio-distal compression and fairly less reduced talonids of M1 and M2. In these respects it might remind the Jebel Zelten bat eventually – a direct comparison is neccessary, of course. The other group that is to be taken in account is a mysterious African clade of early vespertilionids, family Philisidae Sigé, 1985. The family and its type species, Philisis sphingis Sigé, 1985, were described based on two jaw fragments (P4–M3, (P2–)P4–M2) obtained during 1961–1962 excavation in famous early Oligocene site of Jebel El Quatrani in Fayum, Egypt. The detailed analysis by Sigé (1985) can be for purpose of the present comparisons tentatively summarized as follows: the remains belonged to a very large bat (P4–M3 9.29, M1–M2 6.08 mm) exhibiting a striking differences from any genus then available in its combination of the following characters: (a) a broad basis of orbit with lateral extension of zygomata at level of M1–M2, (b) oval shaped and vertically oriented foramen infraorbitale at level of P4, (c) robust Fig. 3. M1,2 dental morphology in Scotophilisis libycus gen. nov. et sp. nov., Jebel Zelten, leg. O. Fejfar (1ab), Scotophilus cf. viridis, Khartoum, Sudan, leg. P. Štys (2b), Scotophilus kuhlii, CAM 40, Pakse, Laos, leg. I. Horáček (3ab), Eptesicus anatolicus, NMP 48192, Bavineh, Iran, leg. P. Benda (4a), Eptesicus (serotinus) turcomanus, CT84/19, Frunze, Kirghizia, leg. I. Horáček (5ab), and Otonycteris hemprichii, E40, Egypt. a – lateral view, b – occlusal view. Not in a scale. Obr. 3. Morfologie M1 a M2 u Scotophilisis libycus gen. nov. et sp. nov., Džebel Zelten, leg. O. Fejfar (1ab), Scotophilus cf. viridis, Chartúm, Sudan, leg. P. Štys (2b), Scotophilus kuhlii, CAM 40, Pakse, Laos, leg. I. Horáček (3ab), Eptesicus anatolicus, NMP 48192, Bavineh, Iran, leg. P. Benda (4a), Eptesicus (serotinus) turcomanus, CT84/19, Frunze, Kirgisie, leg. I. Horáček (5ab), and Otonycteris hemprichii, E40, Egypt. a – lateralní pohled, b – oklusální pohled. Upraveno na stejnou velikost. 136 137 but mesiodistally shortened upper molariforms (P4–M3), (d) very high inner wall of upper molars with postprotocrista terminating with a sharp sweep at a hypocone region, (e) shalow fossa without para- or metalophi, (f) split of mesostyle into mutually separated lateral terminations of inner crests of para- and metacone at M1 and M2 but (g) retaining complete plagiocrista at M3 that connects parastyle, paracone, mesostyle and metacone that is preserved at M3 despite the tooth is moderately reduced, (h) a short but broad P4 with robust mesiopalatal thickening and distinct inflexion of the distal crown margin, (i) a robust mandible body, (j) three premolars with the middle one reduced and displaced lingually, (k) haevily build lower molars with (l) high lingual walls, (m) relatively low paraconid and shalow protofossid, (n) indistinct undulation of lingual base of the teeth at level of paraconid, (o) a distinct but low standing hypoconulide, (p) hypocone is relatively lower in M2 than in M1, (r) distal alveolus of M3 not reduced, alveolar length of M3 seems to be nearly equal to that of M1. Except for (h), the mandibular characters of Philisis sphingis resemble clearly the situation demonstrated in the Jebel Zelten bat except for its size which is nearly twice larger that in the latter form. Further data concerning Philisidae arose of the analyses of bat remains from the Tunisian early Eocene fauna in Chambi (for its stratigraphical setting see Hartenberger et al. 1997): Sigé (1991) demonstrated that at least two of four bats remains found there (M1, M1–M2) fit well the diagnostic characters of the family (in general architecture of molars, particularly at M1) but differ from Philisis sphingis in its very small size (length of M1: 1.23, width of M1: 1.71, length of M2: 1.12 ), and, in particular, nyctalodont condition of the lower molars. The form described as Dizzya exsultans Sigé, 1991 further differs in surprisingly high degree of talonid reduction at M2 contrasting to broad and shalow talonid at M1. Alveoli of M3 suggests a considereble degree of reduction, too. Thus, in comparison to Philisis sphingis, Dizzya exsul tans shows a combination of both primitive characters (nyctalodoncy, small size) and greatly advanced characters (reduction od M2 and M3) for which it cannot be regarded a direct ancestor of the former one. In respect to the present comparisons, the most pertinent data are those by Sigé et al. (1994) reporting further two forms of Philisidae from the early Oligocene site Taqah from Oman. Philisis sevketi Sigé, 1994 described from there based on isolated M1 and seven other teeth resembles our specimen in available characters (comp. Fig 18 in Sigé et al. 1994) but differs in comparativelly larger size (M1 length: 2.28, talonid width: 1.91). Besides it, of course, Sigé et al. (1994) reports from the site also two incomplete fragments of upper molars (M3 and M1) identified as “cf. Philisis sp.” which are of a smaller size roughly corresponding to that of our specimen (length of M3 ca. 1.14, length of M1 ca. 1.48). Summing up the above comparisons we can conclude: (i) the Jebel Zelten bat differs clearly from the extant genera which come here in account mostly for specificities in its molar architecture and for its robust mandible, its symphyseal region, and a considerable degre of reduction of premolar and incisor rows. (ii) To a considerable degree these specificities are shared with extant genus Scotophilus. Nevertheless, all of the six species examined essentially differ in degree of reduction and positional rearrangements of talonids in which they exceed corresponding tendencies in any other vespertilionid genus including Philisis and/or the Jebel Zelten bat. (iii) The specimen in study shows a broad measure of agreement with the Early Eocene to Early Oligocene Afro-Arabian genus Philisis, particularly in shape and general architecture of M1 and M2. It markedly differ from Philisis by its smaller size and a reduced premolar row (P3 absents at all, P4 compressed). 138 Fig. 4. Type specimen of Samonycteris majori Revilliod, 1922 (Mytilini, Samos, Greece), Mus. Lausanne 945(S)/21908. Above – lateral view of the specimen, middle left – cochlear region (comp. also Horáček 1991), others – details of dentition. Obr. 4. Typový exemplář Samonycteris majori Revilliod, 1922 (Mytilini, ostrov Samos, Řecko), Mus. Lausanne 945(S)/21908. Nahoře – laterální pohled na exemplář, střed vlevo – kochleární oblast (viz také Horáček 1991), ostatní – detaily dentice. 139 Taxonomy For the above mentioned reasons, the Jebel Zelten bat cannot be coidentified with any of the yet described genera both extant and fossil. At the same time, it is clear, of course, that a more robust conclusions on its actual affinities could be drawn only after a more complete material will be available, particularly after it would support the above predictions with at least some of the upper jaw characters. Thus, it would be perhaps the most correct to stop our treatment at this point and wait for supplementing of the material with further items. Nevertheless, finally we decided to express the results of the above comparisons in form of a taxonomic opinion not only for apparent improbability of the expected record, but also as a way to emphasize the non trivial aspects of the matter in order to stimulate its further reexamination. Since, the specimen cannot be arranged under any yet described genus, its proper taxonomic treatment necessitates to propose a new genus for it. Scotophilisis gen. nov. Type Species. Scotophilisis libycus sp. nov. Derivatio nominis. Supposedely related to stem line of the extant genus Scotophilus Leach, 1821, and at the same time to a Palaeogene genus Phylisis. Diagnosis. A derived member of Phylisidae Sigé, 1985, sharing the characters of M1 and M2 architecture with Philisis sphingis Sigé, 1985, particularly the heavily build protoconide complex, high lingual crown wall and shalow protofossid of the respective teeth. It differs from Philisis by a reduced premolar row consisting of P2 and P4 only. It is further characterized by three small incisors, large canine, robust and broadly oval symphysis not extended behind a level of P2, and indistinct angulus mandibulae. It resembles Scotophilus in the latter characters but differs from it by a lesser degree of reduction and specific rearrangement of talonids, diagnostic for the extant genus (Miller 1907, Koopman 1994). From eptesicoid genera Eptesicus, Hespero ptenus or Scotomanes which share the corresponding degree of reduction of unicuspid row, it differs mainly in the molar characters shared with Philisis, in shape of symphyseal region, more advanced degree of compresion of incisor row, and smaller angle of angulus mandibulae. Scotophilisis libycus sp. nov. Holotype. Right mandible with M1 and M2, alveoli of I1 to P4 and of M3, well preserved mandible body including the symphyseal region and anterior part of ramus mandibulae, deposited in collections of National Museum Praha under number NMPC/OF.J.Zel./Chi1. Diagnosis. Same as diagnosis of the genus. Type locality and stratum. Jebel Zelten, MS2, Northern Libya, 28° 28’ N, 20° 00’E, Early Miocene, MN4–5 (see Wessels et al. 2003 and Fejfar & Horáček 2006 for details). Details. For measurements, description and comparisons see above. Discussion Vespertilionid bats represent a group characterised by a generalised state of dental and cranial characters and a prolonged maintenance of the ancestral states in most of them. The largest genus of the family, Myotis, and several other genera bear the complete dentition with unreduced number of particular tooth types while only few genera reach the level of advanced dental rearrangements at the extent common in other chiropteran families. Simply said, dental 140 evolution in vespertilionid bats was mostly restricted to clade-specific variation in relative size of particular dental elements, typically including enlargements of the molariform sector and/or canines, reductions of relative size of individual incisors or premolars, their displacements from a toothrow up to their disapperance in several clades, eventually, often accompanied with reduction of distal elements at M3/3 and relative enlargements of rostral part of skull. Worth mentioning is that until middle Miocene a vast majority of vespertilionid fossil record consisted of the forms exhibiting the ancestral dental pattern corresponding to the state in extant genus Myotis though – as demonstrated elsewhere (Horáček 2001) – they most probably belonged to the clades which extant members are characterized by rather derived dental pattern (Eptesicus, Vespertilio, a. o.). Parallel appearance of the same trends in dental rearrangements (reduction and mesio-distal compression of premolar and incisor rows, reduction of distal elements of M3/3) seems to be perhaps the most characteristic phenomenon in Late Caenozoic history of the family. Despite such characters are undoubtedly relevant (and in most instances reliable) for distinguishing of extant vespertilinid genera, for phylogenetic interpretations of the Tertiary record of the family are of a limited significance only. Nevertheless, under presumption that the respective morphoclines are irreversible, an appearance of a derived state exceeding the maximum degree observed in a Recent taxon, to which the fossil item could be assigned, can be used as an indirect argument against such decision. Such a sylogism was applied also in the above comparisons. In contrast to the above mentioned traits, the form of molars and molar design (except for M3/3, of course) represent in vespertilionid bats an extremelly conservative character which variation operates within very narrow limits characterizing particular genera and/or suprageneric clades. It provides perhaps the most reliable information for generic assignment and morphological comparisons at suprageneric level (Menu 1985). As pointed out by Menu (o.c.), in vespertilionids, the most primitive conditions in molar characters are in Myotis (which at the same time exhibits relativelly large intrageneric variation), while far the most derived state appears in Scotophilus. Scotophilus: a neontologic perspective The genus Scotophilus Leach, 1821 is distributed throughout sub-Saharan Africa, in southern Arabia, Madagascar and Reunion (Koopman 1984, 1994, Robbins et al. 1985, Gaucher 1993, Simmons 2005), and throughout considerable part of southern and southeast Asia including Phillipines, Borneo and Celebes (Corbet & Hill 1992, Bates & Harrison 1997, Kitchener at al. 1997). All species included in the genus are nearly uniform in their cranial, dental and external characters while, at the same time, in almost all dental and cranial characters they exhibit the most derived state of all vespertilionid bats (Menu 1987). This separates the genus quite distinctly from the other clades but, simultaneously, it provides almost no chance to comprehend its true affinities. The genus represents a taxonomic puzzle in all classifiations and numerous confusions accompanied also further aspects of its study. The genus Scotophilus Leach, 1821 was described based on a juvenile individual bearing milk teeth what subsequently caused numerous confusions on actual meaning and content of genus. The name was later applied nearly to all vespertilionids with a derived dental pattern (comp. Hoofer et al. 2006) until its current meaning was established by Miller (1907), who, at the same time, of course, suggested a homonymy of the name Scotophilus Leach, 1821 and Scotophila Hübner, 1816 (Lepidoptera) and denoted the genus with the name Pachyotus Gray, 141 1831. Tate (1942) proposed to include Scotophilus together with Scotomanes, Stotoecus, Scotei nus, Nycticeius, Rogeessa, Baeodon and Otonycteris (i.e. those sharing the most derived dental characters) into a separate tribe Nycticeini diagnosed among other by apomorphic condition in upper incisors with (“the outer incisor is obsolete, the inner one usually lacks the supplementary cusp and is simply conical”). Hill & Harrison (1987) suggested that Nycticeini as assembled by Tate (1942) is not a natural group and reports that Scotophilus and Scotomanes have bacula reminiscent of the flattened triangular structure of Eptesicus and its immediate associates. Because of derived dental and cranial characters they placed Scotophilus and Scotomanes in a separate tribe Scotophilini, while Otonycteris, Nycticeius, Rhogeessa and Baedon placed in a tribe Plecotini. Despite that Koopman (1994) in regard to his former proposals on reality of the tribe Nycticeini retained the traditional concept and stressed relations of Scotophilus to Scotomanes and Otonycteris. Results of molecular analyses by Hoofer & Van Den Bussche (2003) strongly contradicts any close association between Scotomanes and Scotophilus. The former genus was found to be closely related to Eptesicus, while Scotophilus is, in its mtDNA, the most derived genus within Vespertilioninae (Hoofer & Van Den Bussche 2003: 28). Our morphological comparisons supports this conclusion quite a well: Scotomanes lacks the derived characters of Scotophilus on upper molars (C type molars in sense of Menu 1985) and also its lower molars exhibit the characters of Eptesicus (low lingual wall of trigonid, deep and sharply bordered protofossid). In respect to their results, Hoofer & Van Den Bussche (2003) assigned Scotophilus to its own tribe Scotophilini and expressed serious doubt on its relationship to other vespertilionid clades (commented as “sedis mutabilis”). The detailed karyological comparisons undertaken by Vol leth et al. (2006) convincingly support the same view, too. The phylogenetic analyses perfomed with cyt b mtDNA sequences of major vespertilinid genera (Fig. 5, Appendix) suggests for Scotophilus a position at basal split of the vespertilionid stem group (worth mentioning that it immediately follows that of Miniopterus Bonaparte, 1837 and Cistugo Thomas, 1912, whose African origin is quite probable too. The genus is uniform in apomorphic condition of its dental and cranial specificities but greatly variegated in other respects (body size, pelage colouration, age and sex variation etc.). Its members can be quite easily distinguished from other vespertilionid bats but taxonomic structure of the genus, status of particular forms and their variation and phylogenetic relationships present traditionally quite confused, controversial and largerly unresolved topics. Koopman (1984, 1994) who first revised the topics in detail suggested an extensive synonymization over the many named forms and reduced the content of the genus onto 5 phenotypically well defined species (three African: borbonicus (Geoffroy, 1803), leucogaster, nigritta (Schreber, 1774), and two Asian: kuhlii, heathii). Fig. 5. Bayesian tree based on complete 1140 bp sequence of cytochrome b showing relationships among extant vespertilionid genera. Note basal position of Scotophilus. For details see Appendix. Obr. 5. Fylogenetický strom vybraných zástupců čeledi Vespertilionidae spočtený na základě sekvencí kompletního genu pro cytochrom b (technika Bayesianské analysy). Povšimněte si basální posice rodu Scotophilus. Detaily analysy jsou v appendixu. 142 143 The most detailed morphometric analysis on the African Scotophilus was undertaken by Robbins et al. (1985), who examined over 2000 specimens from throughout sub-Saharan Africa including 11 type specimens. They recognized six species of Scotophilus occurring in sub-Saharan Africa: S. dinganii, S. leucogaster, S. nigrita, S. nucella Robbins, 1983, S. nux Thomas, 1904, and S. viridis. Kitchener et al. (1997) revised the taxon in Indo-Malayan region and demonstrated a distinct species status of S. celebensis Sody, 1928 and S. collinus Sody, 1936 apart from widespread Oriental species S. kuhlii. With these additions, the results by Robbins et al. (1985) have been accepted in the most recent account of the genus by Simmons (2005) who recognized 12 species within it: S. borbonicus (Geoffroy, 1803); S. celebensis Sody, 1928; S. collinus Sody, 1936; S. dinganii (Smith, 1833); S. heathi (Horsfield, 1831); S. kuhlii Leach, 1821; S. leucogaster (Cretzschmar, 1830); S. nigrita (Schreber, 1774); S. nucella Robbins, 1983; S. nux Thomas, 1904; S. robustus Milne-Edwards, 1881; and S. viridis (Peters, 1852). The list that should be further completed with S. tanrefana Goodman, Jenkins et Ratrimomanarivo, 2005. The phylogenetic relations among particular species of the genus present another topic quite confused. Koopman (1984) and Robbins et al. (1985) proposed different phenetic groupings for the African taxa while their relations to the Asian species has not been discussed. Hoofer & Van Den Bussche (2003) based on molecular analysis of three mtDNA genes suggested a close relationship among the Ethiopian species but a distant relationship between the two Oriental species (S. heathi and S. kuhlii). The topics was recently reexamined into great details by Trujillo (2005) who analyzed 138 specimens of 10 spp. with aid od large segments of mtDNA and the Y-chromosome zfy gene. His results represent undoubtedly the most comprehensive recent contribution to taxonomy and phylogeography of the genus. He confirmed the above mentioned conclusions by Hoofer & Van Den Bussche (2003), provided evidence for two additional cryptic species within S. viridis and S.dinganii (by which the number of species incereased to 16 species) and further draw a detailed view on phylogenetic structure of the genus and its phylogeographic specificities. The study demonstrated S. kuhlii as the most basal taxon, monophyly of African Scotophilus with S. nux as the most basal African taxon, and a considerable distance between the two Indomalayan species suggesting multiple origins of Asian Scotophilus (S. heathi represents an inner clade of the African radiation related to viridis-dinganii group). It was also demonstrated that Malagasy Scotophilus appear to have affinities to different African species and likely originated from multiple colonization events from Africa. Further cryptic species was recently described from Madagascar (Goodman et al. 2006) and also the most recent studies on African Scotophilus (Ja cobs et al. 2006) demonstrated appearance of further cryptic species within the south African S. dinganii group. All that suggest that the genus may produce a considerable cladogenetic activity without marked effects on the morphometric characters of the once stabilized phenotypes. Summing up the neontologic data on the genus, it seems that: Scotophilus is a product of a very early divergence from ancestor stock of other vespertilionid clades (tribes), it is apparently of the Palaeotropic origin and its major radiation occured in Africa from where it colonized either Madagascar and neighbouring islands (Reunion with S. bourbonicus) and the Indian region (the most recently with S. heathii). In these connection it should be reminded that Scotophilus bats rank among the top vespertilionid specialist in aerial fast hawking what dispose them to distant migrations quite a well. Despite that they respresent a very common element of bat communities both in Sub-Saharan Africa and in Oriental region, they nowhere (except for the southernmost Arabia, e.g. Gaucher 1993) crossed the southern border of the Palaearctic region (Horáček et al. 2000) and entered the temperate zone. 144 Scotophilus and Philisidae: a stem line? The above surveyed neontologic evidences suggest that Scotophilus originated by early divergence from ancestral vespertilionids and its major radiation is confined to Africa. Unfortunately, as to our knowledge, any fossil record which would support these statements absents except for those from the middle Miocene of central Europe which are extralimital and should be reexamined with particular care. Nevertheless, the mandible from MN6 Steinheim figured by Engesser (1972: 129) resembles the extant forms of the genus quite a much, perhaps except for less reduced talonids. In contrast, M2 from MN8 Anwil, figured by Engesser (1972: 130) exhibits a well developed mesostyle at a buccal position instead of the deep inflexion characteristic for extant Scotophilus. If the generic affiliation of these items, suggested by Engesser (o.c.) is correct then they would suggest that as late as in the late middle Miocene the dental specificities of the genus were not attained completely, what, at the first sight, contradicts the conclusion on early divergence of the genus. The Jebel Zelten bat supports that view too (at least if the prediction of its relations to Scotophilus is actually correct, of course). It shares with the extant genus derived characters of incisor and premolar reduction, robustness of the mandible and symphyseal region and massive body of molars with high lingual walls and shallow fossids. Nevertheless, the major dental apomorphy of the extant genus – extensive reduction of talonids and their shift to the base of a tooth is here apparently not attained. Alternatively, this fact migh fit to a theoretical expectation that the most derived character states appear just at the latest stage of phylogenetic morphocline of the respective characters, at least in a case of the characters with conservative variation dynamics as in the molar shape in bats. Further extension of the same sylogism would result in a prediction that the ancestor of Scotophilisis -Scotophilus clade should share with it the essential specificities such as robustness of molars and mandible body but exhibit an ancestral stage of the common morphocline, i.e. unreduced number of premolars, incompressed incisors etc. Exactly this characterize the mandibular pattern in Philisis in which the robustness of the respective structures is moreover related to extremely large overall size. Philisidae reported as endemic African Late Paleogene group (Sigé 1985, 1991, 1994) would perfectly fit both the predictions on ancestry of the clade in question: an early divergence and the African origin. It seems that also the other specificities of Philisis can well be explained in the same way: split of the mesostyle of upper molars can well be looked upon as the state preceding the complete reduction of that structure and deep undulation of buccal crown wall at its position, characteristic of Scotophilus, similarly as the broad base of orbit with conspicuous zygomatic extension of maxilla, reported for Philisis by Sigé (1985) is partly retained in Scotophilus where it is restricted from rostral side by excessive enlargement of infraorbital region, apparently connected with conspicuous enlargement of dorsal structures of skull, the another character in which Scoto philus reached the most derived state among all vespertilionid bats. The respective concept of phylogenetic morphocline of the infraorbital region is further supported by a specific shape of infraorbital foramen in Scotophilus which is large, fissure-like and vertically tappered. In conclusion: despite the actual evidence is largerly incomplete and no direct support for the following statemetnts is available, we hypothesize that Philisidae in sense of Sigé (1985, 1991, 1994) and the newly described genus represent the stem line of Scotophilus and as suggested by distant position of the extant genus in molecular analyses, the group should be classified as separate subfamily of Vespertilionidae for which the prior available name is Philisinae Sigé, 1985. Under such a view, proposed content of the clade would be then: 145 Vespertilionidae Gray, 1821 Philisinae Sigé, 1985 stat. nov. †Dizzya Sigé, 1991 (Eocene, Africa) †Philisis Sigé, 1985 (Oligocene, Africa) †Scotophilisis gen. nov. (Miocene, Africa) Scotophilus Leach, 1821 (middle Miocene, Europe – Recent, Africa, S and SE Asia, Madagascar, Reunion). Palaeobiogeographic notes The above surveyed alternative view of relationship among the genera in question suggests also several non-trivial palaeobiogeographic hypotheses that are worth of a brief comment. To give a “safe” background to them first we have to repeat the facts which seem to be invariant in the present respects: (a) the deepest divergence within the Recent Scotophilus is between the Asian S. kuhlii group and diversified groups of the African clade including the Asian S. heathi, (b) the former Asian taxon (S. kuhlii group) seems to be more primitive both in morphological and molecular respects, (c) extensive radiation of the genus in Africa may suggest the African origin of the genus, (d) the only records beyond limits of its Recent range (i.e. Palaeotropics) are from MN7–8 in central Europe, i.e. from a retreat period of the Miocene Climatic Optimum (Böhme 2003), some 12–13 My, (e) the mtDNA distance between S. kuhlii and the African species is 16–22% (Trujillo 2005) what indicates the divergence time of some 10–12 My, according to the most widely used calibration for animal mtDNA (Brown et al. 1979) – about two percent sequence divegence between pairs of lineages per million years or one percent divergence per lineage per million years, (f) Philisis and related taxa exhibit even at the early Oligocene much higher degree of dental specialisation than any vespertilionid clade known from the Early and Middle Miocene. This holds true also for the Early Miocene Jebel Zelten bat. (g) The Asian and/or North Tethyan elements in Jebel Zelten fauna (Muridae a.o.) suggest closing of the Tethyan seaway in the eastern Mediterranean and Iranian region (comp. Rögl 1999, Hrbek & Mayer 2003) as well as expansion of open habitats and onset of aridisation of the Eastern Mediterranean in the early Miocene (Tschernov 1992). It cannot be excluded that both these factors (immigration of North Tethyan elements and spread of open ground habitats) contributed to extensive phenotypical rearrangement in the above discussed African vespertilionid clade and finally established the apomorphic design of Scotophilus and its subsequent spread to the North Tethyan and Asian range. Both the molecular clock estimate and the above mentioned records from Central Europe suggest dating of that event to about 12 My, i.e. some 4 My after the situation illustrated by the Jebel Zelten bat. Worth mentioning is that the supposed transition from the stem line to the crown taxon (Scotophilus) and spread of its range is situated in the period when this most probably occurred also in other vespertilionid clades such as Eptesicus, Hypsugo etc. (Sigé & Legendre 1983, Ziegler 2000, Horáček 2001), i.e. in the peak of the Miocene Climatic Optimum (Böhme 2003, Zachos et al. 2001). Souhrn Dobře zachovaná spodní čelist vespertilioidního letouna je popsána z naleziště MS2 z Džebelu Zelten v Libyi. Nález vykazuje vysoce odvozenou vývojovou úroveň ve většině dentálních znaků, ale liší se zároveň od současných rodů s odpovídajícím stupněm redukce zubů (Eptesicus, Scotomanes, Hesperoptenus), a to v utváření molárů a symfysy. V určitém ohledu připomíná současný rod Scotophilus a svrchnopaleogénní 146 africký rod Philisis. V kontextu výkladů molekulárních analys je v článku diskutována možnost, že formy Scotophilus, Phylisis a letoun z Džebelu Zelten, popsaný zde jako Scotophilisis libycus gen. nov. et sp. nov. vytvářejí spoečnou vývojovou linii. Acknowledgements We wish to thank to all collegues who kindly enabled us to study the comparative materials under their care: Petr Benda & Vladimír Hanák (Praha), Jiří Gaisler & Jan Zima (Brno), John Edwards Hill (London), Dieter Kock & Gerhard Storch (Frankfurt am Main), Renate Angermann and Robert Asher (Berlin), Kurt Bauer & Friederike Spizenberger (Wien), and Burkhard Engesser (Basel). Petr Benda kindly discussed previous version of the manuscript. Special thanks go to Bernard Sigé, a leading personality in study of fossil bats, who kindly discussed many details of the topics though he is in no way responsible for the opinions presented here. The study was supported by grant GAČR 206/05/2334 and COST B23.3 (IH). References Bates P. J. J. & Harrison D. L., 1997: Bats of the Indian Subcontinent. Harrison Zoological Museum, Seveoaks, 258 pp. Böhme M., 2003: The Miocene Climatic Optimum: evidence from ectothermic vertebrates of Central Europe. Paleaogeography, Paleaoclimatology, Palaeoecology, 195: 389–401. Brown W. M., George M. Jr. & Wilson A. C., 1979: Rapid evolution of animal mitochondrial DNA. Proc. Natl. Acad. Sci. USA, 76: 1967–1971. Corbet G. B. & Hill J. E., 1992: The Mammals of the Indomalayan Region. A Systematic Review. Oxford University Press, Oxford, 488 pp. Engesser B., 1972: Die obermiozäne Säugetierfauna von Anwil (Baselland). Inauguraldissertation. University of Basel, Basel. Fejfar O. & Horáček I., 2006: The Early Miocene mammalian assemblages in Jebel Zelten, Libya. Lynx, n. s., 37: 95–105. Gaucher P., 1993: First record of Scotophilus leucogaster (Cretzschmar, 1826) (Mammalia: Chiroptera: Vespertilionidae) in Saudi Arabia. Mammalia, 57: 146–147. Goodman S. M., Jenkins R. K. B. & Ratrimomanarivo F. H., 2005: A review of the genus Scotophilus (Chiroptera: Vespertilionidae) on Madagascar, with the description of a new species. Zoosystema, 27: 867–882. Goodman S. M., Ratrimomanarivo F. H. & Randrianandrianina F. H., 2006: A new species of Scotophilus (Chiroptera: Vespertilionidae) from western Madagascar. Acta Chiropterol., 8: 21–37. Hartenberger J.-L., Crochet J.-Y., Martinez C., Feist M., Godinot M., Mannai Tayech B., Marandat B. & Sigé B., 1997: Le gisement de mammiferes de Chambi (Éocene, Tunisie centrale) dans son contexte géologiquie, apport a la connaissance de l’évolution des mammiferes en Afrique. Mém. Trav. E. P. H. E. Inst. Montpellier, 21: 163–274. Huelsenbeck J. P. & Ronquist F. R., 2001: MRBAYES: Bayesian inference of phylogenetic trees. Bio informatics, 17: 754–755. Hill J. E. 1980: The status of Vespertilio borbonicus E. Geoffroy, 1803 (Chiroptera: Vespertilionidae). Zool. Mededel., 55: 287–295. Hill J. E. & Harrison D. L., 1987: The baculum in the Vespertilioninae (Chiroptera: Vespertilionidae) with a systematic review, a synopsis of Pipistrellus and Eptesicus, and the descriptions of a new genus and subgenus. Bull. Brit. Mus. Natur. Hist., Zool., 52: 225–305. Hoofer S. R. & Van Den Bussche R. A., 2003: Molecular phylogenetics of the chiropteran family Vespertilionidae. Acta Chiropterol., 5(Suppl.): 1–63. Horáček I., 1991: Enigma of Otonycteris: ecology, relationship, classification. Myotis, 29: 17–30. Horáček I., 2001: On the early history of vespertilionid bats in Europe: the Lower Miocene record from the Bohemian Massif. Lynx, n. s., 32: 123–154. 147 Horáček I., Hanák V. & Gaisler J., 2000: Bats of the Palearctic region: a taxonomic and biogeographic review. Pp.: 11–157. In: Wołoszyn B. W. (ed.): Proceedings of the VIIIth European Bat Research Symposium. Vol. I. Approaches to Biogeography and Ecology of Bats. Chiropterological Information Center, Institute of Systematics and Evolution of Animals PAS, Kraków, 280 pp. Hrbek T. & Meyer A., 2003: Closing of the Tethys Sea and the phylogeny of Eurasian killifishes (Cyprinodotiformes: Cyprinodontidae). J. Evol. Biol., 16: 17–36. Jacobs D. S., Eick G. N., Schoeman M. C. & Matthee C. A., 2006: Cryptic species in an insectivorous bat, Scotophilus dinganii. J. Mammal., 87: 161–170. Kitchener D. J., Packer W. C. & Maryanto I., 1997: Morphological variation among populations of Scophilus kuhlii (sensu lato) Leach, 1821 (Chiroptera: Vespertilionidae) from the Greater and Lesser Sunda Islands, Indonesia. Trop. Biodiver., 4: 53–81. Koopman K. F., 1984: A Progress Report on the Systematics of African Scotophilus (Vespertilionidae). Pp.: 102–113. In: Proceedings of the Sixth International Bat Research Conference. Ile-Ife, Nigeria. Koopman K. F., 1994: Chiroptera: Systematics. Handbook of Zoology. Volume VIII. Mammalia. Walter de Gruyter, Berlin and New York, 224 pp. Menu H., 1985: Morphotypes dentaires actuels et fossiles des chiroptéres vespertilioninés. 1e partie: Ètude des morphologies dentaires. Palaeovertebrata, 15: 71–128. Menu H., 1987: Morphotypes dentaires actuels et fossiles des chiroptéres vespertilioninés. 2eme partie: Implications systematiques et phylogenetiques. Palaeovertebrata, 17: 77–150. Menu H. & Sigé B., 1971: Nyctalodontie te myotodontie, importants characteres de grades evolutifs chez les chiropteres entomophages. Compt. Rend. Acad. Sci. Paris, 272: 1735–1738. Miller G. S., 1907: The families and genera of bats. U. S. Natl. Mus. Bull., 57: i–xviii+1–282. Murphy W. J., Eizirik E., O’Brien S. J., Madsen O., Scally M., Douady C. J., Teeling E., Ryder O. A., Stanhope M. J., de Jong W. W. & Springer M. S., 2001: Resolution of the early placental mammal radiation using Bayesian phylogenetics. Science, 294: 2348–2351. Posada D. & Crandall K. A., 1998: Modeltest: testing the model of DNA substitution. Bioinformatics, 14: 817–818. Robbins C. B., De Vree F. & Van Cakenberghe V., 1985: A systematic revision of the African bat genus Scotophilus (Vespertilionidae). Zool. Wetensch., 246: 51–84. Rögl F., 1999: Circum Mediterranean Miocene Paleogeography. Pp.: 9–24. In: Rössner G. & Heissig K. (eds.): The Miocene Land Mammals of Europe. Pfeil Verlag, München, 516 pp. Savage R. J. G. & Hamilton W. R., 1973: Introduction to the Miocene mammalian fauna of Gebel Zelten, Libya. Bull. Brit. Mus. Natur. Hist., Geol., 22: 515–527. Sigé B., 1975: Les Insectivores et Chiropteres du Paleogene moyen d’Europe dans l’histoire des faunes de Mammiferes sur ces continent. J. Palaeontol. Soc. India, 20: 178–190. Sigé B., 1985: Les chiroptéres oligocenes du Fayum, Egypte. Geol. Palaeontol., 19: 161–189. Sigé B., 1991: Rhinolophoidea et Vespertilionoidea (Chiroptera) du Chambi (Eocene inférieur de Tunisie). Aspects biostratigraphique, biogéographique et paléoécologique de l’origine des chiropteres modernes. N. Jb. Geol.-Paläontol. Abh., 182: 355–376. Sigé B. & Legendre S., 1983: L’histoire des peuplements de chiropteres du bassin méditerranéen: l’apport comparé des remplissages karstiques ez des dépot fluvio-lacustres. Mém. Biospél., 10: 209–225. Sigé B., Thomas H., Sen S., Gheerbrant E., Roger J. & Al-Sulaimani Z., 1994: Les chiropteres de Taqah (Oligocene inferieur, Sultanat d’Oman). Premier inventaire systématique. Münch. Geowiss. Abh. (A), 26: 35–48. Simmons N. B., 2005: Order Chiroptera. Pp.: 312–529. In: Wilson D. E. & Reeder D. M. (eds.): Mammal Species of the World. A Taxonomic and Geographic Reference. Third Edition. Volume 1. The John Hopkins University Press, Baltimore, xxxviii+743 pp. Solounias N., 1981: Mammalian fossils of Samos and Pikermi. Part 2. Resurrection of a classic Turolian fauna. Ann. Carnegie Mus., 50: 231–270. Storch G., 1999: Order Chiroptera. Pp.: 81–90. In: Rössner G. & Heissig K. (eds.): The Miocene Land Mammals of Europe. Pfeil Verlag, München, 516 pp. 148 Swofford D. L., 2001: PAUP* 4.0b10. Smithsonian Institution, Sunderland, Massachusetts. Tate G. H. H., 1942: Results of the Archbold Expeditions. No. 47. Review of the vespertilionine bats : with special attention to genera and species of the Archbold collections. Bull. Am. Mus. Natur. Hist., 80: 221–297. Trujillo R. G., 2005: Phylogenetics of the genus Scotophilus (Chiroptera: Vespertilionidae): perspectives from paternally and maternally inherited genomes with emphasis on African species. PhD Disseration, Texas A&M University, 91 pp. Volleth M., Heller K.-G. & Fahr J., 2006: Phylogenetic relationship of three “Nycticeiini” genera (Vespertilionidae, Chiroptera, Mammalia) as revealed by karyological analysis. Mammal. Biol., 71: 1–12. Wessels W., Fejfar O., Paláez-Camponars P., Van Der Meulen A. & De Bruijn H., 2003: Miocene small mammals from Jebel Zelten, Libya. Pp.: 699–715. In: López-Martínez N., Peláez-Campomanes P. & Hernández Fernández M. (eds.): Coloquios de Paleontología. En Honor al Dr. Remmert Daams. Volumen Extraordinario 1. Tschernov E., 1992: Eurasian-African biotic exchanges through the Levantine corridor during the Neogene and Quaternary. Cour. Forsch.-Inst. Senckenberg., 153: 103–123. Zachos J., Pagani M., Sloan L., Thomas E. & Billups K., 2001: Trends, rhythms, and aberrations in global climate 65 Ma to present. Science, 292: 686–693. Ziegler R., 2000: The bats (Chiroptera,Mammalia) from the Late Oligocene Fissure Fillings Herrlingen 8 and Herrlingen 9 near Ulm (Baden-Württemberg). Senckenberg. Lethaea, 80: 642–683. Appendix Molecular phylogenetics Sequences were aligned by eye since there were no ambiguities in coding sequences of cytochrome b. Phylogenetic analyses were performed in PAUP* 4.0b2 (Swofford 1998). The model of sequence evolution was inferred using Modeltest 3.06 (Posada & Crandall 1998) and used for computing of Bayesian tree. Bayesian analysis was chosen because of its ability of processing large number of taxa with complex model of sequence evolution, effective exploration of posterior probability landscape and straightforward interpretation of Bayesian probabilities (see e.g. Murphy et al. 2001). Bayesian analysis was performed in MrBayes (Huelsenbeck & Ronquist 2001) using MCMC analysis with one cold and three incrementally heated chains with length of 1.000,000 replicates and under the GTR+I+G model of DNA substitution with R-matrix = (0.2397, 13.7245, 0.3104, 0.5302, 9.9049, 1.0000), base frequencies = (0.3922, 0.3018, 0.0562, 0.2498), gamma shape parameter = 0.484 and burnin = 10,000 based on empirical evaluation. List of taxa included in the analysis and GenBank accession numbers of respective sequences (in alphabetical order): Chalinolobus tuberculatus (Forster, 1844): AF321051; Cistugo seabrae (Thomas, 1912) 1: AJ841962; Cistugo seabrae 2: AY485685; Cistugo lesueuri Roberts, 1919: AY485687; Eptesicus diminutus Osgood, 1915: AF376833; Eptesicus fuscus (Beauvois, 1796): AF376835; Eptesicus hotten totus (Smith, 1833): AJ841963; Eptesicus nilssonii (Keyserling et Blasius, 1839): AF376836; Eptesicus serotinus (Schreber, 1774): AF376837; Harpiocephalus mordax Thomas, 1923: AJ841971; Hypsugo savii (Bonaparte, 1837): AJ504450; Kerivoula papillosa (Temminck, 1840) 1: AJ841969; Kerivoula papillosa 2: AJ841970; Laephotis wintoni Thomas, 1901: AJ841964; Lasiurus Gray, 1831 sp.: AF376838; Miniopterus fraterculus Thomas et Schwann, 1906: AJ841975; Miniopterus fuliginosus Hodgson, 1835: AB085735; Miniopterus natalensis (Smith, 1834): AJ841977; Miniopterus schreibersii (Kuhl, 1817): AY208139; Murina cyclotis Dobson, 1872: AJ841972; Murina leucogaster Milne-Edwards, 1872: AB085733; Myotis albescens (Geoffroy, 1806): AF376839; Myotis annectans (Dobson, 1871): AJ841956; Myotis bechsteinii (Kuhl, 1817): AF376843; Myotis blythii (Tomes, 1857): AF376842; Myotis brandtii (Eversmann, 1845): AF376844; Myotis capaccinii (Bonaparte, 1837): AF376845; Myotis chinensis (Tomes, 1857): AB106588; Myotis dasycneme (Boie, 1825): AF376846; Myotis dominicensis Miller, 1902: AF376848; Myotis emar ginatus (Geoffroy, 1806): AF376849; Myotis formosus (Hodgson, 1835): AB106592; Myotis frater Allen, 1923: AB106593; Myotis hasseltii (Temminck, 1840): AF376850; Myotis horsfieldii (Temminck, 1840): 149 AF376851; Myotis ikonnikovi Ognev, 1912: AB106603; Myotis keaysi Allen, 1914: AF376852; Myotis levis (Geoffroy, 1824): AF376853; Myotis lucifugus (Le Conte, 1831): AF376854; Myotis macrodactylus (Temminck, 1840): AB085736; Myotis macrotarsus (Waterhouse, 1845): AF376856; Myotis montivagus (Dobson, 1874): AF376858; Myotis muricola (Gray, 1846): AF376859; Myotis myotis (Borkhausen, 1797): AF376860; Myotis mystacinus (Kuhl, 1817): AF376861; Myotis nattereri (Kuhl, 1817): AF376863; Myotis nigricans (Schinz, 1821): AF376864; Myotis oxyotus (Peters, 1867): AF376865; Myotis petax Hollister, 1912: AB106590; Myotis pruinosus Yoshiyuki, 1971: AB085737; Myotis ricketti (Thomas, 1894): AB106608; Myotis riparius Handley, 1960: AF376866; Myotis ruber (Geoffroy, 1806): AF376867; Myotis sicarius Thomas, 1915: AJ841951; Myotis thysanodes Miller, 1897: AF376869; Myotis velifer (Allen, 1890): AF376870; Myotis volans (Allen, 1866): AF376872; Myotis welwitschii (Gray, 1866): AF376874; Myotis yanbarensis Maeda et Matsumura, 1998: AB106610; Myotis yumanensis (Allen, 1864): AF376875; Neoromicia capensis (Smith, 1829): AJ841965; Nyctalus leisleri (Kuhl, 1817): AF376832; Nyctalus noctula (Schreber, 1774): AJ841967; Perimyotis subflavus (Cuvier, 1832): AJ504449; Pipistrellus abramus (Temminck, 1838): AB085739; Pipistrellus hesperidus (Temminck, 1840): AJ841968; Pipistrellus kuhlii (Kuhl, 1817): AJ504444; Pipistrellus nathusii (Keyserling et Blasius, 1839): AJ504446; Pipistrellus pipistrellus (Schreber, 1774): AJ504443; Pipistrellus pygmaeus (Leach, 1825): AJ504441; Plecotus auritus (Linnaeus, 1758): AB085734; Scotophilus heathi (Horsfield, 1831): AF376831; Vespertilio murinus (Linnaeus, 1758): AF376834; Vespertilio sinensis (Peters, 1880): AB085738. 150 Lynx (Praha), n. s., 37: 151–159 (2006). ISSN 0024–7774 Bats of the middle and upper Kızılırmak regions, Central Anatolia, Turkey (Chiroptera) Netopýři oblasti středního a horního Kizilirmaku, střední Anatolie, Turecko (Chiroptera) Ahmet Karataş1 & Mustafa Sözen2 1 Department of Biology, Faculty of Science-Arts, Niğde University, TR–51100 Niğde, Turkey; rousettus@hotmail.com 2 Department of Biology, Faculty of Science-Arts, Zonguldak Karaelmas University; TR–67100 Incivez-Zonguldak, Turkey; spalaxtr@hotmail.com received on 11 July 2006 Abstract. In this study, we found 15 bat species in the middle and upper Kızılırmak regions of central Anatolia. Four of these species were new for this region, while four previously recorded species were not found. Our new results raise the total number of species recorded in these subregions to 19. The taxonomic position of Plecotus bats, which is under discussion recently, was evaluated and P. teneriffae was recorded under its correct name for the first time in this region. Among the provinces in the area, Niğde and Kayseri have more bat species than the others by 11 species. Introduction The Central Anatolian region, one of seven geographic regions of Turkey, is divided into four subregions. Two of these, the middle and upper Kızılırmak subregions are situated in the eas tern part of the region. There are the following provinces in these subregions: Çankırı, Kayseri, Kırıkkale, Kırşehir, Nevşehir, Niğde, Sivas, and Yozgat (Fig. 1). The area has a xeric continental climate. Bat species recorded in this region have been reviewed by Benda & Horáček (1998), who also added some new species for this region. According to Benda & Horáček (1998), 15 bat species have been found in this region. However, individuals of Plecotus auritus and P. austriacus recorded in these areas are under discussion and may actually belong to other species of the genus (see Spitzenberger et al. 2006). This study is a part of a large project on bat fauna of Turkey. Material and Methods In total, 102 specimens were captured using mist net or hand net in the middle and upper Kızılırmak subregions of central Anatolia in 1994–2006. 38 individuals were released after measuring. The voucher samples were prepared in a standard way, the skins, skulls and bacula were deposited in the Mammal Collection of the Department of Zoology, Niğde University, Turkey (ZDNU). 151 ReCORDS We recorded altogether 15 bat species in the middle and upper Kızılırmak subregions of central Anatolia (Table 1). Rhinolophus ferrumequinum (Schreber, 1774) Kayseri: İncesu, Kırkinler Cave, 22 May 2005: 1 female (ZDNU 2005/35) (leg. O. Kizilcik); Sarnıcın Dere vicinity, 20 May 2005: 1 male (ZDNU 2005/36) (leg. O. Kizilcik). – Nevşehir: Avanos, Karakaya vicinity (ancient rocky settlement), 5 August 1997: 1 sad. female (ZDNU 1997/01), Avanos, between Sarılar and Özkonak, Hemdionun Cave, 19 September 2001: 3 ad. females (ZDNU 2001/224–226). – Niğde: Gümüşler, Epçik Cave, 13 October 2001: 1 ad. female (ZDNU 2002/174); near Gebere Dam Lakelet (1411 m), 21–22 July 2005: 1 ad. female (lact.) (released); 23–24 July 2005: 1 ad. female (lact.) (ZDNU 2005/66), 1 ad. female (lact.) (released); Ulukışla, Çiftehan, Maden Village, mines, 17 June 2002: 1 ad. male (ZDNU 2002/41). – Sivas: Akıncılar, Deliklitaş Cave, 9 September 2004: 1 ad. male (leg. R. Bİlgİn & A. Karataş) (ZDNU 2004/346), 1 ind. (obs.); Hâfik, Koşutdere Village (mines), 31 July 2003: 1 ad. ind. (mummy) (ZDNU 2003/52). – Yozgat: Hacıbekir Farm, ruins of Çeşka Castle, 12 April 2002: 1 ad. female (ZDNU 2002/22) (leg. H. C. Öztekİn); Darıcı Village, 17 July 2001: 1 ad. male (ZDNU 2001/81) (leg. H. C. Öztekİn). Rhinolophus hipposideros (Bechstein, 1800) Kayseri: Yeşilhisar, Soğanlı, 7 July 2001: 4 ad. females (ZDNU 2001/39–42) (leg. Y. Karakaya). – Niğde: Gümüşler, Epcik Cave, 13 October 2001: 1 ad. male (ZDNU 2002/175); Çamardı, Akpınar Alabalık Fishfarm, 10 September 2006: 1 sad. male (ZDNU 2006/93), ca. 10 sad. ind. (obs.); Ulukışla, Gümüş Village (cave), May 1996: 1 ind. (obs.); – Sivas: Hâfik, 3–4 km NW, Yıkılgan vicinity, 10 August 2003: 1 ad. male (ZDNU 2003/51). Myotis myotis (Borkhausen, 1797) Kayseri: Kocasinan, Kuşçu, Sarıağıl Cave, 12 July 2001: 1 ad. female (ZDNU 2001/52) (cf. Karataş et al. 2003); Melikgazi, 2 km NW of Gürpınar, 13 July 2001: 1 ad. male (ZDNU 2002/81); Bünyan, Karadayı Village, Karatay Hanı Caravanserai, 19 September 2002: 1 ad. male (ZDNU 2002/132); Talas, Başakpınar Örenyeri vicinity, 13 July 2001: 1 ad. male (ZDNU 2001/65). – Nevşehir: Avanos, between Sarılar and Özkonak, Hemdionun Cave, 19 September 2001: 1 ad. male, 1 ad. female (ZDNU 2001/227, 228); Gülşehir, Açıksaray, Açıksaray Ruins (ca. 1150 m), 28 August 2001: 1 ad. female (ZDNU 2001/147); Hacıbektaş, Kütükçü Village, 28 August 2001: 1 ad. female (ZDNU 2001/146); Kozaklı, Kaşkışla Village (cave), 28 August 2001: 1 ad. male, 1 ad. female (ZDNU 2001/148, 152). – Niğde: Gümüşler, Epcik Cave, 28 May 2001: 1 pregnant female (measured); 29 July 2002: 2 ad. males (ZDNU 2002/103, 110); Uluağaç Village, Uluağaç Dam Lakelet (artificial cave), 2 May 1996: 1 ad. female (ZDNU 1996/40); Ulukışla, Öküz Mehmet Paşa Caravanserai, 15 August 1999: 2 ad. males (ZDNU 1999/32–33). – Yozgat: Yozgat (center), 26 May 2002: 1 ad. male (ZDNU 2002/31). Myotis blythii (Tomes, 1857) Kayseri: Kocasinan, Kuşçu, Sarıağıl Cave, 12 July 2001: 1 sad. male, 1 ad. female (ZDNU 2001/53, 60) (cf. Karataş et al. 2003). – Niğde: near Gebere Dam Lakelet (1411 m), 21–22 July 2005: 1 ad. male (ZDNU 2005/61); Aşlama (artificial cave), 21 April 2001: 1 ad. male (ZDNU 2001/17); Çamardı, Çukurbağ Village, Ecemiş Stream (ca. 1445 m), 14–15 August 2005: 1 male (net. Ş. Özkurt & A. Karataş) (ZDNU 2005/89); Ulukışla, Öküz Mehmet Paşa Caravanserai, 15 August 1999: 1 ad. male (ZDNU 1999/31). –Yozgat: Şefaatli, Armağan Village, 15 July 2001: 1 ad. male (ZDNU 2001/82) (leg. H. C. Öztekİn). Myotis aurascens Kusjakin, 1935 Niğde: N. U. Campus, Milli Piyango Hostel, 6 May 2003: 1 ad. female (ZDNU 2003/06). 152 Myotis brandtii (Eversmann, 1845) Yozgat: Hacıbekir Farm, Çeşka Castle ruins, 12 April 2002: 1 ad. female (ZDNU 2002/23) (leg. H. C. Öztekİn) (cf. Benda & Karataş 2005). Eptesicus serotinus (Schreber, 1774) Niğde: near Gebere Dam Lakelet (1411 m), 23–24 July 2005: 2 ad. males (ZDNU 2005/70, 71); 1 ad. male, 1 sad. female (released); 12–13 August 2006: 1 ad. male (released). Fig. 1. Map of the area under study. Obr. 1. Mapa studovaného území. Legend / legenda: Kayseri: 1. Bünyan, Karadayı, 2. İncesu, Kırkinler Cave, 3. Sarnıcındere, 4. Kocasinan, Kuşçu, 5. Melikgazi, Gürpınar, 6. Talas, Başakpınar, 7. Yahyalı, 8. Yeşilhisar, Soğanlı; Kırıkkale: 9. Kırıkkale (centrum); Kırşehir: 10. Kırşehir (centrum); Nevşehir: 11. Nevşehir (centrum), 12. Avanos, Hemdionun Cave, 13. Derinkuyu, Suvermez, 14. Gülşehir, Açıksaray, 15. Tozköy, 16. Hacıbektaş, Kütükçü, 17. Kozaklı, Kaşkışla, 18. Ürgüp, Sarıhıdır; Niğde: 19. Niğde (centrum), Kayabaşı, 20. Niğde University Campus, 21. Gebere Dam, 22. Aşlama, 23. Gümüşler Monastery, 24. Gümüşler, Eski Gümüş, 25. Gümüşler, Epcik Cave, 26. Sazlıca, Akkaya Dam, 27. Uluağaç, 28a. Çamardı, Çukurbağ, 28b. Akpınar Fishfarm, 29. Ulukışla, 30. Gümüş, 31. Maden, 32. Bolkar mines; Sivas: 33. Akıncılar, 34. Hâfik, Koşutdere, 35. Hâfik, Yıkılgan, 36. Zâra; Yozgat: 37. Yozgat (centrum), 38. Çeşka Castle, 39. Darıcı, 40. Şefaatli, Armağan. 153 Myotis capaccinii (Bonaparte, 1837) Kayseri: Kocasinan, Kuşçu, Sarıağıl Cave, 12 July 2001: a maternity colony of ca. 200 ind. (obs.); 3 ad. males (ZDNU 2001/61–63) (cf. Karataş et al. 2003). – Nevşehir: Gülşehir, Tozköy Airport (leg. H. Bİl gen), 22 September 2003: 1 ad. male (ZDNU 2003/108); Ürgüp, Sarıhıdır Village, Armutludelik (a tunnel near Kızılırmak River) (890 m), 6 September 2004: 1 ad. male (ZDNU 2004/344), 2 ind. from a colony of ca. 20–30 ind. (released). Pipistrellus pipistrellus (Schreber, 1774) s. l. Kırıkkale: Kırıkkale (centrum), Gürler Quarter (house), 7 November 2003: 1 ad. male (ZDNU 2003/138) (leg. H. Yurtseven). – Kırşehir: Kırşehir (centrum), 9 October 2004: 1 male (ZDNU 2004/363); 1 male, 2 females (leg. Ş. Özkurt) (released). – Nevşehir: Nevşehir (centrum) (house), 12 November 2003: 1 ad. female (leg. H. Bİlgen) (ZDNU 2003/137). – Niğde: Kayabaşı Quarter, 1996: 1 ad. cf. female (ZDNU 55/1996); June 2001: 1 ind.; near Gebere Dam Lakelet (1411 m), 21–22 July 2005: 1 ad. male (ZDNU 2005/62), 1 ad. male (released); 23–24 July 2005: 2 ad. males (released); Aşlama (house), 21 April 2001: 1 ad. female (ZDNU 2001/17); 2 ind. (deposited in Ankara Univ.); Sazlıca, near Akkaya Dam Lakelet; spring 2002: net. 2 ad. males (released); Ulukışla, Çiftehan, Maden Village (house), 23 May 2001: 1 sex ? (ZDNU 2001/17), 1 male (deposited in Ankara Univ.), a colony of 20–30 ind. (obs.). Pipistrellus kuhlii (Kuhl, 1817) Kayseri: ca. 2 km N of Yahyalı, a stream along road to Develi, 14–15 July 2001: net 2 ad. males. Hypsugo savii (Bonaparte, 1837) Niğde: near Gebere Dam Lakelet (1411 m), 23–24 July 2005: 3 ad. males (ZDNU 2005/67–69); 12–13 Au gust 2005: 2 ad. females (net. A. Karataş & Ş. Özkurt) (ZNDU 2005/83–84); 13–14 August 2005: 1 ad. female (net. A. Karataş & Ş. Özkurt) (ZDNU 2005/87). Plecotus macrobullaris Kuzjakin, 1965 Kayseri: Bünyan, Karadayı Village, Karatay Hanı Caravanserai, 19 September 2002: 1 ad. female (ZDNU 2002/133); Talas, Başakpınar, Örenyeri, 13 July 2001: 1 ad. male (ZDNU 2001/64; as P. auritus in Karataş et al. 2003). – Nevşehir: Derinkuyu, Suvermez, 15 June 2002: 1 ad. male (ZDNU 2002/40); – Niğde: Gümüşler, Eski Gümüş (house under construction), August 1999: a colony of 10–15 ind. (obs.); Gümüşler Monastery, 20 July 1996: 1 ad. male, 2 ad. females (ZDNU 42–44/1996; as P. auritus in Karataş et al. 2003); Ulukışla, Çiftehan, Maden Village, Bolkar mines, snow tunnels, 27 June 2001: 1 ad. male (coll. A. Karataş & K. Ünsal; as P. auritus in Karataş et al. 2003). – Sivas: Hâfik, 3–4 km NW, Yıkılgan vicinity, 10 August 2003: 1 ad. female, 1 sad. male (ZDNU 2003/53–54), 1 sad. female (released). Plecotus teneriffae Barrett-Hamilton, 1907 Kayseri: Melikgazi, 2 km NW of Gürpınar, 13 July 2001: 1 sad. male, 1 sad. female (ZDNU 2001/69, 70; as P. austriacus in Karataş et al. 2003); Yeşilhisar, Soğanlı, Eski Soğanlı, Ballık vicinity, 06 July 2001: 3 juv. males, 2 ad. females (lact.) (ZDNU 2001/49–51, 72, 80; leg. Y. Karakaya; as P. austriacus in Karataş et al. 2003). – Niğde: N. U. Campus, Milli Piyango Hostel (leg. N. İpek), 28 September 2003: 1 ad. female (ZDNU 2003/101). Miniopterus schreibersii (Kuhl, 1817) Kayseri: Kocasinan, Kuşçu, Sarıağıl Cave, 12 July 2001: a maternity colony of ca. 300 ind. (obs.); 2 males, 4 females (ZDNU 2001/54–59) (cf. Karataş & Sözen 2004). – Nevşehir: Ürgüp, Sarıhıdır Village, Armutludelik (tunnel) (890 m), 6 September 2004: 1 ad. male (ZDNU 2004/341), 9 ind. from a colony of ca. 200 ind. (released) (cf. Karataş & Sözen 2004). – Niğde: N. U. Campus, 22 September 2003: 1 male (found dead) (ZDNU) (cf. Karataş & Sözen 2004); near Gebere Dam Lakelet (1411 m), 23–24 July 154 Table 1. List of bat species recorded in the study area; + literature records (see Benda & Horáček 1998, Spitzenberger et al. 2006); * first recorded; ** previously recorded under different names Tab. 1. Přehled druhů netopýrů nalezených ve studovaném území: + publikované nálezy (viz Benda & Horáček 1998, Spitzenberger et al. 2006); * nalezen poprvé; ** původně hlášen pod odlišným jménem Sivas Yozgat Rhinolophus ferrumequinum Rhinolophus hipposideros Rhinolophus euryale Rhinolophus mehelyi Myotis blythii Myotis myotis Myotis capaccinii Myotis aurascens Myotis brandtii Myotis mystacinus s. l. Eptesicus serotinus Hypsugo savii Pipistrellus kuhlii Pipistrellus pipistrellus s. l. Plecotus macrobullaris Plecotus teneriffae Barbastella cf. barbastellus Miniopterus schreibersii Tadarida teniotis species \ district Çankırı KayseriKırıkkaleKırşehirNevşehir Niğde + + – – + – – – – + + – – + – – – – – * * – – * * * – – – + – + – ** ** – * + + – – + – – – – – – – – – * – – – – – + – – + + + – – – – – – – * – – – – – * – – – + + * – – – – – – + ** – + * – * + – – + + – * – – * * – * ** * – + – + * + – + – – – – – + – – – * – – – + * – – – + * – – * – + – – – – – – – – total 6 11 3 5 8 11 7 5 2005: 1 ad. female (lact.) (ZDNU 2005/43); Gümüşler, Epcik Cave, 12 August 1999: 1 sad. male, 1 sad. & 1 ad. females (ZDNU 1999/28–30); 29 July 2002: 2 ad. males, 4 ad. females (ZDNU 2002/104–109), 3 males, 6 females (released) (cf. Karataş & Sözen 2004). Tadarida teniotis (Rafinesque, 1814) Sivas: Zâra, 9–10 August 2003: min. 20–50 ind. (obs.). RESULTS AND Discussion Rhinolophus ferrumequinum is the most frequently recorded species in Turkey and has been recorded in all eight provinces in the study area. Another abundant species is Myotis blythii which has been recorded in seven provinces in the study area but not in the Kırşehir province (cf. Benda & Horáček 1998). We did not find Myotis mystacinus s. l. which was recorded at the Devrez River (Çankırı: Ilgaz) by von Helversen (1989). Benda & Tsytsulina (2000) and Benda & Karataş (2005) revised the mystacinus group and stated that this record might be incorrect and most likely pertained to M. aurascens. According to these studies, M. mystacinus s. str. is very rare in Anatolia. The closest record to the study area was made near Kızılcahamam which is situated close to the northeastern part of the area (Benda & Karataş 2005). 155 Table 2. Comparison of occurrence of bat species in Turkey with respect to geographic regions (Central Anatolia is divided into two subregions, the eastern part [= the area under study] and the western parts) Tab. 2. Srovnání výskytu netopýřích druhů v Turecku podle zeměpisných oblastí (střední Anatolie je rozdělena na dvě podoblasti, východní [= studované území] a západní) species \ region Mediter- Aegean Marmara Black E SE CW CE ranean Sea AnatoliaAnatolia AnatoliaAnatolia Rousettus aegyptiacus Taphozous nudiventris Rhinolophus ferrumequinum Rhinolophus hipposideros Rhinolophus euryale Rhinolophus blasii Rhinolophus mehelyi Myotis bechsteinii Myotis myotis Myotis blythii Myotis nattereri Myotis emarginatus Myotis mystacinus Myotis aurascens Myotis nipalensis Myotis brandtii Myotis capaccinii Myotis daubentonii Nyctalus noctula Nyctalus leisleri Nyctalus lasiopterus Eptesicus serotinus Eptesicus bottae Vespertilio murinus Pipistrellus pipistrellus Pipistrellus pygmaeus Pipistrellus kuhlii Pipistrellus nathusii Hypsugo savii Plecotus auritus Plecotus macrobullaris Plecotus austriacus Plecotus teneriffae Barbastella barbastellus Barbastella cf. barbastellus Otonycteris hemprichii Miniopterus schreibersii Tadarida teniotis 1 0 1 1 1 1 1 1 1 1 1 1 1? 1 0 0 1 0 1 1 0 1 1 0 1 0 1 1 1 0 0 0 1 0 0 0 1 1 0 0 1 1 1 1 1 0 1 1 1 1 1? 1 0 0 1 0 0 0 0 1 1 0 1 0 ? 0 1 0 0 0 1 0 0 0 1 1 0 0 1 1 1 1 1 1 1 1 1 1 1 0 0 0 1 1 1 1 1 1 0 0 1 1 1 1 1 1 0 1 0 1 0 0 1 1 0 0 1 1 1 1 1 1 1 1 1 1 1 1 0 1 1 1 0 1 1 1 0 1 1 0 1 0 1 1 1 0 0 1 0 0 1 1 0 0 1 1 1? 0 1 0 1 1 1 0 1? 1 1 0 1 0 0 0 0 1 0 1 1 0 1 1 1 ? 1 0 0 0 0 0 1 1 0 1 1 1 1 0 1 0 1 1 1 1 1? 0 0 0 1 0 1 0 0 1 1 1 1 0 1 0 1 0 0 0 0 0 1 1 1 1 0 0 1 1 1? 0 1 0 1 1 0 0 1 1 0 0 0 0 0 0 0 1 0 0 1 0 0 0 1 0 1 0 1 0 0 0 1 1 0 0 1 1 0 0 1 0 1 1 0 0 1? 1 0 1 1 0 0 0 0 1 0 0 1 0 1 0 1 0 1 0 1 0 1 0 1 1 Benda & Horáček (1998) recorded Barbastella cf. barbastellus near Ürgüp (Nevşehir), howe ver, they released the specimen without examining it in detail. It is not clear whether this bat was the boreal B. barbastellus or the steppe B. leucomelas. The Ürgüp area is a steppe region and so it is most likely that it was B. leucomelas. 156 Moreover, Rhinolophus euryale recorded at Zâra (Sivas), and R. mehelyi from Keskin (Kırıkkale) and from the Seyfe Lake (Kırşehir) (see Benda & Horáček 1998) were not found by us in these regions. On the other hand, Myotis brandtii, which had been recorded only in Rize in Turkey, was recorded surprisingly in the steppe Yozgat province (cf. Benda & Karataş 2005). Hypsugo savii, Myotis aurascens, M. capaccinii and Plecotus teneriffae were also recorded for the first time in the study area. Additional specimens of M. capaccinii collected in the study area clearly show that this species is present not only in the Mediterranean (Akdeniz) and Marmara regions but also in suitable habitats along the Kızılırmak River in central Anatolia. Previously, western Palaearctic bats of the genus Plecotus had been included in P. auritus and P. austriacus. Similarly, Plecotus in central Anatolia had been considered to be P. auritus and P. austriacus (see Benda & Horáček 1998, Karataş et al. 2003). The genus Plecotus has been recently revised by Spitzenberger et al. (2006) and some new species have been described (see Spitzenberger et al. 2001, 2002, 2006, Benda et al. 2004, Juste et al. 2004). After that, the taxonomic position of Plecotus bats in central Anatolia became controversial. According to Spitzenberger et al. (2006), the species present in central Anatolia is P. macrobullaris (Fig. 2), while P. auritus may have a limited distribution area in the eastern Black Sea (Karadeniz) region and P. austriacus in Turkish Thrace. Moreover, according to DNA analysis of the specimens from Karaman, which is situated in the southeastern part of the study area, P. teneriffae is also found in the area (Juste et al. 2004). Fig. 2. Plecotus macrobullaris in a artificial cave at Talas, Başakpınar (Kayseri Prov.) . Obr. 2. Podobnost východní části střední Anatolie podle fauny netopýrů. 157 Fig. 3. Relationship of the eastern part of Central Anatolia with respect to bat fauna. Obr. 3. Podobnost východní části střední Anatolie podle fauny netopýrů. This study shows that 3 of 5 Turkish bat families (60%), 9 of 14 genera (64%), and 19 of 37 species (51.4%) are present in this area (Table 2). According to the analysis using the MVSP 3.1 software (Multivariate Statistical Package, Kovach Computing Services), the species diversity of the eastern part of Central Anatolia mostly resembles that of the western part of the region. It seems that the area shows zoogeographic affinity with Eastern Anatolia (Fig. 3). The results of the present study are consistent with those given by Benda & Horáček (1998). Based on the data on distribution of bats and the above mentioned analysis, the following four zoogeographic zones can be distinguished in Turkey. 1. Mediterranean (Akdeniz) zone: Akdeniz, Ege regions (except for the inner side), western part of the Marmara region with Mediterranean climate; 2. Central Anatolian steppe zone: eastern part of Ege, Central Anatolia, Eastern and Southeastern Anatolia (except for lowlands near the Syrian border) with continental climate; 3. Black Sea (Karadeniz) boreal zone: east of Marmara, Karadeniz, Erzurum and Kars plateau of Eastern Anatolia with very humid climate; 4. Southeastern Anatolian eremial zone: lowlands along the Syrian border. SOUHRN V uvedeném přehledu předkládáme nálezy 15 druhů netopýrů v oblasti středního a horního toku řeky Kizilirmak ve střední Anatolii (Turecko). Čtyři z těchto druhů (Myotis capaccinii, M. aurascens, M. brandtii, Hypsugo savii) byly nalezeny v uvedeném regionu poprvé, zatímco čtyři další druhy netopýrů zaznamenané dřívě nalezeny nebyly. Předložené výsledky zvyšily známý počet netopýrů oblasti na 19. Taxonomická posice netopýrů rodu Plecotus, která je v současné době šířeji diskutována, byla vyhodnocena 158 i ve studovaném regionu; nalezen byl druh P. teneriffae a poprvé tak zaznamenán z regionu pod tímto jménem. Z osmi provincií (vilájetů) střední Anatolie dvě vykazují bohatší faunu netopýrů (11 druhů), Niğde a Kayseri. Acknowledgements We wish to thank M. Ak, H. Baştürk, S. Baştürk, H. Bİlgen, Dr. R. Bİlgİn, K. Doğan, N. İpek, H. Ka rakaya, Y. Karakaya, Dr. Ay. Karataş, O. Kizilcik, Ş. Özkurt, H. C. Öztekİn, F. Telli, K. Ünsal, M. A. Yİğİt, and H. Yurtseven for their help with collecting data and material. A part of this study was supported by the Research Fund of Niğde University (Nr. 01/FEB/023). We dedicate this study to the memory of the Late Res. Assist. Kuddusi Ünsal. References Benda P. & Horáček I., 1998: Bats (Mammalia: Chiroptera) of the Eastern Mediterranean. Part 1. Review of distribution and taxonomy of bats in Turkey. Acta Soc. Zool. Bohem., 62: 255–313. Benda P. & Karataş A., 2005: On some Mediterranean populations of bats of the Myotis mystacinus morpho-group (Chiroptera: Vespertilionidae). Lynx, n. s., 36: 9–38. Benda P. & Tsytsulina K., 2000: Taxonomic revision of Myotis mystacinus group (Mammalia: Chiroptera) in the Western Palearctic. Acta Soc. Zool. Bohem., 64: 331–398. Benda P., Kiefer A., Hanák V. & Veith M., 2004: Systematic status of African populations of long-eared bats, genus Plecotus (Mammalia: Chiroptera). Folia Zool., 53, Monograph 1: 1–48. von Helversen O., 1989: New records of bats (Chiroptera) from Turkey. Zool. Middle East, 3: 5–18. Juste J., Ibáñez C., Muñoz J., Trujillo D., Benda P., Karataş A. & Ruedi M., 2004: Mitochondrial phy logeography of the Long-eared bats (Plecotus) in the Mediterranean Palaearctic and Atlantic Islands. Mol. Phylogenet. Evol., 31: 1114–1126. Karataş A. & Sözen M., 2004: Contributions to karyology, distribution and taxonomic status of the Long-winged bat, Miniopterus schreibersii (Chiroptera: Vespertilionidae), in Turkey. Zool. Middle East, 33: 51–64. Karataş A., Benda P., Toprak, F. & Karakaya H., 2003: New and significant records of Myotis capaccinii (Chiroptera: Vespertilionidae) from Turkey, with some data on its biology. Lynx, n. s., 34: 39–46. Spitzenberger F., Piálek J. & Haring E., 2001: Systematics of the genus Plecotus (Mammalia, Vespertili onidae) in Austria based on morphometric and molecular investigations. Folia Zool., 50: 161–172. Spitzenberger F., Haring E. & Tvrtković N., 2002: Plecotus microdontus (Mammalia, Vespertilionidae), a new bat species from Austria. Natura Croatica, 11: 1–18. Spitzenberger F., Strelkov P. P., Winkler H. & Haring E., 2006: A preliminary revision of the genus Plecotus (Chiroptera, Vespertilionidae) based on genetic and morphological results. Zool. Scripta, 35: 187–230. 159 Lynx (Praha), n. s., 37: 161–172 (2006). ISSN 0024–7774 Jan Svatopluk Presl’s (1821) family-group names of mammals Jména savců skupiny čeledi vytvořená Janem Svatoplukem Preslem (1823) Jiří Mlíkovský Department of Zoology, National Museum (Natural History), Václavské náměstí 68, CZ–115 79 Praha 1, Czech Republic; jiri.mlikovsky@nm.cz received on 5 September 2006 Abstract. Jan Svatopluk Presl created in 1821 eight family-group names of mammals, the existence of which has been overlooked by modern mammalogists. Three of them were not listed in subsequent catalo gues (Catadontidae, Hypsiprymnidae and Lipuridae), while the remaining five were generally attributed to various junior authors in modern literature (Myrmecophagidae, Petauridae, Chironectidae, Georychidae, and Thylacidae). In addition, the family-group name Phyllostomatidae, also used by Presl (1821), was found to be attributable to Goldfuss (1820), not to Gray (1825) as generally believed. Introduction Jan Svatopluk Presl (1791–1849) was a renowned Czech naturalist, whose scientific interests ranged from botany to mammalogy (Hoffmannová 1973). In 1821–1823 he published a syste matic synopsis of vertebrates (Presl 1821, 1822, 1823; see also Presl 1834). In this modernly structured classification he was one of the first naturalists to recognize the family level (cf. Mayr et al. 1953, Bock 1994). Prior to him, the family level was recognized and family-group names were properly formed in mammalogy only by a few authors, such as Karl Illiger (1775–1813) in 1811, Gotthelf Fischer von Waldheim (1771–1853) in 1817, Georg August Goldfuss (1782–1848) in 1820, Wilhelm Friedrich Hemprich (1798–1825) in 1820, and John Edward Gray (1800–1875) in 1821 (Illiger 1811, Fischer von Waldheim 1817, Goldfuss 1820, Hemprich 1820, Gray 1821). Presl’s classification of mammals (Presl 1821) was published in Czech language in the then newly founded, prestigious, Czech scientific journal ‘Krok’ (see Laiske 1959), which might have been the reason, why it has been overlooked by Palmer (1904) and subsequent students of mammalian family-group names (Simpson 1945, Wilson & Reeder 2005). Below I present an account of mammalian family-group names used by Presl (1821), with special respect to nomenclatural issues, which arose from their re-discovery. Family-group names are arranged alphabetically in the systematic part. Presl’s (1821) classification of mammals is given in full in the Appendix. 161 Systematic part Catodontidae Presl Presl (1821: 84) created family Catodonta for Catodon Linnaeus, 1761, which is a junior sub jective synonym of Physeter Linnaeus, 1758 (Mead & Brownell 2005). Catodonta Presl, 1821 is thus a junior subjective synonym of Physeteridae Gray, 1821 (I assume here that the paper by Gray was published earlier than that by Presl). Chironectidae Presl Palmer (1904: 734; see also Simpson 1935: 135; 1945: 41) attributed the family-group name Chironectidae to Anonymous (1897). The name was used already by Presl (1821: 80), who spelled it Cheironectina and based it on Chironectes Illiger, 1811. Cheironectes is a subsequent incorrect spelling of Chironectes Illiger, hence the family-group name should be corrected to Chironectina (ICZN 1999: Art. 35.4.). Water opossums of the genus Chironectes Illiger, 1811 are usually included in the subfamily Didelphinae of the family Didelphidae Gray, 1821 (Gardner 2005). Chironectina Presl, 1823 is thus a junior subjective synonym of Didelphidae Gray, 1821, but it is available if Chironectes opossums are separated at the subfamily level, as recently suggested by Hershkovitz (1997). Georychidae Presl Mole-rats of the genus Georychus Illiger, 1811 are usually included in the family Bathyergidae Waterhouse, 1841 (Simpson 1945, Woods & Kilpatrick 2005). Woods & Kilpatrick (2005: 1538) attributed the family-group name Georychidae to Roberts (1951), but already Simpson (1945: 99) traced the name back to Gravenhorst (1841: facing p. 502), who spelled it Georychina. The name should be attributed to Presl (1821: 81), who spelled it Georychina and based it on the genus Georychus Illiger, 1811. Georychina Presl, 1821 antedates Bathyergidae Waterhouse, 1843, which is the next oldest family-group name available for the family. Georychina Presl was not replaced by Bathyergi dae Waterhouse because it was based on a junior synonym (cf. ICZN 1999: Art. 40). However, Georychina Presl should be set aside, because Bathyergidae Waterhouse is in prevailing use and both conditions of Art. 23.9.1. (ICZN 1999) are met: the name has not been used as valid after 1899 (Art. 23.9.1.1.) and at least 10 authors used Bathyergidae Waterhouse as valid taxon in at least 25 works in immediately preceding 50 years in a period that encompasses over 10 years (Art. 23.9.1.2.). Required citations are as follows: Wood 1958, 1965, de Graff 1975, Jarvis 1978, Nevo 1979, Harvey et al. 1980, Nevo et al. 1987, Honeycutt et al. 1987, 1991, Bennett & Jarvis 1988, Denys 1989, Lovegrove 1989, 1991, Jarvis & Bennett 1990, 1991, Allard & Honeycutt 1992, Janeck et al. 1992, Burda & Kawalika 1993, Filipucci et al. 1994, Buffen stein 1996, Faulkes et al. 1997, McKenna & Bell 1997, Bennett & Faulkes 2000, Spinks et al. 2000, Walton et al. 2000, Oosthuizen et al. 2003. Hypsiprymnidae Presl Presl (1821: 80) based his family Hypsiprimnea [sic!] on the genus Hypsiprimnus, which is a subsequent incorrect spelling of Hypsiprymnus Illiger, 1811, The family-group name thus should be corrected to Hypsiprymnidae (ICZN 1999, Art. 35.4.). Hypsiprymnus Illiger, 1811 is 162 a junior objective synonym of Potorous Desmarest, 1804 (Groves 2005b). Hence, Hypsiprym nida Presl, 1821 is a junior objective synonym of Potoridae Gray, 1821 (I assume here that the paper by Gray was published earlier than that by Presl). Lipuridae Presl Presl (1821: 80) based his family Lipurina on the genus Lipurus Goldfuss, 1817, which is a junior subjective synonym of Phascolarctos Blainville, 1816 (Lee & Carrick 1989). Lipurina Presl, 1821 antedates Phascolarctidae created by Owen (1839), which is the next oldest family-group name available for the family. Lipurina Presl was not replaced by Phascolarcidae Owen because it was based on a junior synonym (cf. ICZN 1999, Art. 40). However, Lipurina Presl should be set aside, because Phascolarctidae Owen is in prevailing use and both conditions of Art. 23.9.1. (ICZN 1999) are met: the name has not been used as valid after 1899 (Art. 23.9.1.1.) and at least 10 authors used Phascolarctidae Owen as valid taxon in at least 25 works in immediately preceding 50 years in a period that encompasses over 10 years (Art. 23.9.1.2.). Required citations are as follows: Turnbull & Lundelius 1970, Imai et al. 1983, Strahan 1983, Nagy & Martin 1985, Aplin & Archer 1987, Haight & Nelson 1987, McKay 1988, Lee & Carrick 1989, Davidson & Young 1990, Harding & Aplin 1990, Gordon 1991, Luckett 1994, Osawa 1993, Szalay 1994, Retief et al. 1996, Black & Archer 1997, Kirsch et al. 1997, Kemper et al. 2000, Sherwin et al. 2000, Fisher et al. 2001, Grand & Barboza 2001, Cardillo et al. 2004, Kavanagh et al. 2004, Archer & Kirsch 2006, Munemasa et al. 2006, Weisbecker & Sánchez-Villagra 2006. Myrmecophagidae Presl The family-group name Myrmecophagidae is widely used and generally attributed to Gray (1825b: 343) (e.g. Gardner 2005: 102). It should be attributed to Presl (1821: 82), who based it on the genus Myrmecophaga Linnaeus, 1758. Petauridae Presl Groves (2005b: 53) attributed the family-group name Petauridae to Bonaparte (1838: 112). It should be attributed to Presl (1821: 79), who spelled it Petaurina and based it on the genus Petaurus Shaw, 1791. Phyllostomatidae Goldfuss Simmons (2005: 395) attributed the family-group name Phyllostomidae [sic!] to Gray (1825a: 242), although most authors attributed it to Gray (1825b: 338). Relative priority of these publications is irrelevant, however, because the name was used prior to Gray (1825a, b) already by Presl (1821: 78) and Goldfuss (1820: 460), of whom the latter spelled it Phyllostomata and based it on the genus Phyllostoma “Geoffr[oy Saint-Hilaire]” = Cuvier, 1800, which is a junior objective synonym of Phyllostomus Lacépède, 1799 (Hemming 1955). The family-group name Phyllostomatidae thus should be attributed to Goldfuss (1820). The family was generally spelled Phyllostomatidae until Kuzjakin (1974) and Handley (1980) argued that the name should be spelled Phyllostomidae. However, this is linguistically incorrect. Following Article 29.3.1. of the International Code of Zoological Nomenclature (ICZN 1999), “the stem for the purposes of the Code is found by deleting the case ending of the appropriate 163 genitive singular”. Here, the genitive singular of ‘stoma’ is ‘stomatis’, and the family-group name thus should be spelled Phyllostomatidae (see also Keržner 1974). Thylacidae Presl Presl (1821: 80) based his family Thylacina on the genus Thylacis Illiger, 1811. The latter genus is generally included in the family Peramelidae Gray, 1825 (Simpson 1945, Grover 2005a). Thylacina Presl, 1821 antedates Peramelidae Gray, 1825, which is the next oldest family-group name available for the family. Thylacina Presl was not replaced by Peramelidae Gray because it was based on a junior synonym (cf. ICZN 1999, Art. 40). However, Thylacina Presl should be set aside, because Peramelidae Gray is in prevailing use and both conditions of Art. 23.9.1. (ICZN 1999) are met: the name has not been used as valid after 1899 (Art. 23.9.1.1.) and at least 10 authors used Peramelidae Gray as valid taxon in at least 25 works in immediately preceding 50 years in a period that encompasses over 10 years (Art. 23.9.1.2.). Required citations are as follows: Lidicker & Follett 1968, Turnbull & Lundelius 1970, Archer & Kirsch 1977, 2006, Vaughan 1978, Stoddart & Braithwaite 1979, Gemmell 1982, Szalay 1982, Strahan 1983, Aplin & Archer 1987, Gordon & Hulbert 1989, Friend 1990, Groves & Flannery 1990, Kem per et al. 1990, Southgate 1990, Menzies 1991, Sherwin et al. 1991, Murphy & Serena 1993, Retief et al. 1995, McKenna & Bell 1997, Short et al. 1998, Muirhead 2000, Westermann et al. 2001, Broughton & Dickman 2002, Chambers & Dickman 2002, Richards & Short 2003, Price 2004, Westerman et al. 2004. Souhrn Jan Svatopluk Presl vytvořil roku 1821 ve své klasifikaci savců osm nových jmen skupiny čeledi. Z nich tři jména vůbec nebyla zahrnuta do pozdějších katalogů (Catadontidae, Hypsiprymnidae a Lipuridae), zatímco autorství ostatních pěti jmen bylo běžně připisováno různým mladším autorům (Myrmecopha gidae, Petauridae, Chironectidae, Georychidae a Thylacidae). Kromě toho bylo zjištěno, že jméno Phyl lostomatidae, zpravidla připisované Grayovi (1825a, b) a taktéž použité Preslem (1821), bylo vytvořeno již Goldfussem (1820). Acknowledgments I am obliged to Petr Benda (Praha) for comments on the manuscript. The study was supported in part by the grants of the Ministry of Culture of the Czech Republic No. DE06P04OMG008 and MK00002327201. References Allard M. W. & Honeycutt R. L., 1992: Nucleotide sequence variation in the mitochondrial 12s rRNA gene and the phylogeny of African mole-rats (Rodentia: Bathyergidae). Molecular Biology and Evolu tion, 9: 27–40. Anděra M., 1999: České názvy živočichů. II. Savci (Mammalia) [Czech Names of Animals. II. Mammals (Mammalia)]. Národní muzeum, Praha, 147 pp (in Czech). Anonymous, 1897: Verzeichnis der im Museum der Naturhistorischen Gesellschaft zu Hannover vorhan denen Säugetiere. Hannover, 30 pp. Aplin K. P. & Archer M., 1987: Recent advances in marsupial systematics with new syncretic classificati on. Pp.: xv–lxi. In: Archer M. (ed.): Possums and Opossums: Studies in Evolution. Surrey Beatty and Sons, Chipping Norton, lxxii+788 pp. 164 Archer M. & Kirsch A. W., 1977: The case for the Thylacomyidae and Myrmecobiidae, Gill, 1872, or why are marsupial families so extended. Proceedings of the Linnean Society of New South Wales, 102: 18–25. Archer M. & Kirsch A. W., 2006: The evolution and classification of marsupials. Pp.: 1–21. In: Ar mati P. J., Dickman C. R. & Hume I. D. (eds.): Marsupials. Cambridge University Press, Cambridge, xiii+373 pp. Bennett N. C. & Faulkes C. G., 2000: African Mole-rats: Ecology and Eusociality. Cambridge Univer sity Press, Cambridge, xiv+273 pp. Bennett N. C. & Jarvis J. U. M., 1988: The social structure and reproductive biology of colonies of the mole-rat, Cryptomys damarensis (Rodentia, Bathyergidae). Journal of Mammalogy, 69: 293–302. Black K. & Archer M., 1997: Nimiokoala gen nov. (Marsupialia, Phascolarctidae) from Riversleigh, northwestern Queensland, with a revision of Litokoala. Memoirs of the Queensland Museum, 41(2): 209–228. Bock W. J., 1994: History and nomenclature of avian family-group names. Bulletin of the American Mu seum of Natural History, 222: 1–281. Bonaparte C. L, 1838: Synopsis vertebratorum systematis. Nuovi Annali delle Scienze Naturali, 2(1): 105–133. Broughton S. K. & Dickman C. R., 1991: The effect of supplementary food on home range of the southern brown bandicoot, Isoodon obesulus (Marsupialia: Peramelidae). Australian Journal of Ecology, 16: 71–78. Buffenstein R., 1996: Ecophysiological responses to a subterranean habitat – a bathyergid perspective. Mammalia, 60: 591–605. Burda H. & Kawalika M., 1993: Evolution of eusociality in the Bathyergidae. The case of the giant mole rats (Cryptomys mechowi). Naturwissenschaften, 80: 235–237. Cardillo M., Bininda-Emonds O. R. P., Boakes E. & Purvis A., 2004: A species-level phylogenetic supertree of marsupials. Journal of Zoology, London, 264: 11–31. Chambers L. K. & Dickman C. R., 2002: Habitat preferences of the long-nosed bandicoot, Perameles nasuta (Mammalia, Peramelidae), in a patchy, urban environment. Australian Ecology, 27: 334–342. Davidson C. V. & Young W. G., 1990: The muscles of mastication of Phascolarctos cinereus (Phascolar ctidae: Marsupialia). Australian Journal of Zoology, 38: 227–240. de Graaff G., 1975: Family Bathyergidae. Pp.: 1–5. In: Meester J. &Setzer H. W. (eds.): The Mammals of Africa: An Identification Manual. Part 6.9. Smithsonian Institution Press, Washington, DC, not con tinuously paginated. Denys C., 1989: A new species of bathyergid rodent from Olduvai Bed I (Tanzania, Lower Pleistocene). Neue Jahrbücher Geologie Palaeontologie, 5: 257–264. Faulkes C. G., Bennett N. C., Bruford M. W., O’Brien H. P., Aguilar G. H. & Jarvis J. U. M., 1997: Ecological constraints drive social evolution in the African mole-rats. Proceedings: Biological Sciences, 264: 1619–1627. Filipucci M. G., Burda H., Nevo E. & Kocka J., 1994: Allozyme divergence and systematics of common mole-rats (Cryptomys, Bathyergidae, Rodentia) from Zambia. Zeitschrift für Säugetierkunde, 59: 42–51. Fischer D. O., Owens I. P. F. & Johnson C. N., 2001: The ecological basis of life history variation in mar supials. Ecology, 82: 3531–3540. Fischer von Waldheim G., 1817: Adversaria zoologica. Memoires de la Société Impériale des Naturalistes de Moscou, 5: 368–428. Friend G. R., 1990: Breeding and population dynamics of Isoodon macrourus (Marsupialia: Peramelidae): studies from the wet-dry tropics of northern Australia. Pp.: 357–365. In: Seebeck J. H., Brown P. H., Wallis R. L. & Kemper C. L. (eds.): Bandicoots and Bilbies. Surrey Beatty & Sons, Chipping Norton, NSW, xxx+392 pp. Gardner A. L., 2005: Order Didelphimorphia. Pp.: 3–18. In: Wilson D. E. & Reeder D. (eds.): Mammal Species of the World. Third Edition. The John Hopkins University Press, Baltimore, 2142 pp. 165 Gemmell R. T., 1982: Breeding bandicoots in Brisbane (Isoodon macrourus; Marsupialia, Peramelidae). Australian Mammalogy, 5: 187–193. G oldfuss G. A., 1820: Handbuch der Zoologie. Vol. 3(2). Johann Leonard Schrag, Nürnberg, xxiv+506 pp. Gordon G., 1991: Estimation of age of the koala, Phascolarctos cinereus (Marsupialia: Phascolarctidae) from tooth wear and growth. Australian Mammalogy, 14: 5–12. Gordon G. & Hulbert H. J., 1989: Peramelidae. Pp.: 603–624. In: Walton D. W. (ed.): Fauna of Austra lia. Volume 1B. Mammalia. Australian Government Publishing Service, Canberra, 827 pp. Grand T. I. & Barboza P. S., 2001: Anatomy and development of the koala, Phascolarctos cinereus: an evolutionary perspective on the superfamily Vombatoidea. Anatomy and Embryology, 203: 211–223. Gravenhorst J. L. C., 1843: Vergleichende Zoologie. Grass, Barth und Co., Breslau, xx+686 pp. Gray J. E., 1821: On the natural arrangement of vertebrose animals. London Medical Repository, 15: 296–310. Gray J. E., 1825a: An attempt at a division of the family Vespertilionidae into groups. Zoological Journal, 2: 242–243. Gray J. E., 1825b: Outline of an attempt at the disposition of the Mammalia into tribes and families with a list of the genera apparently appertaining to each tribe. Annals of Philosophy, (2)10: 337–344. Groves C. P., 2005a: Order Peramelemorphia. Pp.: 38–42. In: Wilson D. E. & Reeder D. (eds.): Mammal Species of the World. Third Edition. The John Hopkins University Press, Baltimore, 2142 pp. Groves C. P., 2005b: Order Diprotodontia. Pp.: 43–70. In: Wilson D. E. & Reeder D. (eds.): Mammal Species of the World. Third Edition. The John Hopkins University Press, Baltimore, 2142 pp. Groves C. P. & Flannery T. F., 1990: Revision of the families and genera of bandicoots. Pp.: 1–11. In: Seebeck J. H., Brown P. H., Wallis R. L. & Kemper C. L. (eds.): Bandicoots and Bilbies. Surrey Beatty & Sons, Chipping Norton, NSW, xxx+392 pp. Haight J. R. & Nelson J. E., 1987: A brain that doesn’t fit its skull: A comparative study of the brain and endocranium of the koala, Phascolarctos cinereus (Marsupialia: Phascolarctidae). Pp.: 331–352. In: Archer M. (ed.): Possums and Opossums: Studies in Evolution. Surrey Beatty, Chipping Norton, NSW, 788 pp. Handley C. O., Jr., 1980: Inconsistencies in formation of family-group and subfamily-group names in Chiroptera. Pp.: 9–13. In: Wilson D. E. & Gardner A. L. (eds.): Proceedings of the Fifth International Bat Research Conference. Texas Tech University, Lubbock, 434 pp. Harding H. R . & Aplin K. P., 1990: Phylogenetic affinities of the koala (Phascolarctidae: Marsupialia): a reassessment of the spermatozoal evidence. Pp.: 1–13. In Lee A. K., Handasyde K. A. & Sanson G. D. (eds.): Biology of the Koala. Surrey Beatty and Sons, Sydney, 788 pp. Harvey P. H., Clutton-Brock T. H. & Mace G. M., 1980: Brain size and ecology in small mammals and primates. Proceedings of the National Academy of Sciences of the United States of America (Biological Sciences), 77: 4387–4389. Hemming F., (ed.) 1955: Direction 24: completion of the entries relating to the names of certain genera in the class Mammalia made on the Official List of Generic Names in Zoology in the period up to the end of 1936. Opinions and Declarations Rendered by the International Commission on Zoological Nomenclature, 3: 219–246. Hemprich W. F., 1820: Grundriß der Naturgeschichte für höhere Lehranstalten. August Rücker, Berlin, & Friedrich Volke, Wien, 432 pp. Hershkovitz P., 1997: Composition of the family Didelphidae Gray, 1821 (Didelphoidea: Marsupialia), with a review of the morphology and behavior of the included four-eyed pouched opossums of the genus Philander Tiedemann, 1808. Fieldiana: Zoology (New Series), 86: 1–103. Hoffmannová E., 1973: Jan Svatopluk Presl, Karel Bořivoj Presl. Melantrich, Praha, 299 pp (in Czech). Honeycutt R. L., Edwards S. V., Nelson K. & Nevo E., 1987: Mitochondrial DNA Variation and the Phy logeny of African Mole Rats (Rodentia: Bathyergidae). Systematic Zoology, 36: 280–292. 166 Honeycutt R. L., Allard M. W., Edwards S. V. & Schlitter D. A., 1991: Systematics and evolution of the family Bathyergidae. Pp.: 45–65. In: Sherman P. W., Jarvis J. U. M. & Alexander R. D. (eds.): The Biology of the Naked Mole-rat. Princeton University Press, Princeton, 518 pp. ICZN, 1999: International Code of Zoological Nomenclature. 4th ed. International Trust for Zoological Nomenclature, London, xxix+306 pp. Illiger C., 1811: Prodromus systematis mammalium et avium additis terminis zoographicis utriusque classis, eorumque versione germanica. C. Salfeld, Berlin, 302 pp. Imai H. T., Maruyama T. & Croizer T. H., 1983 : Rates of mammalian karyotype evolution by the karyo graph method. American Naturalist, 121: 477–488. Janeck L. L., Honeycutt R. L., Rautenbach I. L., Erasmus B. H. & Schlitter D. A., 1992: Allozyme variation and systematics of African mole-rats (Rodentia: Bathyergidae). Biochemical Systematics and Ecology, 20: 401–416. Jarvis J. U. M., 1978: Energetics of survival in Heterocephalus glaber (Rüppell) the naked mole-rat (Ro dentia: Bathyergidae). Bulletin of the Carnegie Museum of Natural History, 6: 81–87. Jarvis J. U. & Bennett N. C., 1990: The evolutionary history, population biology and social structure of African mole-rats: family Bathyergidae. Progress in Clinical and Biological Research, 335: 97–128. Jarvis, J. U. M. & Bennett N., 1991: Ecology and behavior of the family Bathyergidae. Pp.: 66–96. In: Sherman P. W., Jarvis J. U. M. & Alexander R. (eds.): The Biology of the Naked Mole-rat. Princeton: Princeton University Press, xvi+518 pp. Kavanagh J. R., Burk-Herrick A., Westerman M. & Springer M. S., 2004: Relationships among families of Diprotodontia (Marsupialia) and the phylogenetic position of the autapomorphic honey possum (Tarsipes rostratus). Journal of Mammalian Evolution, 11: 207–222. Kemper C., Kitchener D. J., Humphreys W. F., How R. A., Schmitt L. H. & Bradley A. 1990: The biology of the northern brown bandicoot, Isoodon macrourus (Marsupialia: Peramelidae) at Mitchell Plateau, Western Australia. Australian Journal of Zoology, 37: 627–644. Kemper C., Reardon T. & Queale L., 2000: Mammals. Pp.: 13–35. In: Robinson A. C., Casperson K. D. & Hutchinson M. N. (eds.): A List of the Vertebrates of South Australia. 3rd ed. Department for Envi ronment and Heritage, Adelaide, 146 pp. Keržner I. M., 1974: Zamečanija k stat’e A. P. Kuzjakina “O nazvanijach rukokrylych” [Remarks on the paper by A. P. Kuzjakin “On the names of bats”]. P.: 49. In: Strelkov P. P. & Kuzjakin A. P. (eds.): Mate rialy pervogo vsesojuznogo soveščanija po rukokrylym (Chiroptera) [Proceedings of the First All-Union Meeting on Bats (Chiroptera)]. Zoologičeskij Institut AN SSSR, Leningrad, 183 pp (in Russian). Kirsch J. A. W., Lapointe F.-J. & Springer M. S., 1997: DNA-hybridisation studies of marsupials and their implication for Metatherian classification. Australian Journal of Zoology, 45: 211–280. Kuzjakin A. P., 1974: O nazvanijah rukokrylyh [On the names of bats]. Pp.: 45–49. In: Strelkov P. P. & Kuzjakin A. P. (eds.): Materialy pervogo vsesojuznogo soveščanija po rukokrylym (Chiroptera) [Proceedings of the First All-Union Meeting on Bats (Chiroptera)]. Zoologičeskij Institut AN SSSR, Leningrad, 183 pp (in Russian). Laiske M., 1959: Časopisectví v Čechách 1650–1847 [Journalism in Bohemia 1650–1847]. Národní knihovna, Praha, 180 pp (in Czech). Lee A. K. & Carrick F. N., 1989: Phascolarctidae. Pp.: 740–754. In: Walton D. W. & Richardson B. J. (eds.): Fauna of Australia. Vol. 1B. Mammalia. Australian Government Publishing Service, Canberra, 827 pp. Lidicker Jr., W. Z. & Follett W. I., 1968: Isoodon Desmarest, 1817, rather than Thylacis Illiger, 1811, as the valid generic name of the short-nosed bandicoots (Marsupialia: Peramelidae). Proceedings of the Biological Society of Washington, 81: 251–256. Lovegrove B. G., 1989: The cost of burrowing by the social mole-rats (Bathyergidae) Cryptomys dama rensis and Heterocephalus glaber: the role of soil moisture. Physiological Zoology, 62: 449–469. Lovegrove B. G., 1991: The evolution of eusociality in molerats (Bathyergidae): a question of risks, numbers, and costs. Behavioral Ecology and Sociobiology, 28: 37–45. 167 Luckett W. P., 1994: Suprafamilial relationships within Marsupialia: resolution and discordance from multidisciplinary data. Journal of Mammalian Evolution, 2: 255–283. Mayr E., Linsley E. G. & Usinger R. L., 1953: Methods and Principles of Systematic Zoology. McGraw-Hill Book Company, New York, ix+336 pp. McKay G. M., 1988: Phascolarctidae. Pp.: 51–52. In: Walton D. W. (ed.): Zoological Catalogue of Aus tralia. Vol. 5. Mammalia. Australian Government Publishing Service, Canberra, 274 pp. McKenna M. C. & Bell S. K., 1997: Classification of Mammals Above the Species Level. Columbia University Press, New York, 631 pp. Mead J. G. & Brownell Jr., R. L., 2005: Order Cetacea. Pp.: 723–743. In: Wilson D. E. & Reeder D. (eds.): Mammal Species of the World. Third Edition. The John Hopkins University Press, Baltimore, 2142 pp. Menzies J., 1991: A Handbook of New Guinea Marsupials and Monotremes. Kristen Pres, Madang, 138 pp. Muirhead J., 2000: Yaraloidea (Marsupialia, Perameleformia), a new superfamily of marsupial and a description and analysis of the cranium of the Miocene Yarala burchfieldi. Journal of Paleontology, 74: 512–523. Munemasa M., Nikaido M., Donnellan S., Austin C. C., Okada N. & Hasegawa M., 2006: Phylogenetic analysis of diprotodontian marsupials based on complete mitochondrial genomes. Genes & Genetic Systems, 81: 181–191. Murphy J. A. & Serena M., 1993: Results of radio tracking eastern barred bandicoots, Perameles gunnii, (Marsupialia: Peramelidae) at Gellibrand Hill Park, Victoria. Australian Mammalogy, 16: 51–54. Nagy K. A. & Martin R. W., 1985: Field metabolic rate, water flux, food consumption and time budget of koalas, Phascolarctos cinereus (Marsupialia: Phascolarctidae) in Victoria. Australian Journal of Zoology, 33: 655–665. Nevo E., 1979: Adaptive convergence and divergence in subterranean mammals. Annual Review of Eco logy and Systematics, 10: 269–308. Nevo E., Ben-Shlomo R., Beiles A., Jarvis J. U. M. & Hickman G. C., 1987: Allozyme differentiation and systematics of the endemic subterranean mole rats of Africa (Rodentia, Bathyergidae). Biochemical Systematics and Ecology, 15: 409–502. Oosthuizen M. K., Cooper H. M. & Bennett N. C., 2003: Circadian rhythms of locomotor activity in solitary and social species of African mole-rats (Family: Bathyergidae). Journal of Biological Rhythms, 18: 481–490. Osawa R., 1993: Dietary preference of koalas Phascolarctos cinereus (Marsupialia: Phascolarctidae) for Eucalyptus spp. with a specific reference to their simple sugar contents. Australian Mammalogy, 16: 85–87. Owen R., 1839: Outlines of a classification of the Marsupialia. Proceedings of the Zoological Society of London, 1839: 5–19. Palmer T. S., 1904: Index generum mammalium: a list of the genera and families of mammals. United States Department of Agriculture, North American Fauna, 23: 1–984. Presl J. S., 1821: Nawržení saustawy žiwočichů dle tříd, řádů a rodů, a spolu pokus zčeštění potřebných w žiwočistwu názwů [Proposition of the system of animals, according to classes, orders and genera, with an attempt to creation of useful Czech names of animals]. Krok, 1(2): 67–84 (in Czech). Presl J. S., 1822: Nawržení saustawy žiwočichů dle tříd, řádů a rodů, a spolu pokus zčeštění potřebných w žiwočistwu názwů (Pokračowání.) [Proposition of the system of animals, according to classes, orders and genera, with an attempt to creation of useful Czech names of animals (continuation)]. Krok, 1(3): 116–129 (in Czech). Presl J. S., 1823: Nawržení saustawy žiwočichů dle tříd, řádů a rodů, a spolu pokus zčeštění potřebných w žiwočistwu názwů (Pokračowání) [Proposition of the system of animals, according to classes, orders and genera, with an attempt to creation of useful Czech names of animals (continuation)]. Krok, 1(4): 104–109 (in Czech). 168 Presl J. S., 1834: Ssawectwo [Mammals]. Kronberg a Webr, Praha, xii+388 pp (in Czech). Price G. J., 2004: Fossil bandicoots (Marsupialia, Peramelidae) and environmental change during the Pleistocene on the Darling Downs, southeastern Queensland, Australia. Journal of Systematic Paleon tology, 2: 347–356. Retief J. D., Krajewski C., Westerman M., Winkfein R. J. & Dixon G. H., 1995: Molecular phylogeny and evolution of marsupial protamine P1 genes. Proceedings: Biological Sciences, 259 7–14. Richards J. D. & Short J., 2003: Reintroduction and establishment of the western barred bandicoot Perameles bougainville (Marsupialia: Peramelidae) at Shark Bay, Western Australia. Biological Con servation, 109: 181–195. Roberts A., 1951: The Mammals of South Africa. Trustees of “The mammals of South Africa” Book Fund, Johannesburg, 700 pp. Sherwin W. B., Murray N. D., Marshall Graves J. A. & Brown P. R., 1991: Measurement of genetic variation in endangered populations: bandicoots (Marsupialia: Peramelidae) as an example. Conserva tion Biology, 5: 103–108. Sherwin W. B., Timms P., Wilcken J. & Houlden B., 2000: Analysis and conservation implication of koala genetics. Conservation Biology, 14: 639–649. Short J. C., Richards J. D. & Turner B., 1998: Ecology of the western barred bandicoot (Perameles bougainville) (Marsupialia: Peramelidae) on Dorre and Bernier Islands, Western Australia. Wildlife Research, 25: 567–586. Simmons N. B., 2005: Order Chiroptera. Pp.: 312–529. In: Wilson D. E. & Reeder D. (eds.): Mammal Species of the World. Third Edition. The John Hopkins University Press, Baltimore, 2142 pp. Simpson G. G., 1935: Note on the classification of recent and fossil opossums. Journal of Mammalogy, 16: 134–137. Simpson G. G., 1945: The principles of classification and a classification of mammals. Bulletin of the American Museum of Natural History, 85: i–xvi, 1–350. Southgate R. I., 1990: Distribution and abundance of the greater bilby Macrotis lagotis Reid (Marsupialia: Peramelidae). Pp.: 293–302. In: Seebeck J. H., Brown P. R., Wallis R. L. & Kemper C. M. (eds.): Bandicoots and Bilbies. Surrey Beatty & Sons, Chipping Norton, xxx+392 pp. Spinks A. C., Jarvis J. U. M. & Bennett N. C., 2000: Comparative pattern of philopatry and dispersal in two common mole-rat populations: implications on the evolution of mole-rat sociality. Journal of Animal Ecology, 69: 224–234. Stoddart D. M. & Braithwaite R. W., 1979: A strategy for utilization of regenerating heathland habitat by the brown bandicoot (Isoodon obesulus; Marsupialia, Peramelidae). Journal of Animal Ecology, 48: 165–179. Strahan R. (ed.), 1983: Complete Book of Australian Mammals. Magnus and Robertson Publ., London, xxi+530 pp. Szalay F. S., 1982: A new appraisal of marsupial phylogeny and classification. Pp.: 621–640. In: Archer M. (ed.): Carnivorous Marsupials. Royal Zoological Society of New South Walers, Mosman, vi+804 pp. Szalay F. S., 1994: Evolutionary History of the Marsupials and an Analysis of Osteological Characters. Cambridge University Press, Cambridge, xi+481 pp. Turnbull W. D. & Lundelius E. L., 1970: The Hamilton fauna. A late Pleistocene mammalian fauna from the Grange Burn, Victoria, Australia. Fieldiana: Geology, 19: 1–163. Vaughan T. A., 1978: Mammalogy. 2nd Edition. W. B. Saunders, Philadelphia, 522 pp. Walton A. H., Nedbal M. A. & Honeycutt R. L., 2000: Evidence from intron 1 of the nuclear transthyre tin (prealbumin) gene for the phylogeny of African mole-rats (Bathyergidae). Molecular Phylogenetics and Evolution, 16: 467–474. Waterhouse G. R., 1841: Observations on the Rodentia. Annals and Magazine of Natural History, 8: 81–84. Weisbecker V. & Sánchez-Villagra M. R., 2006: Carpal evolution in diprotodontian marsupials. Zoolo gical Journal of the Linnean Society, 146: 369–384. 169 Westermann M., Springer M. S. & Krajewski C., 2001: Molecular relationships of the New Guinean bandicoot genera Microperoryctes and Echymipera (Marsupialia: Peramelina). Journal of Mammalian Evolution, 8: 93–105. Westermann M., Springer M. S., Dixon J. & Krajewski C., 2004: Molecular relationships of the pig-footed bandicoot Chaeropus ecaudatus (Marsupialia: Perameloidea) using 12S rRNA sequences. Journal of Mammalian Evolution, 6: 271–288. Wilson D. E. & Reeder D. (eds.), 2005: Mammal Species of the World. Third Edition. The John Hopkins University Press, Baltimore, xxxviii+2142 pp. Wood A. E., 1958: Are there rodent suborders? Systematic Zoology, 7: 169–173. Wood A. E., 1965: Grades and clades among rodents. Evolution, 19: 115–130. Woods C. A. & Kilpatrick C. W., 2005: Infraorder Hystricognathi. Pp.: 1538–1600. In: Wilson D. E. & Reeder D. (eds.): Mammal Species of the World. Third Edition. The John Hopkins University Press, Baltimore, 2142 pp. Appendix Presl’s (1821) classification of mammals. Czech names proposed by Presl (1821) for mammali an orders, families and genera are given in parentheses (most were not listed by Anděra 1999). Note that Presl (1821) did not name families when an order included only a single family. 1. Bimana (dvaurucí) 1.1. [Family not given]: Homo (člowěk). 2. Quadrumana (čtwerorucí) 2.1. Simia (opicowití): Pithecus (op), Hylobates (ramenáč), Lasiopyga (dlakořit), Cercopithecus (koč kodán), Papio (martyška), Poago (mirkuwín), Cynocephalus (psohlaw), Colobus (kykatas), Ateles (chapan), Cebus (malpa), Pithecia (chwostan), Aotus (nočák), Callitrix (pěknowlasec), Hapale (kosman). 2.2. Prosimia (munowití): Lemur (muna), Lichanotus (požast), Stenops (autloň). 2.3. Prehensilia (palcuchowití): Chirogeleus (palcucha). 2.4. Macrotarsia (nártaunowití): Tarsius (nártaun), Otolicnus (uchoš). 3. Cheiroptera (letauni) 3.1. Galeopithecia (letuškowití): Galeopithecus (letuška). 3.2. Phyllostomata (řasonosowití): Phyllostoma (řasonos), Nycteris (šerowec), Rhinopoma (nosalec), Rhinolophus (wrapenec), Megaderma (weloblanec). 3.3. Harpiae (upírowití): Pteropus (upír), Cephalotes (hlawan). 3.4. Noctiliones (nedopírowití): Stenoderma (auzkoblanec), Vespertilio (netopír), Plecotus (ušan), Miopterus (spaka), Nyctionomus (příšerec), Noctilio (mračník), Dysopes (psohubec), Thapazous (wečer ník). 4. Ferae (šelmy) 4.1. Canina (psovití): Megalotis (welouch), Canis (pes), Hyena (hyena), Felis (kot), Rycaena (zeník). 4.2. Mustelina (kunowití): Herpestes (promyka), Mephitis (smrdoš), Mustela (kuna), Ichneumon (mun kos), Lutra (wydra). 4.3. Ursina (nedwědowití): Ursus (nedwěd), Meles (jezwec), Gulo (rosomak), Procyon (mýwal), Nasua (nosál), Cercoleptes (ohonowin). 4.4. Erinacea (ježovití): Centetes (bodlín), Erinaceus (jež). 4.5. Talpina (krtkowití): Condylura (uzloš), Chrysochloris (zlatokrt), Talpa (krt), Scalops (hrabuška). 4.6. Sorexina (reyskowití): Sorex (reysek), Mygale (wychuchol). 5. Marsupialia (waknatí) 5.1. Petaurina (letawcowití): Petaurus (létawec), Phalangista (parowník). 170 5.2. 5.3. 5.4. 5.5. 5.6. Cheironectina (plawákowití): Cheironectes (plawák). Didelphina (wačicowití): Ambliotis (ruhoš), Balantia (tokaun), Didelphis (wačice). Thylacina (torebníkowití): Dasyurus (srstaun), Thylacis (torebník). Lipurina (kwíkolowití): Phascolomys (drapoš), Lipurus (kwíkol). Hypsiprimnea (skokeyšowití): Hypsiprimnus (skokeyš), Halmaturus (klokan). 6. Rosores (hlodawí) 6.1. Dipudina (tarbíkowití): Dipus (tarbík), Pedetes (nohas). 6.2. Sciurina (wewerowití): Myoxus (plch), Tamias (deňka), Pteromys (poletucha), Sciurus (wewer), Cheiromis (letaha). 6.3. Leporina (zajícowití): Lepus (zajíc), Lagomys (pičuha). 6.4. Cavinida (morčowití): Coelogenys (tlamák), Dasyproctes (nahoš), Cavia (morče, also wiska), Hydrochaerus (plawaun). 6.5. Histricina (dikobrazowití): Histrix (dikobraz), Coëndu (ostnoš), Loncheres (ježowec). 6.6. Castorina (bobrowití): Hydromys (woduška), Guillino (dlakaun), Ondatra (ondatra), Castor (bobr). 6.7. Georychina (hrabošowití): Georychus (hraboš), Hypudaerus (pestruška), Fiber (dlakoš). 6.8. Murina (myšowití): Mus (myš), Cricetus (křeček), Arctomys (swišť), Viscacia (wizkacha), Spalax (slepec), Bathyergus (rypoš). 7. Cingulata (pásatí) 7.1. [Family not given]: Dasypus (pásowec), Tolipeutes (chaulan). 8. Pamphracta (Ssupani) 8.1. [Family not given]: Pamphractus (ssupan). 9. Ornithostomata (ptakohubí) 9.1. [Family not given]: Ornithorhynchus (ptakaun). 10. Tachiglossa (rychlozajiční) 10.1.[Family not given]: Tachyglossus (rychlojazan). 11. Vermilinguia (tenkojazyční) 11.1 Manisia (luskaunowití): Manis (luskaun). 11.2.Myrmecophagina (mrawenčíkowití): Myrmecophaga (mrawenčík), Tamandua (tamandua), Orycte ropus (kuťoš). 12. Tardigrada (lenochodi) 12.1.[Family not given]: Bradypus (lenochod), Choloepus (kulhoš). 13. Bisulca (dwaupaznehtní) 13.1.Taurina (beykowití): Bos (beyk), Ovibos (owoskot), Ovis (owce), Capra (kozel), Antilope (sajka). 13.2.Cervina (jelenowití): Cervus (jelen), Moschus (kabarha). 13.3.Girafina (girafowití): Camelopardalis (girafa). 13.4.Camelina (welblaudowití): Auchenia (wikuně), Camelus (welblaud). 14. Solidungula (jednopaznehtní) 14.1.[Family not given]: Equus (kůň). 15. Multungula (mnohopaznehtní) 15.1.Elephantina (slonowití): Elephas (slon). 15.2.Tapiracea (tapírowití): Tapirus (tapír). 15.3.Scrofina (wepřowití): Sus (wepř). 15.4.Hyracina (tlustošowití): Lipura (nehtaun), Hyrax (tlustoš). 15.5.Rhinocerina (rohošowití): Rhinoceros (rohoš). 15.6.Hyppopotamea (hrochowití): Hyppopotamus (hroch). 171 16. Pinnipedia (ploskonozí) 16.1.[Family not given]: Phoca (teleň), Pusa (siwuč), Nepus (ťutě); also nerpa, lachták. 17. Syrtobatica (smeykali) 17.1.[Family not given]: Trichechus (morž). 18. Sirenia (ochechule) 18.1.[Family not given]: Manatus (kapustňak), Halicore (moroň), Rytina (koraun). 19. Cetacea (welrybi) 19.1.Balaenacea (kytowití): Balaena (kyt), Balanopter (pleytwák). 19.2.Catodonta (worwaňowití): Physeter (perutoš), Oryx (zubaun), Cetus (olbroť), Catodon (worwaň), Delphinus (pliskawice), Delphinapterus (běluha), Physalus (sykawice). 172 Lynx (Praha), n. s., 37: 173–177 (2006). ISSN 0024–7774 Diet of the American mink (Mustela vison) in the Czech Republic (Carnivora: Mustelidae) Potrava norka amerického (Mustela vison) v České republice (Carnivora: Mustelidae) Michaela Nováková1,2 & Petr Koubek2 1 Institute of Botany and Zoology, Faculty of Science, Masaryk University, Kotlářská 2, CZ–611 37 Brno, Czech Republic; novakova@ivb.cz 2 Institute of Vertebrate Biology, Czech Academy of Science, Květná 8, CZ–603 65 Brno, Czech Republic; koubek@brno.cas.cz received on 28 June 2006 Abstract. An analysis of 29 full stomachs of the American mink (Mustela vison Schreber, 1777), collected in the Křivoklátsko Protected Landscape Area and in the Horažďovice region (Czech Republic) during autumn and winter time in 2001–2006, showed a great diversity of the mink diet. Fish (Cyprinidae, Percidae, and Esocidae) were the dominant component of their diet (51.7% of occurrence, 41.0% of volume), followed by mammals (Rodentia, Insectivora) (48.7% of occurrence, 31.1% of volume) and birds (20.7% of occurrence, 13.2% of volume). Amphibians, insects, carrions and plants served only as supplementary food. Mink was found to be a typical generalist predator. Mink diet is, most probably, influenced by prey availability, prey behaviour, season, prey abundance and habitat type. INTRODUCTION The American mink Mustela vison Schreber, 1777, is a North American mustelid predator that was brought to Europe for its quality fur. The first wild living populations of American minks appeared throughout Europe in the 1930s, as a large amount of American minks escaped from captivity or were deliberately released into the wild (Sidorovich 1993). At the end of the 1990s, American minks became a common European species living on the banks of rivers, lakes, ponds and water reservoirs (Dunstone 1993, Bartoszewicz & Zalewski 2003). The first report on their existence in the Czech Republic comes from the 1960s (Mazák 1964). The species started to spread across Bohemia at the turn of the millennium. An evaluation of a national questionnaire showed that in 2003 the American mink was present on 29.3% of the area of the Czech Republic (Červený et al. 2005). The American mink is an unspecialised predator (Wise et al. 1981, Br zeziński & Zurowski 1992, Jedrzejewska et al. 2001), hunting in water and also on land. Their presence often leads to a steep decrease in their prey population (Bartoszewicz & Zalewski 2003). They hunt mostly in littoral vegetation and in water (Erlinge 1969). Water invertebrates (crayfish), as well as various types of vertebrate species are their most common prey (Erlinge 1969, Sidorovich 2000, Jedrzejewska et al. 2001, Bartoszewicz & Zalewski 2003). Fish are a dominant component of their diet (Akande 1972, Lodé 1993). The particular compositon of 173 their diet varies throughout the year and depends on the habitat the mink lives in (Gerell 1967, Chanin & Linn 1980, Jenkins & Harper 1980, Jedrzejewska et al. 2001). MATERIAL AND METHODS The diet of the American mink was studied using an analysis of stomach content of individuals coming from Bohemia (Křivoklátsko Protected Landscape Area and Horažďovice region). In total, 51 stomachs were analysed. The samples were continuously collected during autumn (September–November) and winter seasons (December–February) in 2001–2006. All minks were caught in accordance with nature conservation legislation, with the aim to protect populati ons of indigenous animal species. All caught minks were processed using standard zoological methods. Empty stomachs (22 samples, 43%) were not included in further analysis. Stomach contents were ana lysed using standard methods (Hammershøj et al. 2004). Each stomach content was mixed with water on a Petri dish and individual components separated into groups. Various identification guides as well as comparative material were used to determine the prey remains (bones, scales, feather, teeth, fur or hair or other body parts) (Gaffrey 1961, Brom 1986, Teerink 1991). Results are showed as a percentage of volume (V %), indicating the volume percentage of a certain food component in all of the analysed full stomachs. In order to compare our results with those analysing American mink faeces, we also used frequency of occurrence (F %), indicating the occurrence percentage of a certain food component in all of the analysed full stomachs RESULTS AND DISCUSSION The analysis of 29 stomach contents showed that fish with the body length of up to 30 cm (F=51.7%; V=41.0%) were the dominant component in the American mink diet. The most frequently found species were from the families of Cyprinidae (similar to Erlinge 1969, Day & Linn 1972), also Percidae (Erlinge 1969, Bartosewicz & Zalewski 2003) and Esocidae (Erlinge 1969). It is clear that low activity of fish, caused by low water temperature during winter months, was the main reason for their high abundance in the mink diet (Gerell 1967, Erlinge 1969, Chanin & Linn 1980, Chanin 1981, Wise et al. 1981). The list of food compo nents is shown in Table 1. Mammals (Rodentia, Insectivora) were the second most important food component (F=48.7%; V=31.1%). Considering that most predators avoid eating insectivorous mammals (Lockie 1961, Macdonald 1977), occurrence of the lesser white-toothed shrew Crocidura suaveolens (F=6.9%; V=3.8%) in mink stomachs was very interesting. Similar findings are very rare and if there are some insectivores identified in the mink diet, it is usually only in small amounts (Wise et al. 1981, Brzeziński & Zurowski 1992, Maran et al. 1998, Jedrzejewska et al. 2001). The presence of a roe deer fur (Capreolus capreolus) in the stomachs proves the ability of minks to use large mammal carcasses as a food source (F=3.5%; V=3.5%). In comparison with other literature data, the richness of mammal species found in the diet of minks in Bohemia is very low. This could be caused not only by the limited amount of analysed samples and by the season of sample collection, but also, and mainly, by food availability in the study area. For example, Lepus, Rattus or Apodemus species were totally absent from the diet of minks in Bohemia, whereas they are very often hunted by mink populations living elsewhere (Day & Linn 1972, Chanin & Linn 1980, Dunstone & Birks 1987, Bartosewicz & Zalewski 2003). Exceptionally, small weasel predators such as the ermine Mustela erminea and weasel Mustela nivalis (Akande 1972, Chanin & Linn 1980) can also be a prey of the American mink, but neither these were recorded in our study. On the other hand, muskrats (Ondatra zibethicus) 174 Table 1. Diet of the American mink based on the analysis of stomach content. For explanations see Material and methods Tab. 1. Potrava norka amerického podle analýzy obsahu žaludků. Vysvětlivky viz Material and methods component / složka F [%] V [%] Crocidura suaveolens Talpa europea Ondatra zibethicus Microtus sp. Glis glis Rodentia sp. Capreolus capreolus Mammalia unidentified 6.90 3.45 3.45 3.45 3.45 21.05 3.45 6.90 3.79 1.21 3.41 3.45 2.76 13.03 3.45 3.45 mammals birds amphibians 52.10 20.69 3.45 27.55 13.24 1.72 Cyprinidae Esocidae Percidae Pisces unidentified 27.59 6.90 10.34 10.34 19.81 4.13 9.01 6.86 fish invertebrates plant material unidentified 55.17 17.24 41.38 6.90 41.00 0.91 9.76 0.01 are known to be hunted more often than we recorded (Bartosewicz & Zalewski 2003). It is obvious from the literature that mammals are a significant part of the American mink diet. The frequency of mammal component in mink diet ranges from 28.0% (Akande 1972) up to 61.8% (Brzeziński & Zurowski 1992). Bird remains (Passeriformes, Galliformes) were recorded in 20.7% of cases (V=13.2%), whereas Bartosewicz & Zalewski (2003) found bird remains, during the same season, only in 4–16% of cases. Birds happen to be the most common prey during breeding or migrating seasons (Gerell 1967, Erlinge 1969, Akande 1972, Wise et al. 1981, Brzeziński & Zurowski 1992, Maran et al. 1998). Therefore, in spring and summer birds can become the dominant component of the mink diet (Bartosewicz & Zalewski 2003). Frogs are another significant component of the American mink diet (Gerell 1967, Brzeziński & Zurowski 1992, Hammershøj et al. 2004), especially bullfrogs. Minks avoid eating toads (Sido rovich & Pikulik 1997). Despite the fact that minks often dig up wintering frogs (Jedrzejewska et al. 2001), we found frog remains in our samples only once. Remains of invertebrates were found in 17.2% of samples (V=0.9%). They were mostly water arthropods and insects, although some of these could have already been present in the guts of the consumed fish. Unlike water arthropods, especially crabs (Gerell 1967, Chanin & Linn 1980, Dunstone & Birks 1987, Brzeziński & Zurowski 1992, Sidorowich et al. 2001), insects are consumed by minks very rarely (Erlinge 1969, Jedrzejewska et al. 2001). 175 Plant remains (grass, twigs, leaves, roots) were found in 41.38 % of samples (V=9.76%). In six cases they formed the majority of the stomach content (V=20%=100%) and therefore it is highly unlikely that they were eaten by accident. A similar feeding behaviour was described in the pine marten (Martes martes) by Lockie (1961) who found plant remains only in some faeces samples during spring and autumn. The season of sample collection and, in some cases, the used methods of stomach analysis did not allow to prove the presence of bird eggs, earthworms or slugs in the American mink diet, which are common food components of mink populations living elsewhere (Chanin & Linn 1980, Brzeziński & Zurowski 1992, Maran et al. 1998, Jedrzejewska et al. 2001). The most significant factors influencing the American mink diet are season (Jedrzejewska et al. 2001, Bartosewicz & Zalewski 2003) and the food availability in the given area (Chanin & Linn 1980, Bartosewicz & Zalewski 2003). Other factors include prey behaviour, its abundan ce and the structure of mink habitat (Chanin & Linn 1980). For example, mammals, fish and amphibians are the most important food sources in riverine habitats, whereas birds and fish are dominating prey of minks living near lakes and ponds (Jedrzejewska et al. 2001, Bartosewicz & Zalewski 2003). Considering our small material of stomach samples collected mostly in autumn, it was not possible to evaluate seasonal changes of the diet, sex differences in the diet nor differences between sites. Our results show that food composition of American minks living in the Czech Republic is almost identical with that of minks living in Scotland (all year round) (Akande 1972) and very similar to that of minks living in Poland (autumn and winter) (Bartosewicz & Zalewski 2003). SOUHRN Potrava norka amerického, Mustela vison (Schreber, 1777), v České republice byla studována analýzou žaludků pocházejících z podzimního a zimního období z let 2001–2006. Norci pocházeli z území Čech (Horažďovicko, CHKO Křivoklátsko). Celkem bylo získáno 51 žaludků, prázdné (43 %) byly z dalších analýz vyloučeny. Ryby z čeledí Cyprinidae, Percidae a Esocidae tvořily nejvýznamnější část potravy (frekvence výskytu [F] =51.7 %; objemové procento [V] =41.0 %), následované savci (Rodentia, Insectivora) (F=48.7 %; V=31.1 %) a ptáky (F=20.7 %; V=13.2 %). Obojživelníci, hmyz, kadávery velkých savců a rostlinné zbytky doplňovaly potravu. Zajímavý je výskyt bělozubky Crocidura suaveolens a naopak velmi vzácná přítomnost obojživelníků v potravě zkoumaných norků. Z našich výsledků vyplývá, že norek americký se na území Čech živí poměrně širokým spektrem živočišných druhů (tab. 1). Potravní složení je pravděpodobně ovlivněno ročním obdobím, potravní nabídkou, početností a chováním kořisti a strukturou biotopu, který norek obývá. REFERENCES Akande M., 1972: The food of feral mink (Mustela vison) in Scotland. J. Zool., London, 167: 475–479. Bartoszewicz M. & Zalewski A., 2003: American mink, Mustela vison diet and predation on waterfowl in the Słońsk Reserve, western Poland. Folia Zool., 52: 225–238. Brom T. G., 1986: Microscopic identification of feather fragments of palearctic birds. Bijdr. Dierk., 56: 181–202. Brzeziński M. & Zurowski W., 1992: Spring diet of the American mink Mustela vison in the Mazurian and Brodnica Lakelands, northern Poland. Acta Theriol., 37: 193–198. 176 Červený J., Anděra M. & Koubek p., 2005: Co nového v naší fauně (Vyhodnocení dotazníků z let 2001–2003) [What is new in our nature (Evaluation of questionnaires from 2001–2003]. Myslivost, 53(12): 62–66 (in Czech). Day M. G. & Linn I., 1972: Notes on the food of feral mink Mustela vison in England and Wales. J. Zool., London, 167: 463–473. Dunstone N., 1993: The Mink. T & A. D. Poyser Ltd., London, 233 pp. Dunstone N. & Birks J. D. S., 1987: The feeding ecology of mink (Mustela vison) in coastal habitat. J. Zool., London, 212: 69–83. Erlinge S., 1969: Food habits of the otter Lutra lutra L. and the mink Mustela vison Schreber in a trout water in southern Sweden. Oikos, 20: 1–7. Gaffrey G., 1961: Merkmale der wildlebenden Säugetiere Mitteleuropas. Akademische Verlagsgesells chaft, Leipzig, 284 pp. Gerell R., 1967: Food selection in relation to habitat in mink (Mustela vison Schreber) in Sweden. Oikos, 18: 233–246. Hammershøj E., Thomsen E. A. & Madsen B., 2004: Diet of free-ranging American mink and European polecat in Denmark. Acta Theriol., 49: 337–347. Chanin P., 1981: The diet of the otter and its relations with the feral mink in two areas of southwest England. Acta Theriol., 26: 83–95. Chanin P. R. F. & Linn I., 1980: The diet of the feral mink (Mustela vison) in southwest Britain. J. Zool., London, 192: 205–223. Jedrzejewska B., Sidorovich V. E., Pikulík M. M. & Jedrzejewski W., 2001: Feeding habits of the otter and the American mink in Białowieża Primeval forest (Poland) compared to other Eurasian population. Ecography, 24: 165–180. Jenkins D. & Harper R. J., 1980: Ecology of otters in northern Scotland II. Analyses of otter (Lutra lutra) and mink (Mustela vison) faeces from Deeside, N. E. Scotland in 1977–78. J. Anim. Ecol., 49: 737–754. Lockie J. D., 1961: The food of the pine marten Martes martes in west Ross-Shire, Scotland. Proc. Zool. Soc. London, 136: 187–195. Lode T., 1993: Diet composition and habitat use of sympatric polecat and American mink in western France. Acta Theriol., 38: 161–166. Macdonald D. W., 1977: On food preference in the Red fox. Mammal Review, 7: 7–23. Maran T., Kruuk H., Macdonald D. W. & Polma M., 1998: Diet of two species of mink in Estonia: dis placement of Mustela lutreola by M. vision. J. Zool., London, 245: 218–222. Mazák V., 1964: Několik poznámek o rodu Lutreola Wagner v Československu [Some notes on the genus Lutreola Wagner in Czechoslovakia]. Lynx, n. s., 3: 17–29 (in Czech). Sidorovich V. E., 1993: Reproductive plasticity of the American mink Mustela vison in Belarus. Acta Theriol., 38: 175–183. Sidorovich V. E., 2000: Seasonal variation in the habits of riparian mustelids in river valleys of NE Belarus. Acta Theriol., 45: 233–242. Sidorovich V. E. & Pikulik M. M., 1997: Toads Bufo spp. in the diets of mustelid predators in Belarus. Acta Theriol., 42: 105–108. Sidorovich V. E., Macdonald D. W., Pikulik M. M. & Kruuk H., 2001: Individual feeding specialization in the European mink, Mustela lutreola and the American mink, M. vison in north-eastern Belarus. Folia Zool., 50: 27–42. Teerink B. J., 1991: Hair of West-European Mammals. Cambridge University Press, Cambridge, 224 pp. Wise M. H., Linn I. J. & Kennedy C. R., 1981: A comparison of the feeding biology of mink Mustela vison and otter Lutra lutra. J. Zool., London, 195: 181–213. 177 Lynx (Praha), n. s., 37: 179–195 (2006). ISSN 0024–7774 Distribution and status of Myotis bechsteinii in Bulgaria (Chiroptera: Vespertilionidae) Rozšíření a statut netopýra velkouchého (Myotis bechsteinii) v Bulharsku (Chiroptera: Vespertilionidae) Boyan P. Petrov National Museum of Natural History, Tsar Osvoboditel blvd. 1, BG–1000 Sofia, Bulgaria; boyanpp@nmnh.bas.bg received on 23 May 2006 Abstract. The first record of Myotis bechsteinii in Bulgaria dates from 1935. Since then, a total of 55 females, 141 males and two individuals of unknown sex have been recorded. Up to now only three breeding colonies have been found in Bulgaria. At present, the Bechstein’s bat is known from 34 localities (33 UTM squares) situated from sea level up to 1650 m. Since its first discovery, only two males have been found hibernating in caves. Besides that no data are available on wintering sites of the species in Bulgaria. Although most localities were at altitudes below 300 m, the highest number of individuals during summer was found in mountain beech and mixed coniferous woodlands at an elevation between 800 m and 1450 m a. s. l. During the swarming period, some individuals were found to make vertical migrations of ca. 770 m between their roosts and the site of the capture. Myotis bechsteinii was found in Pleistocene and Early Holocene deposits of only two caves in Bulgaria. However, it was one of the most abundant and common species during that time. At present, conservation of mature forests (i.e. sustainable forest management), maintenance of their connectivity and further planting of new forest clearings are considered the most important factors that could promote the occurrence of the species. Introduction There are comparatively less records of bats from the southern (both eastern and western) parts of Europe than those from its central and western parts. This is especially true for the Bechstein’s bat, Myotis bechsteinii (Kuhl, 1817). This species is considered to be distributed continuously in most Central European countries, but has a scattered distribution in Southern Europe and further to the east. Its occurrence in the south and east was reviewed for Portugal and Spain (Benzal & de Paz 1991), Italy (Vergari et al. 1998), Croatia (Dulić 1959, Kovačić & Dulić 1989), Slovenia (Kryštufek & Cerveny 1997), former Yugoslavia (Petrović et al. 1985), Albania (Uhrin et al. 1996), Bulgaria (Benda et al. 2003, Petrov 1997), Romania (Nagy et al. 2005), Greece (Hanák et al. 2001, Helversen & Weid 1990), European Turkey (Kahmann 1962, Furman & Özgül 2004), Asia Minor (Benda & Horáček 1998, Albayrak 2003), the Caucasus region (Rakhmatulina 1990, Tsytsulina 1999) and by DeBlase (1980) who provided data from the easternmost locality of the species in Iran. The Bechstein’s bat is considered a typical species of the temperate humid zone whose range is centred mainly in the western part of the Palaearctic region (Horáček et al. 2000). The distribution of the species is only well known in 179 the western and northern parts of its range. Southeastern records are often quite isolated from the main distribution range. Therefore, additional information on the occurrence at the margins of its range is needed to understand the recent scattered distribution of Myotis bechsteinii in Europe. Furthermore, new records coming from the poorly studied regions, incl. Bulgaria, may allow us to better understand what limits Bechstein’s bats in their occurrence. Besides depicting precise maps of its distribution, chorological data are essential for the analysis of the species’ ecological requirements. This is an important prerequisite to design up-to-date national and international conservation plans for the Bechstein’s bat, which has been listed as vulnerable in the ‘Red List of Threatened Species’ since 1994 (IUCN 2004). Material and Methods This review is based mainly upon literature sources. However, new field records from the period 1997-2006 were added. Bechstein’s bats were caught at cave or mine entrances, above streams and in forest clearings using 3 m, 6 m or 12 m long mistnets. At four sites (7, 18, 24, and 33), radio-telemetry (Regal 2000 telemetry receiver, Titley electronics Ltd.; LB-2 transmitter, Holohil Systems Ltd.) was used to discover the roosts of the tagged individuals. All captured bats were measured, aged, ringed and released afterwards. Additionally, since 1999 a small piece of the bat’s wing membrane was collected, using a sterile biopsypunch for future DNA analyses. All bats were captured under the license of the Ministry of the Environment and Waters (Bulgaria) and permissions from the local departments of forests (2001–2006). Results Review of the records from Bulgaria Heinrich (1936) was the first to record the Bechstein’s bat in the Balkans and particularly on the Bulgarian Black Sea coast. Among other mammals, he reported six bat species, including Myotis bechsteinii, for the first time for the Bulgarian fauna. Later on, Hanak & Josifov (1959) added one locality, showing that the Bechstein’s bat also occurs in the mountains (Rila Mts.). Beron (1961) found that this species hibernates in Bulgarian caves. After several field trips to Bulgaria, Czech zoologists reported new localities thereby showing that the species is not as rare as previously assumed (Horáček et al. 1974, Benda et al. 2003). Studies by Beshkov (1993) raised the number of Bulgarian localities to eight. Petrov (1997) summarized all previous records and added 4 new sites. Some recent regional bat surveys (Ivanova 1998, Pandurska & Beshkov 1998b, Pandurska et al. 1999, Petrov 2001, Popov et al. 2006) reported Bechstein’s bats from eight new localities. The most recent review of the distribution of bats in Bulgaria (including M. bechsteinii) was published by Benda et al. (2003). Only few papers deal with the historical evidence of this bat species in Bulgaria. Wołoszyn (1982) first reported it in cave deposits from the Upper Pleistocene. Until today there has been no contemporary record of the Bechstein’s bat from this locality (Bacho Kiro cave), although environmental conditions are still suitable. Popov (2000) found abundant remains of this species in sediments dated to originate from the Upper Pleistocene and Holocene (cave 15 and 16, Karlukovo village). He also suggested that the species was common and widely distributed in periods with mild and humid climate, when forests still covered larger areas of the Balkans and elsewhere. Recent occurrence of the Bechstein’s bat in the caves close to Karlukovo was supported with field data by Benda et al. (2003). In summary, the current knowledge of the distribution of Myotis bechsteinii shows that in Bulgaria, the species inhabits diverse habitats, covering a broad altitudinal range. However, at 180 most places only single individuals were caught. A higher population density was estimated for some of the localities (e. g., 7, 17, 18, and 33, see below) but only three maternity colonies have been found so far in Bulgaria. D i s t r i b u t i o n o f t h e B e c h s t e i n’ s b a t i n B u l g a r i a The Bechstein’s bat is known to occur at 34 localities (33 UTM squares of 10×10 km) from sea level up to 1650 m a. s. l. (Fig. 1, Table 1). All records before 1971 were more or less accidental findings. With few exceptions, density of the records from the mountainous regions (e.g. Stara Planina, Rhodopes, Pirin) is higher compared to the scattered distribution in the lowlands. There are no records of Myotis bechsteinii from the Bulgarian part of Dobrogea or the upper parts of the Thracian plain where open agricultural landscape is prevailing. Occurrence of the species in the lowlands is mostly associated with larger forested areas (e.g. Strandja Mts., 6, 7, 24, 26, and 34) or presence of a diverse mosaic of habitats (e.g. 10, 19, 21, 23, etc.). In many regions, the species has not yet been found or has been only occasionally proved. For example, no records are available from the northern slopes of the Rhodopes, eastern slopes of the Pirin and Rila, Slavyanka, Belasitsa and other mountain ranges, which all offer suitable environmental conditions. Fig. 1. Records of the Bechstein’s bat (Myotis bechsteinii) in Bulgaria (1935–2006), see Table 1 for explanation of numbers. Shaded are mountains above 1200 m a. s. l. Obr. 1. Nálezy netopýra velkouchého (Myotis bechsteinii) v Bulharsku (1935–2006), vysvětlení čísel viz tab. 1. Stínována jsou horská území nad 1200 m n. m. 181 182 district site m a. s. l. method m f UTM NH75 Heinrich 1936 Hanak & Josifov 1959 NH78 Hanak & Josifov 1959 GM18 Beron 1961 FP33 Horáček et al. 1974 KG71 Benda et al. 2003 KG71 B. Petrov, present paper KG71 B. Petrov, G. Kerth & KG71 B. Koenig, present paper B. Petrov, present paper KG71 Benda et al. 2003 NG69 B. Petrov, G. Kerth & B. Koenig, present paper NG69 B. Petrov, G. Kerth & D. NG69 Dechmann, present paper B. Petrov, present paper NG69 B. Petrov & G. Kerth, NG69 present paper B. Petrov & G. Kerth, NG69 present paper B. Petrov & T. Stoyanov, NG69 present paper B. Petrov & T. Stoyanov, NG69 present paper B. Petrov, G. Kerth & T. NG69 Ivanova, present paper B. Petrov, G. Kerth & T. NG69 Ivanova, present paper Benda et al. 2003 KH68 Benda et al. 2003 KH78 Benda et al. 2003 KH68 Benda et al. 2003 KH68 Benda et al. 2003 KH68 date reference 1 Kamtchiya Dolni Chiflik lower course of the river 10 shot-gun 1 1 June 1935 2 Varna Varna pallace Evksinovgrad 10 hand 1 2 3. 8. 1935 3 Samokov Sofia Borovetz 1350 hand 18. 7. 1950 4 Belogradchik Montana Gornata propast cave 600 hand 1 6. 2. 1960 5 Yagodina Devin Yagodinskata peshtera cave 1015 mist-net 1 2. 8. 1971 5 Yagodina Devin Yagodinskata peshtera cave 1015 mist-net 1 15. 8. 1978 5 Yagodina Devin Yagodinskata peshtera cave 1015 mist-net 1 4. 8. 1997 5 Yagodina Devin Yagodinskata peshtera cave 1015 mist-net 5 14. 9. 2001 5 Yagodina Devin Yagodinskata peshtera cave 1015 mist-net 2 17. 9. 2005 6 Primorsko Bourgas Arkutino swamp 10 mist-net 2 6. 6. 1972 7 Primorsko Bourgas Ropotamo Reserve, 10 mist-net 1 12. 9. 2001 Veliov vir 7 Primorsko Bourgas Ropotamo Reserve 10 mist-net 3 20. 4. 2002 7 Primorsko Bourgas Ropotamo Reserve 10 bat-box 2 4. 6. 2002 7 Primorsko Bourgas Ropotamo R., entrance gate 5 mist-net 1 1 25. 4. 2003 7 Primorsko Bourgas Ropotamo Reserve, 10 funnel trap 5 26. 4. 2003 Veliov vir 7 Primorsko Bourgas Ropotamo Reserve, 10 mist-net 1 25. 9. 2003 Veliov vir 7 Primorsko Bourgas Ropotamo Reserve, 5 mist-net 1 7. 5. 2004 entrance gate 7 Primorsko Bourgas Ropotamo Reserve, 5 mist-net 1 1 11. 6. 2004 entrance gate 7 Primorsko Bourgas Ropotamo Reserve, mouth 3 funnel trap 11 15. 6. 2004 of the Ropotamo river 8 Karlukovo Cherven bryagrocky amphitheatre 200 mist-net 1 15. 6. 1977 8 Karlukovo Cherven bryagrocky amphitheatre 200 mist-net 1 6. 8. 1978 8 Karlukovo Cherven bryagcave behind the monastery 200 mist-net 2 8. 8. 1978 8 Karlukovo Cherven bryagcave behind the monastery 200 mist-net 1 9. 8. 1978 8 Karlukovo Cherven bryagcave in the monastery 200 mist-net 4 9. 8. 1978 No.village Table 1. List of record localities of the Bechstein’s bat (Myotis bechsteinii) in Bulgaria (1935–2006) Tab. 1. Přehled lokalit nálezů netopýra velkouchého (Myotis bechsteinii) v Bulharsku (1935–2006) 183 9 Krivnya Razgrad Bozkova dupka cave 200 hand 1 15. 3. 1989 1 0 Buhovtsi Targovishte Prilepnata dupka cave 200 hand 1 25. 4. 1989 11 Leshko Blagoevgrad Leshtanskata peshtera cave 1000 mist-net 1 7. 6. 1996 12 Gorna Bela Varshets mine 1000 mist-net 1 3. 4. 1996 13 Breze Svoge Travninata cave 1050 mist-net 1 15. 9. 1996 14 Gintzi Godech Dinevata peshtera cave 1200 mist-net 1 5. 9. 1994 15 Divchovoto Teteven Grazdenitza cave 800 mist-net 1 28. 9. 1997 15 Divchovoto Teteven Grazdenitza cave 800 mist-net 3 19. 9. 2001 16 Karnare Karlovo Mazata cave 1350 mist-net 1 25. 9. 1997 17 Vidima Aprilitsi Pleven hut, Vodnite dupki c. 1400 mist-net 9 15. 8. 1997 17 Vidima Aprilitsi Pleven hut, Vodnite dupki c. 1400 mist-net 101 28. 8. 2001 17 Vidima Aprilitsi Pleven hut, Vodnite dupki c. 1400 mist-net 4 16. 8. 2003 18 Bov Svoge Izdremets, mine 1450 mist-net 5 11. 9. 1998 18 Bov Svoge Izdremets, mine 1450 mist-net 7 14. 9. 1999 18 Bov Svoge Izdremets, mine 1450 mist-net 9 2 16. 9. 2001 18 Bov Svoge Izdremets, mine 1450 mist-net 2 2. 10. 2001 18 Bov Svoge Izdremets, mine 1450 mist-net 2 3. 8. 2002 18 Bov Svoge Izdremets, mine 1450 mist-net 2 2 2. 9. 2002 18 Bov Svoge Izdremets, mine 1450 mist-net 3 18. 9. 2002 18 Bov Svoge Izdremets, mine 1450 mist-net 10 22. 8. 2003 18 Bov Svoge Izdremets, mine 1450 mist-net 3 17. 9. 2003 18 Bov Svoge Izdremets, mine 1450 mist-net 5 19. 9. 2003 18 Bov Svoge Izdremets, mine 1450 mist-net 1 22. 10. 2003 19 Ribino Kroumovgrad Samara cave 340 mist-net 1 20. 4. 1995 Beshkov 1993 MJ54 Beshkov 1993 MH89 Pandrurska & Beshkov FM64 1998a Pandurska & Beshkov FN98 1998b Pandurska & Beshkov FN86 1998b Pandurska 1999 FN77 Ivanova 1998 KH64 B. Petrov, G. Kerth & KH64 B. Koenig, present paper Ivanova 1998 LH04 Ivanova 1998 LH23 Schunger et. al. 2004 LH23 Schunger et. al. 2004 LH23 R. Pandurska & V. Beshkov FN97 present paper B. Petrov, V. Beshkov & Y. FN97 Gorelov, present paper B. Petrov, G. Kerth & FN97 B. Koenig, present paper B. Petrov & S. Beshkov, FN97 present paper B. Petrov & V. Beshkov, FN97 present paper B. Petrov, present paper FN97 B. Petrov, present paper FN97 B. Petrov & V. Beshkov, FN97 present paper B. Petrov & T. Stoyanov, FN97 present paper B. Petrov & T. Stoyanov, FN97 present paper B. Petrov & T. Stoyanov, FN97 present paper Petrov 1997 LF78 184 district site m a. s. l. method m f UTM LF78 C. Dietz & I. Schunger, present paper Petrov 1997 FN97 Petrov 1997 FM73 Petrov 2001 FM73 Petrov 1997 KH95 Benda et al. 2003 MG55 B. Petrov, present paper NG78 B. Petrov, G. Kerth & NG69 T. Ivanova, present paper Benda et al. 2003 KH58 B. Petrov & G. Stoyanov, KH58 present paper Benda et al. 2003 NG26 C. Dietz & I. Schunger, NG26 present paper Benda et al. 2003 GM32 B. Petrov, present paper FM92 B. Petrov, present paper FM91 B. Petrov, present paper FM91 C. Dietz & I. Schunger, LJ23 present paper Benda et al. 2003 LG90 E. Tilova, present paper NH45 B. Petrov & G. Kerth, GN06 B. Petrov & G. Kerth, GN06 B. Petrov & G. Kerth, GN06 B. Petrov & G. Kerth, GN06 B. Petrov & G. Kerth, GN06 all five records by the present paper Popov et al. 2006 NG85 date reference 1 9 Ribino Kroumovgrad rocky bridge 320 hand 1 18. 9. 2002 20 Lakatnik Svoge Svinskata dupka cave 500 mist-net 1 24. 8. 1995 21 Kresna Blagoevgrad Kresna gorge, on the road 200 hand 1 5. 10. 1995 21 Kresna Blagoevgrad Kresna gorge, Sheitan dere 200 mist-net 1 2. 7. 1997 22 Golyama Jelyazna, Troyan Toplya cave 700 hand 1 2. 2. 1997 23 Ustrem Topolovgrad Bozkite cave 200 mist-net 2 10. 4. 1998 24 Primorsko Burgas residence Perla 10 mist-net 1 18. 4. 1998 24 Primorsko Burgas oak forest 95 mist-net 1 14. 6. 2004 25 Brestnitsa Yablanitsa Seeva dupka cave 500 mist-net 1 1. 5. 1999 25 Brestnitsa Yablanitsa Seeva dupka cave 500 mist-net 4 15. 5. 2003 26 Mladezko Malko TarnovoLeyarnitzata cave ? 4 160 mist-net 1 25. 8. 1999 26 Mladezko Malko TarnovoLeyarnitzata cave ? 4 160 mist-net 2 11. 6. 2003 27 Ribnovo Gotse DeltchevManoilovata peshtera cave 1000 mist-net 2 22. 6. 2000 28 Kresna Blagoevgrad Peshterata, mine 1250 mist-net 2 1 25. 9. 2001 29 Ilindentsi Sandanski Sharaliiskata peshtera cave 1650 hand 1 1 7. 4. 2002 29 Ilindentsi Sandanski Sharaliiskata peshtera cave 1650 mist-net 1 25. 6. 2002 30 Muselievo Nikopol Nanin kamuk cave 140 mist-net 1 10. 6. 2002 31 Dolno Cherkovishte, HaskovoSedemte peshteri-Oreshari c. 320 observ. 1 30. 9. 2003 32 Golitsa Dolni Chiflik oak forest (Quercus cerris) 300 mist-net 4 13. 5. 2003 33 Gabrovnitsa Svoge Sedemte prestola Monastery 650 mist-net 3 7. 5. 2003 33 Gabrovnitsa Svoge Sedemte prestola Monastery 650 funnel trap 5 9. 5. 2003 33 Gabrovnitsa Svoge Sedemte prestola Monastery 650 mist-net 1 15. 4. 2006 33 Gabrovnitsa Svoge Sedemte prestola Monastery 650 funnel trap 9 16. 4. 2006 33 Gabrovnitsa Svoge Sedemte prestola Monastery 650 funnel trap 2 19. 4. 2006 34 Sinemorets Tsarevo Silistar Reserve 3 mist-net 1 August 1998 No.village Table 1. (continued) Tab. 1. (pokračování) Table 2. Vertical distribution of the localities of the Bechstein’s bat (Myotis bechsteinii) in Bulgaria (1935–2006) Tab. 2. Vertikální rozšíření lokalit nálezů netopýra velkouchého (Myotis bechsteinii) v Bulharsku (1935–2006) altitude (m) sites (see Table 1) 0–300 301–600 601–900 901–1200 1201–1500 1501–1800 1, 2, 6, 7, 8, 9, 10, 21, 23, 24, 26, 30, 32, 34 4, 19, 20, 25, 31 15, 22, 33 5, 11, 12, 13, 14, 3, 16, 17, 18, 28 29 total sites 14 5 3 6 5 1 rel. share [%] 41.2 14.7 8.8 17.6 14.7 2.9 New data In this study I report new data from eight previously unknown localities. The species was proved to occur in the Pirin Mts. (loc. 28, 29), close to the Danube (loc. 30), in the western Stara Planina east of the Iskar river gorge (loc. 18, 33), in the Ropotamo Reserve at the Black Sea coast (loc. 7, 24) and in the eastern Stara Planina Mts. (loc. 32). At the localities Ropotamo Reserve (Loc. 7), Vodnite dupki cave (loc. 17), Izdremets (loc. 18) and Sedemte Prestola monastery (loc. 33), Bechstein’s bats were captured more than twice. These sites (among others) were therefore supposed to hold higher population density and bat boxes (Scwengler 2 FN) have recently been installed at localities No. 7, 18 and 33 (2001–2002, Petrov & Kerth unpubl.). Localities No. 18, 28 and 29 broaden the altitudinal range where Myotis bechsteinii was found in Bulgaria (see below). Furthermore, at four localities that were already known before (5, 15, 19, and 25), the presence of the species was confirmed. While the majority of the other papers (cf. Benda et al. 2003) deal with the geographic distribution of the species, the present survey analyses some details of its occurrence. To provide better description of the habitats where the species was found, 32 localities (out of 34 known) were personally visited. Among many other bat localities in the country visited by the author, mistnetting was also attempted at some previously known sites (e.g. loc. 8, 11, 12) but no Bechstein’s bats were captured. Some of the known sites were visited in winter as well (e.g. loc. 9, 14, 17, 20, 22, 27, 29) but the species was not found. The maximum number of Bechstein’s bats captured per night was 11 specimens (loc. 18, 17; Schunger et. al. 2004). Since its first discovery in 1935, 55 females, 141 males and 2 specimens (loc. 3) of unknown sex have been captured. Out of these numbers, 129 males and 46 females (i.e. 89% of the total number) were caught between 1994 and 2006 (i.e. in a period when intensive regional bat surveys in Bulgaria were carried out mostly by local researchers including the author of this paper). Altitudinal distribution Most of the localities (n=14; 41.2%) are situated between sea level and 300 m a. s. l. (Table 2). This high rate of occurrence at lowland sites is partly due to the fact that numerous bat surveys in Bulgaria were carried out in this altitudinal range. On the other hand, Bulgarian bat fauna reaches its greatest species diversity per area between 100 m and 300 m (Pandurska 1996, Petrov 2001, Popov & Ivanova 1995) and regional fauna can consist of 17–20 bat species, which inhabit these 185 Table 3. List of the localities of the Bechstein’s bat (Myotis bechsteinii) in Bulgaria sorted by altitude. Composition of the dominant vegetation cover is after Bondev (1991). Temporal water presence is abbreviated as “temp.” Tab. 3. Přehled lokalit nálezů netopýra velkouchého (Myotis bechsteinii) v Bulharsku řazený podle nadmořské výšky. Složení dominujícího vegetačního pokryvu podle Bondeva (1991). Nestálá přítomnost vody je naznačena zkratkou “temp.” No. village, site m a. s. l. dominant vegetation cover 34 Sinemorets, Silistar Reserve 3 1 Kamtchiya, river 10 2 Varna, Evksinovgrad pallace 10 6 Primorsko, Arkutino swamp 10 7 Primorsko, Ropotamo Reserve 10 24 Primorsko, Rerla residence 10–95 30 Muselievo, Nanin kamuk cave 140 26 Mladezko, Leyarnitzata N 4 cave 160 8 Karlukovo, caves, ridge 200 9 Krivnya, Bozkova dupka cave 200 10 Buhovtsi, Prilepnata dupka cave 200 21 Kresna, Kresna gorge 200 23 Ustrem, Bozkite cave 200 32 Golitsa, oak forest (Q. cerris) 300 31 Dolno Cherkovishte, Sedemte peshteri- 320 Oreshari cave 19 Ribino, Samara cave 340 20 Lakatnik, Svinskata dupka cave 500 25 Brestnitza, Seeva dupka cave 500 4 Belogradchik, Gornata propast cave 600 33 Gabrovnitsa, Sedemte prestola Mon. 650 22 Golyama Jelyazna, Toplya cave 700 15 Divchovoto, Grazdenitza cave 800 11 Leshko, Leshtanskata peshtera cave 1000 12 Gorna Bela rechka, mine 1000 27 Ribnovo, Manoilovata peshtera cave 1000 5 Yagodina, Yagodinska peshtera cave 1015 186 water m f Quercus cerris, Q. frainetto river 1 with Mediterranean elements Quercus cerris, Q. freinetto river 1 1 Quercus pubescnes, Q. sea 1 2 virgiliana, C. orientalis Quercus spp., Acer campestre swamp 2 Quercus spp., Acer campestre river 1018 Querceta freinetti with sea 1 1 Mediterranean elements arable lands, Quercus spp., Carpinus orientalis river 1 Quercus cerris, Q. frainetto with Mediterranean elements river 3 Paliureta spina-christi, Quercus cerris, Q. frainetto river 5 4 Carpinus betulus, Q. cerris, river 1 Q. dalechampii Carpinus betulus, Q. cerris, lake 1 Q. dalechampii Platanus orientalis, Alnus glutinosa river 2 Qurcus dalechampii, Carpinus river 2 orientalis Querceta polycarpae river 4 Carpitneta orientalis with river 1 Mediterranean elements river 2 Carpitneta orientalis with river 2 Mediterranean elements Carpinteta orentalis river 1 Carpinteta orentalis temp. 5 Quercus cerris, Q. freinetto none 1 Fagus sylvatica moesiaca river20 Fagus sylvatica moesiaca river 1 Fagus sylvatica moesiaca river 4 Quercus dalechampii (+ temp. 1 Carpinus orientalis) Fageta sylvaticae river 1 Pinus sylvestris, Fagus sylvatica river 2 Pinus sylvestris, Fagus river 10 sylvatica 13 Breze, Travninata cave 1050 14 Gintsi, Dinevata peshtera cave 1200 28 Kresna, Peshterata, mine 1250 3 Samokov, Borovetz 1350 16 Karnare, Mazata cave 1350 17 Vidima, Vodnite dupki cave 1400 18 Bov, Izdremets, mine 1450 29 Ilindentsi, Sharaliiskata peshtera cave 1650 Carpineta orientalis (+ none 1 Fagus sylvatica) Carpineta orientalis (+ Fagus sylvatica) river 1 Fagus sylvatica moesiaca river 2 Picea abies, Fagus sylvatica river (+ Pinus sylvestris) Fageta sylvaticae none 1 Fageta sylvaticae river 23 Fagus sylvatica moesiaca, lake 49 Carpinus betulus Pinus heldreichi, P. sylvestris none 2 1 1 4 1 areas seasonally or permanently. Higher population density of Bechstein’s bats in the lowlands was reported also from Switzerland (Zingg 1982), Germany (Kerth 1997, Weishaar 1996) and the Czech Republic (Červený & Bürger 1989). The high number of localities in the lowlands of Bulgaria does not correlate with the species abundance. Considering all captured bats (n= 196), the average number of individuals from the 22 localities below 1000 m a. s. l. is 4.3 specimens per site, versus 8.2 from 12 localities between 1000 m and 1650 m (Table 1). Two male Bechstein’s bats were caught and radio-tracked at the swarming site of Izdremets (loc. 18) at 1450 m in August and September. On the next day after capture, both were found to roost in beeches (Fagus sylvatica) at lower altitudes (630–680 m, loc. 33). These individuals thus migrated vertically about 770 m (2.7 km one way straight distance) for swarming and turned back to their roosts. These observations are the first that demonstrate short-distance vertical flights in the species. The highest altitude at which Myotis bechsteinii was recorded in Bulgaria is 1650 m a. s. l. (loc. 29). The species (1 female, 2 males) was encountered at this location twice in three months. Other findings at comparable altitudes in central Spain (1500 m, Benzal & de Paz 1991), in the Swiss Alps at Montreux (1560 m, Chapuisat & Ruedi 1993), Jumelles (grotte au Tichodrome, 1750 m) (Arlettaz et al. 1993) indicate that the species can be occasionally found near the timberline. However in most of these high-altitude cases, bats probably went so high for swarming or feeding rather than for seasonal roosting. A skull was found at 1950 m (Pigna, cave No. F 7-813) in Italy (Liguria, Amelio 1973). Holocene findings of the Bechstein’s bat in Austria in caves from 1800 m (Bauer & Walter 1977) up to 2100 m (Spitzenberger 2001) show its historic occurrence in this region. The subfossil Italian locality ‘Pigna’ together with the record from 2100 m in Austria are the highest ever recorded localities for Myotis bechsteinii. Seasonal occurrence and wintering The highest number of Bechstein’s bats was captured in September, followed by August. At that time, 100 males were caught (i.e. 71% of all males reported in this study) versus only 12 females. The highest number of females was recorded in late April (19 ind.) and in early June (16 ind.) due to the discovery of three breeding colonies (Loc. No. 7, 33) found by radio-tracking of female bats. Bechstein’s bats were caught only twice in July (1950, 1997), though at the 187 locality No. 7 mistnetting in the forest was performed for 10 nights in 2002 and no Bechstein’s bats were captured. It is worth mentioning that since 1935, only two specimens have been found to hibernate in caves (loc. 4: 6 February 1960, loc. 22: 2 February 1997). Both were male and were found in crevices in the wall near the entrance. Only one of those records comes from the last ten years even though bat research in the country has been relatively intensive during this period. Myotis bechsteinii has never been observed during recent winter censuses of bats in numerous caves (about 60), mines (about 10) and other underground roosts, which host hibernating colonies of various vespertilionid species (cf. Benda et al. 2003). Habitat and roost preferences Forests and scrubs dominated mainly by Quercus spp., Carpinus spp. and occasionally by Platanus orientalis cover all lowland localities (i.e. 0–600 m) where the species was found in Bulgaria. In many cases (e.g. loc. 13, 17, 18, 28) the Bechstein’s bats were not caught in a roost but at a swarming site. Thus our knowledge on the preferred habitats is biased by the seasonal activity patterns. Oak and oriental plane forests offer relatively thick trees and Bechstein’s bats are thought to occupy mostly tree holes and crevices (cf. von Helversen & Weid 1990). Findings of the Bechstein’s bat in the Carpinus forests, where suitable trees are much less common could suggest that the species uses alternative roosts such as caves, mines, bunkers, tunnels, rock fissures, etc. Male specimens were occasionally recorded in a crevice between bricks of a rocky bridge (loc. 19) and in a rock crevice (loc. 31). Roosts of male Bechstein’s bats and two breeding colonies have recently been found in tree holes (2001–2006, Petrov & Kerth, new records). In the lowland forest of the Ropotamo Reserve, most of the used roosts were discovered in the common maples (Acer campestre) and rarely in Quercus cerris or Q. polycarpa. In the area of the Sedemte Prestola monastery (western Stara Planina Mts.), where the third breeding colony was found, all roosts were in beeches (Fagus sylvatica), though oak trees were also present. Single individuals were found in shallow cavities in relatively slender trees (DBH = 13–20 cm) at the height of 0.7–5 m above the ground (Table 4). Roosting groups (5–55 ind.) were found in thicker trunks (DBH = 40–55 cm) at the height of 5–10 m above the ground. A colony of 55 individuals has recently been observed emerging at dusk in this region but none of the bats was captured and included in the present analysis. This colony however is the largest known so far from the southeastern Europe. Bechstein’s bats were found only once (4 June 2002) in a bat box (Schwegler 2FN) in the Ropotamo Reserve. The two individuals were the first bats found in a bat box in Bulgaria, only a year after the boxes were installed. Since then no Bechstein’s bats have been found in the boxes (n=55) at this site but temporal roosting seems possible at times when no census was made. The majority of the capture sites were situated next to a permanent water body: river (n=23), sea (n=2), lake (n=2) and swamp (n=1). Only few of the localities lacked water (n=4) or had only temporal water sources (n=2) (Table 3). Climatic features of the localities: In Bulgaria, Myotis bechsteinii was found at sites with high average diurnal temperature during summer (22–25 °C) and very little annual precipitation (550 mm and lower) (loc. 21, 23), as well as at sites with low summer diurnal temperature (12–16 °C) and heavy precipitation (1000 mm and higher) (loc. 5, 17). Abundance was higher at the latter sites, where temperatures are moderate and the climate is clearly humid throughout the year. 188 Table 4. Tree roosts of the Bechstein’s bat (Myotis bechsteinii) in Bulgaria (2001–2006). CF – circumference of the trunk [cm]; DBH – trunk diameter at breast height [cm]; DG – distance to the ground [m] Tab. 4. Stromové úkryty netopýra velkouchého (Myotis bechsteinii) v Bulharsku (2001–2006). CF – obvod kmene [cm]; DBH – průměr kmene ve výši prsou [cm]; DG – vzdálenost od země [m] site date tree species Ropotamo (7) 13 September 2001 18 April 2002 20 April 2002 22 April 2002 23 April 2002 26 April 2003 27 April 2003 28 Sept. 2003 15 June 2004 Monastery (33) September 2001 9 May 2003 16 April 2006 17 April 2006 18 April 2006 CF DBH DG Acer campestre Quercus cerris Acer campestre Acer campestre Acer campestre Quercus polycarpa Acer campestre (dead trunk) Acer campestre Carpinus betulus 44 78 62 48 40 170 121 76 161 14 25 19 15 13 54 39 24 51 3.0 1.5 1.6 2.5 0.7 9.0 6.0 3.5 2.5 Fagus sylvatica Fagus sylvatica Fagus sylvatica Fagus sylvatica Fagus sylvatica 95 131 87 153 139 30 42 28 49 44 3.5 5.0 5.0 10.0 8.0 number of bats 1 male 1 male 1 male 1 male 1 male 6 females 1 female 1 male 11 females 1 male 5 females 10 females 1 female 55 individuals Capture methods Almost all specimens reported during the last 30 years in Bulgaria were caught using mistnets at cave and mine entrances. Captures at cave mouths (20 cases) make up the majority of the samples. The highest abundance of male bats was found during the swarming season (late summer and early autumn), while mistnetting at caves and mines in old growth mesophile mountain forests (e.g. loc. 17, 18). In the recent years (after 2001), females from several breeding groups/colonies have been caught at tree holes using a funnel trap. When the latter was set well, almost no escapes were observed. Captures with bare hands happened only occasionally. Bechstein’s bat has been found killed on the road only once in Bulgaria (Loc. No. 21), though many other species (some of them in high numbers) have been found in a road mortality survey in the Kresna gorge (SW Bulgaria, B. Petrov, unpubl.). The highest capture success of mistnetting in forests was proved when the net was set above a stream or shallow riverbed. In the latter case, captures of males were slightly prevailing (9 males, 7 females, loc. 7, 24, 33, and 34) compared to strongly male dominated catches at the swarming sites (see below). Discussion Few studies on Myotis bechsteinii provide detailed data on the vegetation and habitat type in the surroundings of its roosts. The existing data suggest that this bat is a typical inhabitant of the old growth forests in western Europe (Kerth 1997, Schlapp 1990). In central parts of the continent, the Bechstein’s bat also occurs in open river valleys with groves in their vicinity, 189 and in small woodland patches, which alternate with arable and parklands (Červený & Bürger 1989, Harmata 1969). In southern Europe, Myotis bechsteinii was found in more or less dense xerophile forests of the Mediterranean type in Portugal and Spain (Benzal & de Paz 1991), Greece (Hanák et al. 2001, von Helversen & Weid 1990) and Corsica (Libois & Franken 1981). In Bulgaria, the habitats vary with altitude, ranging from dry mixed lowland oak forests to humid mountain beech and mixed coniferous woodlands. As Bechstein’s bats feed mostly on non-airborne invertebrates and water independent flying insects (Dondini & Vergari 1999b, Krochko 1990, Wolz 1993), the presence/absence of water (streams, lakes, etc.) probably does not restrict the species distribution. On the other hand, water availability favours insect diversity, which is presumed to promote the occurrence of bats. The Bechstein’s bat is known as a non-migratory species. Most local movements cover less than 10 km and the females exhibit extreme philopatry (Kerth et al. 2000, 2002). The longest known dispersal flights reach 39 km (Haensel 1991), 43 km (Rudolf et al. 2004) and exceptionally up to 60 km (Kerth & Petit 2005). The maximum flights proved so far by radiotracking in Bulgaria are between 2 and 3 km (loc. 7, 18, 24, and 33). At the swarming sites with the highest capture rate in Bulgaria, more males than females were captured. At loc. 17 only one of the 24 specimens was a female (4.2%). At loc. 18, only 4 out of 54 specimens were females (7.5%). This could indicate higher mobility or longer dispersal flights of the males during the pre- and postmating period compared to the females (cf. Kerth et al. 2003). The high number of records in late summer probably reflects both the bat research activity, which usually is at its maximum during these months, and the fact that the intensity of bat swarming at caves and mines is highest during this period. The latter was proved in many countries (cf. Parsons et al. 2003, Kerth et al. 2003), including Bulgaria (cf. Schunger et al. 2004). The scarce winter records of Myotis bechsteinii in Bulgaria suggest that the species rarely uses underground roosts in this part of its life cycle. In the Czech Republic (Šumava Mts.), Bechstein’s bats were frequently found to hibernate in caves and galleries only during harsh winters (Červený & Bürger 1989). In Great Britain, more records of this bat from underground sites have been reported during severe winters (Harrington et al. 1995). Findings of the Bechstein’s bat all over Europe are relatively rare compared to records of the other species of the continental bat fauna (Baagøe 2001). Many of the recent peripheral localities remain isolated from each other and presumably some marginal populations could become allopatric. Only in a few regions of Central Europe (e.g. parts of Germany and the Czech Republic) it occurs in higher population density and substantial signs of a dramatic decline are not yet recognised (Červený & Bürger 1989, Kerth 1997, Schlapp 1990, Weishaar 1996). In Bulgaria, the Bechstein’s bat ranks among the common bat species with a continuous distribution in regions with larger forest coverage (e.g. Strandja Mts. along the Black Sea coast). On the other hand, only single individuals were caught in majority of the other sites. At present, the species could be considered “rare but locally common” with regard to distribution and “vulnerable” with regard to its environmental sensitivity and low colonisation abilities. Subfossil evidence from Austria (Bauer & Walter 1977, Rabeder 1972, Spitzenberger 2001), Great Britain (Yalden 1999), Bulgaria (Popov 2000), Czech Republic (Kowalski 1962), Hungary (Topál 1959), France (Mein 1975, Sevilla 1990), Italy (Kotsakis & Petronio, 1980), Poland (Kowalski 1956, Ochman 1999), Spain (Sevilla 1989) and Ukraine (Krochko 1990) show that Myotis bechsteinii was one of the most common bat species in the Pleistocene and Early Holocene deposits. Milder, humid climate and the presence of continuous deciduous wo190 odlands all over the continent and particularly in Bulgaria presumably favoured this abundance especially during the Interpleniglacial and the Early Holocene (Peshev et al. 2004). Since then the climate has changed many times and forests were cut in vast regions including clearance of deciduous woodlands in Europe during the post-Neolithic (Yalden 1999). This probably led to the present fragmentation of the species range and population decline. One possible reason of the current rarity of the Bechstein’s bat in most parts of Europe may be the limited distribution and size of old growth forests as well as their low connectivity. Deforestation and unsustainable forest management practices led to a reduction of roost availability, which seems to be the most important environmental factor for the species occurrence (Harrington et al. 1995, Hutson et. al. 2001). Another possible reason could be the concentration of bat research activities, which are not evenly spread over the species range. Although the research coverage varies from year to year, it probably plays a significant role in accuracy of the population assessments at local and national level. The Bechstein’s bat is protected by law in at least 31 of the European countries, which have joined the EUROBATS agreement, including Bulgaria. Considering the global fragmentation of forest habitats preferred by the species, its low population density and poor dispersal abilities, Myotis bechsteinii was also classified as a “vulnerable” species in the new edition of the Bulgarian Red Data Book (Petrov in press). Because of its sedentary life style, a “colonyorientated“ conservation approach was suggested in Germany (Meschede & Heller 2000). Bat box projects in Europe, e.g. in Italy (Dondini & Vergari 1999a), Germany (Kerth 1997, Taake & Hildenhagen 1989), United Kingdom (Schofield et al. 1997) and recently in Bulgaria (Petrov & Kerth, unpublished) proved that Myotis bechsteinii is among the first bat species, which occupy these artificial shelters, especially those that resemble woodpecker’s holes (e.g. 2 FN Schwegler’s boxes). Furthermore, restoration and planting of new forest clearings will contribute to the connectivity of suitable habitats and thus to spreading of the Bechstein’s bat at least in the centre of its range. SOUHRN První nález Myotis bechsteinii v Bulharsku pochází z roku 1935 a od té doby bylo zaznamenáno celkem 55 samic, 141 samců a dva jedinci neznámého pohlaví, avšak do současné doby byly v Bulharsku nalezeny jen tři mateřské kolonie. Netopýr velkouchý je znám z Bulharska z celkem 34 lokalit (33 čtverců UTM) ležících v nadmořské výšce od hladiny moře po 1650 m. Jen dva jedinci (samci) byli dosud nalezeni zimující v jeskyních, jiné nálezy ze zimovišť nejsou známy. Většina lokalit leží ve výšce do 300 m n. m., nejvyšší počet jedinců byl ale zaznamenán v horských bukových a smíšených lesích ve výšce mezi 800 a 1450 m n. m. V průběhu “swarmovacího” období byly zaznamnány vertikální migrace o zhruba 770 m mezi denním úkrytem a místem odchytu. Myotis bechsteinii byl doložen z pleistocénních a spodnoholocénních vrstev dvou jeskyní v Bulharsku, ovšem v těchto údobích představoval jednoho z nejhojnějších a nejběžnějších netopýrů. Ochrana původních lesních porostů (tj. udržitelné lesnické hospodaření), udržování jejich propojení a obnova hospodářských lesů jsou faktory v současné době nejvíce považovány za nezbytné pro ochranu druhu v Bulharsku. Acknowledgments I wish to thank T. Ivanova (NMNH, Sofia), T. Stoyanov (Sofia), A. Gueorguieva (BRPG, Sofia), B. Barov (BSPB, Sofia), N. Simov (NMNH, Sofia), R. Pandurska-Whitcher (Sofia), C. Dietz and I. Schunger (Tübingen), E. Tilova (Green Balkans) as well as all other colleagues who provided material or helped me during the field research. V. Beshkov (Institute of Zoology, Sofia) offered logistic support and assisted my 191 activities with endless enthusiasm. V. Popov (Institute of Zoology, Sofia) gave some valuable suggestions on the manuscript. I also thank A. Hutson (BCT, London), G. Dondini (Florence), H. J. Baagøe (Copenhagen), I. Rakhmatulina (Baku), K.-G. Heller (Magdeburg), M. Paunović (Belgrade), M. Zagmajster (Ljubljana), H. Schofield (The Vincent Wildlife Trust, London), P. Boye (Bonn) and K. Safi (Zürich) for providing me with some of the literature sources. Field travels between 2001 and 2004 were supported by the Swiss National Science Foundation (c/o SCOPES program, grant No. 7BUPJ062292). My sincere thanks go to G. Kerth and B. König (Zürich), who took part in two of the research trips and kindly shared their experience and literature on the Bechstein’s bat. D. Dechmann (Zürich) and G. Kerth commented earlier versions of the manuscript and especially D. Dechmann generously provided linguistic help. References Albayrak İ., 2003: The Bats of the Eastern Black Sea Region in Turkey (Mammalia: Chiroptera). Turk. J. Zool., 27: 269–273. Amelio M., 1973: Su alcuni ritrovamenti di crani di Chirotteri. Boll. Gruppo Speleol. Imper. CAI, 3: 50–51. Arlettaz R., Lugon A., & Sierro A., 1993: Inventaire des chauves-souris du Valais. Catalogue de sites. Seconde contribution a l’inventaire des sites valaisans abritant des chauves-souris. Rapport final de la campagne de prospection 1985–1990 du Groupe Valaisan pour l’etude et la protection des chauvessouris, 146 pp. Baagøe H. J., 2001: Myotis bechsteinii (Kuhl, 1818) – Bechsteinfledermaus. Pp.: 443–471. In: Krapp F. (ed.): Handbuch der Säugetiere Europas. Band 4: Fledertiere. Teil I: Chiroptera I. Rhinolophidae, Vespertilionidae 1. Aula-Verlag, Wiebelsheim, x+604 pp. Benda P. & Horáček I., 1998: Bats (Mammalia: Chiroptera) of the Eastern Mediterranean. Part 1. Review of distribution and taxonomy of bats in Turkey. Acta Soc. Zool. Bohem., 62: 225–313. Benda P., Ivanova I., Horáček I., Hanák V., Červený J., Gaisler J., Gueorguieva A., Petrov B. & Vohralík V., 2003: Bats (Mammalia: Chiroptera) of the Eastern Mediterranean. Part 3. Review of bat distribution in Bulgaria. Acta Soc. Zool. Bohem., 67: 245–357. Benzal J. & de Paz O., 1991: Los murcielagos de España y Portugal. Colleccion Tecnica, Icona, Madrid, 330 pp. Beron P., 1961: Contribution a la connaissance des chauves-souris Bulgares. Fragm. Balcan., 24(3): 189–195. Beshkov V., 1993: Bats. Pp.: 631–644. In: Sakalyan M. (ed.): The National Biological Diversity Conserva tion Strategy. Biodiversity Support Program, Washington, D.C., vii+663 pp (in Bulgarian). Bondev I., 1991: The Vegetation of Bulgaria. St. Kliment Ohridski University Press, Sofia, 183 pp (in Bulgarian). Červený J. & Burger P., 1989: Bechstein’s bat, Myotis bechsteini (Kuhl, 1818), in the Šumava region. Pp.: 591–598. In: Hanak V., Horáček I. & Gaisler J. (eds.): European Bat Research 1987. Charles University Press, Praha, xxii+718 pp. Chapuisat M. & Ruedi M., 1993: Les chauves-souris dans le canton de Vaud: status et evolution des populations. Le Rhinolophe, 10: 1–37. DeBlase A., 1980: The bats of Iran: Systematics, distribution, ecology. Fieldiana Zool. (N. S.), 4: 1–424. Dondini G. & Vergari S., 1999a: First data on the use of bat-boxes in a forest area of Central-North Italy. Pp.: 15–16. In: Cruz M. & Kozakiewicz K. (ed.): VIIIth European Bat Research Symposium. 23–27 August 1999. Kraków – Poland. Abstracts. Chiropterological Information Center, Institute of Animal Systematics and Evolution PAS, Kraków, 86 pp. Dondini G. & Vergari S., 1999b: First data on the diets of Nyctalus leisleri (Kuhl, 1817) and Myotis bechsteinii (Kuhl, 1817) in the Tuscan-Emilian Appennines (North-central Italy. Pp.: 191–195. In: Dondini G., Papalini O. & Vergari S. (eds.): Atti del I° Covegno Italiano sui Chirotteri. Castell’Azzara (Grosseto), 28–29 Marzo 1998. Tipografi a Ceccarelli, Grotte di Castro, 360 pp. 192 Dulić B., 1959: Zweiter Nachweis der Bechsteinfledermaus Myotis bechsteini Kuhl, 1818, für Jugoslawien. Säugetierk. Mitt., 7(2): 75. Furman A. & Özgül A., 2004: The distribution of cave-dwelling bats and conservation status of underground habitats in Northwestern Turkey. Biol. Cons., 120: 243–248. Haensel J., 1991: Vorkommen, Überwinterungsverhalten und Quartierwechsel der Bechsteinfledermaus (Myotis bechsteini) in Land Brandenburg. Nyctalus (N. F.), 4(1): 67–78. Hanak V. & Josifov M., 1959: Zur Verbreitung der Fledermäuse Bulgariens. Säugetierk. Mitt., 7: 145–151. Hanák V., Benda P., Ruedi M., Horáček I. & Sofianidou T., 2001: Bats (Mammalia: Chiroptera) of the Eastern Mediterranean. Part 2. New records and review of distribution of bats in Greece. Acta Soc. Zool. Bohem., 65: 279–346. Harrington L. A., Catto C. M. C. & Hutson A. M., 1995: The Status and Distribution of Barbastelle Bat (Barbastella barbastellus) and Bechstein’s Bat (Myotis bechsteinii) in the UK, with Recovery Plans. Report of the Bat Conservation Trust, English Nature’s Species Recovery Programme, London, 55 pp. Harmata W., 1969: Summer colony of Bechstein’s bat Myotis bechsteini (Kuhl) in Szymbark near Gorlice, Rzeszow District. Remarks about biology and occurrence. Przegl. Zool., 13(2): 233–237 (in Polish). Heinrich G., 1936: Ueber die von mir im Jahre 1935 in Bulgarien gesammelten Saugetiere. Mitt. Koenigl. Naturwissensch. Inst., Sofia, 9: 33–48. Horáček I., Červený J., Taušl A. & Vítek D., 1974: Notes on the mammal fauna of Bulgaria (Insectivora, Chiroptera, Rodentia). Vest. Čs. Společ. Zool., 38: 19–31. Horáček I., Hanák V. & Gaisler J., 2000: Bats of the Palearctic region: a taxonomic and biogeographic review. Pp.: 11–157. In: Wołoszyn B. W. (ed.): Proceedings of the VIIIth European Bat Research Symposium. Vol. I. Approaches to Biogeography and Ecology of Bats. Chiropterological Information Center, Institute of Systematics and Evolution of Animals PAS, Kraków, 280 pp. Hutson A. M., Mickleburgh S. P. & Racey P. A., 2001: Microchiropteran Bats: Global Status Survey and Conservation Action Plan. IUCN/SSC Chiroptera Specialist Group. IUCN, Gland, Switzerland and Cambridge, UK, x+258 pp. Ivanova T., 1998: First data on bats of the Central Balkan Mts., Bulgaria. Vespertilio, 3: 29–36. IUCN, 2004: 2004 IUCN Red List of Threatened Species. <www.iucnredlist.org>. Kahmann H., 1962: Neue Ergebnisse in der Säugetierforschung in der Türkei. Säugetierk. Mitt., 10: 112–116. Kerth G., 1997: Verbreitung und Schutz waldlebender Fledermausarten unter besonderer Berücksichtigung der Bechsteinfledermaus (Myotis bechsteinii) in den Laubwäldern Mainfrankens. Proc. Naturschutzzentr. Wasserschloss Mitwitz, 1: 27–29. Kerth G. & Petit E., 2005: Colonization and dispersal in a social species, the Bechstein’s bat (Myotis bechsteinii). Mol. Ecol., 14: 3943–3950. Kerth G., Mayer F. & König B., 2000: MtDNA reveals that female Bechstein’s bats live in closed societies. Mol. Ecol., 9: 793–800. Kerth G., Safi K. & König B., 2002: Mean colony relatedness is a poor predictor of colony structure and female philopatry in the communally breeding Bechstein’s bat (Myotis bechsteinii). Behav. Ecol. Sociobiol., 52: 203–210. Kerth G., Kiefer A., Trappmann C. & Weishaar M., 2003: High gene diversity at swarming sites suggest hot spots for gene flow in the endangered Bechstein’s bat. Cons. Genet., 4: 491–499. Kotskais T. & Petronio C., 1980: I Chiroterri del Pleistocene superiore della grotta di Spinagallo (Siracusa, Sicilia). Boll. Serv. Geol. Ital., 51: 49–76. Kovačić D. & Dulić B., 1989: Contribution to the knowledge of bats (Chiroptera, Mammalia) of Central Dalmatia. Biosistematika, 14(2): 31–40 (in Croatian). Kowalski K., 1956: Insectivores, bats and rodents from the Early Pleistocene bone breccia of Podlesice near Koczyce, Poland. Acta Paleontol. Polon., 1: 331–394. Kowalski K., 1962: Bats of the Early Pleistocene of Koneprusy (Czechoslovakia). Acta Zool. Cracov., 9: 145–156. 193 Krochko Y. I., 1990: Biology of the Bechstein’s bat (Myotis bechsteini) at the west of the Ukraine. Pp.: 80–82. In: Il’in V. Ju. (ed.): Rukokrylye. Materialy pjatogo vsesojuznogo soveshchaniya po rukokry lym (Chiroptera) [Bats. Proceedings of the Fifth Federal Bat Conference (Chiroptera)]. Vsesojuznoe teriologičeskoe obščestvo (Leningradskoe otdelenie), Penzenskij Gossudarstvennyj Pedagogičeskij Institut imeni V. G. Belinskogo, Penza, 174 pp (in Russian). Kryštufek B. & Cerveny J., 1997: New and noteworthy records of bats in Slovenia. Myotis, 35: 89–93. Libois R. M. & Vranken M., 1981: Myotis bechsteini en Corse. Mammalia, 45: 380. Mein P., 1975: Les Chiropteres (Mammalia) du gisement pleistocene des Abimes de la Fage a Noailles (Correze). Nouv. Arch. Mus. Hist. Natur., Lyon, 13: 57–67. Meschede A. & Heller K.-G., 2000: Ökologie und Schutz von Fledermäusen in Waldern. Schriftenreihe Landschaftspflege und Naturschutz 66. Bundesamt für Naturschutz, Bonn, 374 pp. Nagy Z. L., Barti L., Dóczy A., Jére Cs, Postawa T., Szántó L., Szodoray-Parádi A. & Szodoray-Parádi F., 2005: Survey of Romania’s Underground Bat Habitats. Status and distribution of cave dwelling bats. Report for BP Conservation Programme, 44 pp. Ochman K., 1999: Analysis of holocene bat fauna from Pod Sokola gora cave in systematics and zoogeographical aspects. P.: 44. In: Cruz M. & Kozakiewicz K. (ed.): VIIIth European Bat Research Symposium. 23–27 August 1999. Kraków – Poland. Abstracts. Chiropterological Information Center, Institute of Animal Systematics and Evolution PAS, Kraków, 86 pp. Pandurska R., 1996: Altitudinal distribution of bats in Bulgaria. Myotis, 34: 45–50. Pandurska R. & Beshkov V., 1998a: Bats (Chiroptera) of high mountains of southern Bulgaria. Obser. Mont. Moussala, Sofia, 7: 135–140. Pandurska R. & Beshkov V., 1998b: Species diversity of bats in underground roosts in the Western Stara planina Mts (Bulgaria). Vespertilio, 3: 81–91. Pandurska R., Beshkov V. & Pandurski I., 1999: Bats (Chiroptera) from the karst region of Gintzi village (North-West Bulgaria). Acta Zool. Bulgar., 55: 69–72. Parsons K. N., Jones G. & Greenaway F., 2003: Swarming activity of temperate zone microchiropteran bats: effects of season, time of night and weather conditions. J. Zool., London, 261: 257–264. Peshev T., Peshev D. & Popov V., 2004: Fauna of Bulgaria, 27, Mammalia. Academic Publisher “Marin Drinov”, 632 pp Petrov B., 1997: Distribution of Myotis bechsteini (Leisler in Kuhl, 1818) (Vespertilionidae, Chiroptera) in Bulgaria. P.: 37. In: Book of abstracts, Biodiversity and Ecological Problems of Balkan Fauna. International Conference, Sofia, 26–29 May 1997. Petrov B., 2001: Bats (Mammalia, Chiroptera) in Kresna gorge, SW Bulgaria. Pp.: 325–330. In: Beron P. (ed.). Biodiversity of Kresna gorge. National Museum of Natural History, Institute of Zoology, Sofia, 349 pp (in Bulgarian, English summary). Petrov B., in press: Bechstein’s bat (Myotis bechsteinii). In: Golemansky M. (ed.): Red Data Book of Bulgaria. Petrović P., Dzukić G. & Milenković M., 1985: Neue Angaben zur Verbreitung der Bechsteinfledermaus, Myotis bechsteini Kuhl, 1818 (Chiroptera), Mammalia) in Serbien. Proc. Fauna Serbia, 4: 227–231. Popov V., 2000: The mammals (Mammalia: Insectivora, Chiroptera, Lagomorpha, Rodentia) from Cave 16 (North Bulgaria) and paleoenvironmental changes during the Late pleistocene. Pp.: 159–240. In: Ginter B., Kozlowski J. K. & Laville K. (eds.): Temnata dupka. Excavations in Karlukovo Karst Area, Bulgaria. Jagellonian University Press, Krakow, 418 pp. Popov V. & Ivanova T., 1995: Morphoecological analysis and late quaternary history of a bat community in a karstic landscape in North Bulgaria. Myotis, 32–33: 21–31. Popov V., Pandurski I., Pandurska-Whitcher R. & Beshkov V., 2006: Small mammals (Insectivora, Chiroptera, Lagomorpha, Rodentia) in the area of Strandzha Mountain, South-Eastern Bulgaria. In: Pp.: 87–104. In: Chipev N. (ed.): Challenges in Establishment and Management of a Trans-Border Biosphere Reserve Between Bulgaria and Turkey in Stranja Mountain, Bourgas, 10–13 November 2005. Burgas. Rabeder G., 1972: Die Insektivoren und Chiropteren (Mammalia) aus dem Altpleistozän von Hudsheim (Niederösterreich). Ann. Naturhistor. Mus. Wien, 76: 375–474. 194 Rakhmatulina I. K., 1990: Discovery of the Bechstein’s bat (Myotis bechsteini) and the Grey long eared bat (Plecotus austriacus) in Azerbajan. Pp.: 174–175. In: Novosti faunistiki i sistematiki [Advances in Faunistics and Systematics]. Institute of Zoology, Kiev (in Russian). Rudolf B.-U., Kerth G., Schlapp G. & Wolz I., 2004: Bechsteinfledermaus, Myotis bechsteinii (Kulh, 1817). Pp.: 188–202. In: Meschede A. & Rudolf B.-U. (eds.): Fledermäuse in Bayern. Eugen Ulmer Verlag, München, 411 pp. Schlapp G., 1990: Populationdichte und Habitatansprüche der Bechsteinfledermaus Myotis bechsteini (Kuhl, 1818) im Steigerwald (Forstamt Erbach). Myotis, 28: 39–58. Schober W. & Grimmberger E., 1998: Die Fledermäuse Europas: Kennen, bestimmen, schützen. Kosmos, Stuttgart, 265 pp. Sevilla P., 1989: Quaternary fauna of bats in Spain: paleoecologic and biogeographic interest. Pp.: 349–355. In: Hanak V., Horáček I. & Gaisler J. (eds.). European Bat Research 1987. Charles University Press, Praha, xxii+718 pp. Sevilla P., 1990: The fauna of bats from Upper Pleistocene locality of Santenay (Cote d’or, France). Quaternaire, 2: 101–110. Schofield H., Greenaway F. & Morris C. J., 1997: Preliminary studies on Bechstein’s bat. Pp.: 71–74. In: The Vincent Wildlife Trust Review of 1996 and an overview of the Trust’s mammal work since its inception. The Vincent Wildlife Trust, London, 92 pp. Schunger I., Dietz C., Merdschanova D., Merdschanov S., Christov K., Borissov I., Staneva S. & Petrov B., 2004: Swarming of bats (Chiroptera, Mammalia) in the Vodnite Dupki Cave (Central Balkan National Park, Bulgaria). Acta Zool. Bulgar., 56: 323–330. Spitzenberger F., 2001: Die Säugetierfauna Österreichs. Bundesministerium für Land- und Forstwirschaft, Umwelt und Wasserwirtschaft, Graz, 895 pp. Stebbings R. E. & Griffith F., 1986: Distribution and Status of Bats in Europe. Institute of Terrestrial Ecology, Abbots Ripton, Huntington, 142 pp. Taake K-H. & Hildenhagen U., 1989: Nine years’ inspection of different artificial roosts for forest-dwelling bats in Northern Westfalia: some results. Pp.: 487–493. In: Hanák V., Horáček I. & Gaisler J. (eds.): European Bat Research 1987. Charles University Press, Praha, xxii+718 pp. Topál G., 1959: Die subfossile Fledermausfauna der Felnische von Istallosko. Verteb. Hungar., 2: 215–226. Tsytsulina E. A., 1999: Novye nahodki rukokrylyh na Zapadnom Kavkaze [New records of bats (Chiroptera) in the Western Caucasus]. Plecotus et al., 2: 79–83 (in Russian, with a summary in English). Uhrin M., Horáček I., Šíbl J. & Bego F., 1996: On the bats (Mammalia: Chiroptera) of Albania: survey of the new records. Acta Soc. Zool. Bohem., 67: 63–71. von Helversen O. & Weid R., 1990: Die Verbreitung einiger Fledermausarten in Griechenland. Bonn. Zool. Beitr., 41: 9–22. Yalden D., 1999: The History of British Mammals. Poyser, London, 305 pp. Vergari S., Dondini G. & Ruggieri A., 1998: On the distribution of Myotis bechsteinii (Kuhl, 1817) in Italy (Chiroptera: Vespertilionidae). Hystrix (n. s.), 10(2): 49–56. Weishaar M., 1996: Status der Bechsteinfledermaus (Myotis bechsteini) im Westen von Rheinland-Pfalz. Nyctalus (N. F.), 6(2): 121–128. Wołoszyn B. W., 1982: Chiroptera. Pp.: 40–45. In: Kozlovski J. (ed.): Excavations in the Bacho Kiro cave, Bulgaria. PKW, Warszawa. Wolz I., 1993: Das Beutespektrum der Bechsteinfledermaus Myotis bechsteini (Kuhl, 1818) ermittelt aus Kotanalysen. Myotis, 31: 27–68. Zingg P. E., 1982: Die Fledermäuse (Mammalia, Chiroptera) der Kantone Bern, Freiburg, Jura und Solothurn. Systematische und geographische Übersicht zu den bisher gesammelten und beobachteten Chiropteren. Lizenziatsarbeit, Diplomathesis, Zoologisches Institut Universität Bern, 45 pp. 195 Lynx (Praha), n. s., 37: 197–200 (2006). ISSN 0024–7774 First record of Myotis blythii in Poland (Chiroptera: Vespertilionidae) Pierwsze stanowisko Myotis blythii w Polsce (Chiroptera: Vespertilionidae) Krzysztof Piksa Cracow Pedagogical University, Institute of Biology, Podbrzezie 3, PL–31-054 Kraków, Poland; krzychu@ap.krakow.pl received on 30 June 2006 Abstract. The first record of M. blythii in Poland is described. It slightly extends the known distribution range of the species in Central Europe northwards. Three large species of the genus Myotis live in Europe. Two of them, the greater mouse-eared bat, M. myotis (Borkhausen, 1797), and the lesser mouse-eared bat, M. blythii (Tomes, 1857), are widely distributed throughout southern and central Europe. The distribution range of the third one, the Maghrebian mouse-eared bat, M. punicus Felten, 1977, is restricted to several Mediterranean islands, besides its main range in north-western Africa (Castella et al. 2000, Güttinger et al. 2001, Topal & Ruedi 2001). In Poland, M. myotis is widespread and its continual distribution range covers the southern and partly the western and central parts of the country. In addition, scattered records have been reported from most of the rest of Poland (Sachanowicz et al. 2006). However, M. blythii has not yet been reported from Poland. In Europe, the northern margin of its range reaches roughly 50 °N in the Czech Republic and Slovakia (Topál & Ruedi 2001), so the species has been known from near the southern border of Poland. Therefore, under favourable circumstances, the species was likely to occur also in Poland. During the field studies on swarming activity of bats in the Tatra and Beskidy Mts. carried out in 1999–2005, one individual of M. blythii was recorded. An adult male of M. blythii was captured on 12 October 2005 into a net installed inside the Czarna Cave, several dozens of metres below the northern cave entrance at the altitude of 1294 m, during an in-flight. This site is situated in the Organy Massif, in the Kościeliska Valley. The total length of known corridors of this cave is about 6,500 m, with denivelation of 303.5 m. It has three openings situated at 1326 m, 1294 m and 1404 m a. s. l. (Grodzicki et al. 1995). It is the largest bat hibernaculum in the Tatra Mts (Piksa & Nowak 2000). In order to identify the bat species, the basic morphometric features enabling to differentiate between M. myotis and M. blythii were considered: forearm length (FaL), ear length (EaL), and ear width (EaW) (see Arlettaz et al. 1991, Arlettaz 1995, Dietz & von Helversen 2004). Accroding to Arlettaz et al. (1991), the Z function was calculated: Z = 0.1084×FaL + 1.4166×EaL–40.5907 (if Z< 0, then M. blythii). At first sight, the captured bat seemed smaller and more subtle than M. myotis. As far as the coloration is concerned, it was similar to the earlier and later captured greater mouse-eared bats, with a little bit lighter fur on the belly. The white spot on the hair on the head, which is typical 197 of some M. blythii males, was not visible. The ears were narrower and thinner. The teeth of the captured bat were significantly rubbed off (e.g. the top left canine was broken in the distal part), indicating that it was an adult specimen. The right FaL was 58.1 mm, the left FaL 58.9 mm. The EaW and EaL of the right ear were 8.9 and 22.6 mm, respectively; the left EaW was 8.9 mm. The left EaL was not measured as its tip was slightly cut. The lengths of the forearms and ears of the individual caught were compared with the dimensions of other greater mouse-eared bats captured in the years 2004–2005 during the research on swarming activity in the Polish parts of the Carpathians (Fig. 1). The value of the formula compiled by Arlettaz et al. (1991) was –2.28 in this bat, i.e. the value typical for M. blythii, and differed from the values found in the captured individuals of M. myotis (0.21–3.34). In this respect the examined bat differed remarkably from the other mouse-eared bats captured at that time in the Polish Carpathians. The Czarna Cave is the first locality of M. blythii in Poland. However, the species does not seem to be a permanent element of the Polish fauna. Most probably, this was just an acciden tally present individual whose occurrence, especially in the Polish Tatra Mts, is surprising. On the southern side of the Tatra Mts. in Slovakia, the species has been recorded only at two sites during hibernation and at lower altitudes: in winter 1964 in the Belianska Cave (890 m a. s. l.) (Mošanský & Gaisler 1965, Gaisler & Hanák 1972) and in the winter season 1994/1995 in the Lučivianska Cave (800 m a. s. l.). However, the accuracy of identification of the latter record has been doubted (Pjenčák et al. 2003). Therefore, the last certain record of this species in the Slovak Tatra Mts. was made more than 40 years ago. Since then, despite intensive Fig. 1. Scatter plot of the forearm length against the ear length (n=63) recorded in Myotis myotis (triangles) and M. blythii (square) captured in 2004 and 2005 in the Polish Carpathians. Rys. 1. Relacja między długością przedramienia (Forearm length) a długością ucha (Ear length) (n=63) u Myotis myotis (trójkąty) i M. blythii (kwadrat) schwytanych w 2004 i 2005 roku w okresie swarmingu, w Tatrach, Beskidzie Wyspowym, Beskidzie Sądeckim, Beskidzie Niskim, Pogórzu Ciężkowickim i Podhalu. 198 research, especially in the recent years, both in the periods of activity and of hibernation, the species has not been found (Pjenčák et al. 2003). In other parts of Poland, in the regions close to the northern range of M. blythii, where its occurrence should be possible, it has not yet been recorded (author’s unpubl. data, Paszkiewicz et al. 1998, Postawa & Wołoszyn 2000, Węgiel et al. 2001, 2004). However, it cannot be excluded that single specimens of this species might occasionally occur in Poland. Streszczenie Dorosły samiec nocka ostrousznego Myotis blythii został odłowiony 12 października 2005 roku w Tatrach Polskich w Jaskini Czarnej (1294 m n. p. m.). To pierwsze stanowisko tego gatunku nietoperza w Polsce. Acknowledgements I would like to thank the members of the Caving Club of University of Science and Technology in Cracow, as well as Elżbieta Wiejaczka and Wojciech Gubała for their help in the field. This study was supported by the grant from the State Committee and Scientific Research. REFERENCES Arlettaz R., 1995: Ecology of the Sibling Mouse-eared Bats (Myotis myotis and Myotis blythii): Zooge ography, Niche, Competition, and Foraging. Ph.D. Thesis, University of Lausanne. Horus, Martigny, 206 pp. Arlettaz R., Ruedi M. & Hausser J., 1991: Field morphological identification of Myotis myotis and Myotis blythi (Chiroptera, Vespertilionidae): a multivariate approach. Myotis, 29: 7–16. Castella V., Ruedi M., Excoffier L., Ibáñez C., Arlettaz R. & Hausser J., 2000: Is the Gibraltar Strait a barrier to gene flow for the bat Myotis myotis (Chiroptera: Vespertilionidae)? Mol. Ecol., 9: 1761–1772. Dietz C. & von Helversen O., 2004: Illustrated Identification Key to the Bats of Europe. Electronic pub lication. Version 1.0, Tuebingen & Erlangen, 72 pp. Güttinger R., Zahn A., Krapp F. & Schober W., 2001: Myotis myotis (Borkhausen, 1797) – Großes Mausohr, Großmausohr. Pp.: 123–207. In: Krapp F. (ed.): Handbuch der Säugetiere Europas. Band 4: Fledertiere. Teil I. Chiroptera I. Rhinolophidae, Vespertilionidae 1. AULA-Verlag GmbH, Wiebelshe im, x+603 pp. Paszkiewicz R., Szkudlarek R., Węgiel A., Węgiel J. & Węgiel W., 1998: Materiały do chiropterofauny Pienin – letnie stanowiska nietoperzy [Materials about the Pieniny bats – summer bat sites]. Pieniny Przyr. Czł., 6: 31–46 (in Polish, with a summary in English). Piksa K. & Nowak J., 2000: The bat fauna of the Polish Tatra Caves. Pp.: 181–190. In: Wołoszyn B. W. (ed.): Proceedings of the VIIIth European Bat Research Symposium. Vol. 1. Approaches to Biogeogra phy and Ecology of Bats. Chiropterological Information Center, Kraków, 280 pp. Pjenčák P., Danko Š. & Matis Š., 2003: Netopiere Tatranského národného parku a širšieho okolia [Bats of the Tatra National Park and its wider surroundings (Northern-central Slovakia)]. Vespertilio, 7: 139–160 (in Slovak, with an abstract in English). Postawa T. & Wołoszyn B. W., 2000: Fauna nietoperzy Bieszczadów Zachodnich [The bat fauna of Western Bieszczady Mountains]. Monogr. Bieszczad., 9: 91–101 (in Polish, with a summary in English). Sachanowicz K., Ciechanowski M. & Piksa K., 2006: Distribution patterns, species richness and status of bats in Poland. Vespertilio, 9–10: 151–174. Topal G. & Ruedi M., 2001: Myotis blythii (Tomes, 1857) – Kleines Mausohr. Pp.: 209–255. In: Krapp F. (ed.): Handbuch der Säugetiere Europas. Band 4: Fledertiere. Teil I. Chiroptera I. Rhinolophidae, Vespertilionidae 1. AULA-Verlag GmbH, Wiebelsheim, x+603 pp. 199 Węgiel A., Paszkiewicz R. & Szkudlarek R., 2001: Nietoperze Beskidu Wyspowego, Beskidu Sądeckiego, Beskidu Niskiego i Pogórza Karpackiego – letnie schronienia nietoperzy w budynkach [Bats of Beskid Wyspowy, Beskid Sądecki, Beskid Niski and Pogórze Karpackie – summer roosts in buildings]. Nietoperze, 2: 75–84 (in Polish, with a summary in English). Węgiel A., Szkudlarek R. & Gottfried T., 2004: Wyniki odłowów nietoperzy przy otworach niektórych jaskiń w Beskidach [Species composition, activity and population structure of bats netted in summer at the entrances of some caves in Beskidy Mts]. Nietoperze, 5: 93–105 (in Polish, with a summary in English). 200 Lynx (Praha), n. s., 37: 201–228 (2006). ISSN 0024–7774 Areal and altitudinal distribution of bats in the Czech part of the Carpathians (Chiroptera) Plošné a výškové rozšíření netopýrů v české části Karpat (Chiroptera) Zdeněk Řehák Institute of Botany and Zoology, Masaryk University, Kotlářská 2, CZ–611 37 Brno, Czech Republic; rehak@sci.muni.cz received on 20 December 2006 Abstract. All available bat records from the Outer Carpathians, situated in the eastern part of the Czech Republic, were summarized, divided into old (before 1956), winter and summer records and evaluated with respect to orographical units, quadrats of grid maps and elevation. In total, 20 bat species and/or two pairs of sibling species were recorded in 9 orographical units and 41 quadrats covering the area under study. Hitherto, thirteen species of bats were recorded as hibernating and in 10 species maternity colonies were found in the Czech Carpathians. The areal distribution of particular species was presented via grid maps. Plecotus spp. (50.0% of the studied area), Myotis myotis (46.7%) and M. daubentonii (40.0%) can be considered the most distributed of them. Altitudinal distribution of sites where bats were recorded, expressed as medians of their elevation, shows that P. austriacus significantly preferred low elevation both in the winter and summer periods (259 and 298 m above sea level, respectively) while the hibernacula of the M. mystacinus group and M. nattereri prevailed at higher elevations (872 m and 1050 m, respectively). During the non-hibernation period the highest medians of elevation were recorded in M. brandtii and again in M. nattereri (765, resp. 665 m a. s. l.). The causes of preferences for such high elevated sites are discussed. Introduction The knowledge of the distribution of bats enables to estimate their species diversity over reasonable geographical units and, consequently, to assess the ecological value of individual segments within a given territory. At the same time, bats’ distribution reveals the differences in conditions favourable to the life of these mammals among various areas of the same territory. Differences in the composition and abundance of bat communities then reflect different habitat demands of individual species the communities consist of. Information on the distribution of bats, therefore, represents a starting point for both synecological and autecological studies of the respective chiropteran fauna. Faunistic review of all available records of bats in a particular area is a condition sine qua non to analyse the distribution of the respective bat assemblage or assemblages. Within the territory of the Czech Republic, the Carpathians were relatively neglected in this respect until quite recently. The history of chiropterological research there, not very ample so far, was dealt with in previous papers by the author (Řehák 1998, 2001). The first faunistic review of bats on the territory of the Czech (Moravian-Silesian) Carpathians was published by Řehák (1998). 201 In the present paper, the data of the former one gathered up to 1998 are supplemented by new records till 2006, both published and unpublished, to get a real image on the bat fauna of the easternmost part of the Czech Republic. New data were obtained mainly by the results of systematic research in the Hostýnské vrchy Mts (Lučan 2000, Lučan & Svačina 2001), of acoustic detectoring in Wallachia (Řehák 2001 and unpublished) or of an intense research of bat activity at the Hranická propast Chasm, including a maternity colony roosting in a cave of the chasm (Baroň & Řehák 2001, Řehák & Baroň 2002 and unpublished). Further data resulting from long-term monitoring of hibernacula, Myotis myotis maternity colonies and bat-detectoring of forest bats and sibling species Pipistrellus pipistrellus and P. pygmaeus (Řehák et al. 2005) were included as well. Numerous data, even from regions omitted so far, were obtained thanks to the improvement of species determination from bat detector samples, by tape recording the ultrasound signals and their subsequent digitalization and computer analysis. Although mountain and hilly elevations (up to 1323 m a. s. l.) prevail on the territory, large underground spaces serving to bats as their natural hibernacula are relatively rare there, compared to similar mountain regions elsewhere. Warm pseudo-karstic caves at higher elevations, situated mainly in the eastern part of the territory, are exception. Some of them have been known for years as bat hibernacula (Rumler 1985, Baroň & Řehák 1997, Wagner 2001). Neither the mining was common in the Moravian-Silesian Carpathians, except for a few slate quarries and galleries (Kirchner & Řehák 1990, Řehák & Baroň 1997, Šafář & Rumler 2001). Winter records of bats are particularly rare in the southern and western hilly grounds where only subterranean vaults and cellars of houses are known as bat hibernacula. In contrast, summer records of bats are relatively numerous. They concern checks of bats roosting in lofts, mainly of sacral buildings and castles, netting of bats in the field and, during the last 15 years, acoustic bat-detectoring. It was demonstrated that the occurrence of bats in summer was not limited to warmer lower elevated sites with a higher food supply, but bats occurred also at higher elevations, mainly in the second half of summer. Records of bats were made close to underground spaces where some of them later hibernate, bat detectoring in the massif of the Lysá hora Mt. revealed bats foraging on the mountain ridges at the elevation of 900 m and more. There are two goals of the present paper: (1) to evaluate all records and to show the geographical distribution of bat species on maps with a standard girid, and (2) to analyse the altitudinal distribution of individual species with respect to main periods of their life cyle. Material and methods Study area As in the previous paper (Řehák 1998), the territory on the right bank of the Morava River was excluded from the study area (Fig. 1). The area under study consists of 10 orographic units which are, approximately in the north-south direction: Slezské Beskydy Mts, Jablunkovská brázda Furrow, Jablunkovská vrchovina Highland, Moravskoslezské Beskydy Mts, Podbeskydská pahorkatina Upland, Rožnovská brázda Furrow, Javorníky Mts, Hostýnsko-vsetínská hornatina Mts, Vizovická vrchovina Highland and Bílé Karpaty Mts (Demek & Střída 1971). No research concerning bats was made in only one of them, the Jablunkovská brázda Furrow, and there have been no records of bats there. The whole territory with its orographic and hydrologic situation is shown by a map (Fig. 2). Mountains at the border between the Czech Republic and Slovakia represent the highest part of the area, the elevation then decreases towards the west down to the river valleys of Bečva and Morava. The elevation decreases also in the southward direction where it is difficult to find the border between the Carpathians and the Dolnomoravský úval Valley. 202 Table 1. List of bat species recorded in the area of the Czech Carpathians and abbrevations for bat taxa Tab. 1. Seznam druhů netopýrů zjištěných na území českých Karpat a zkratky jejich názvů species Rhinolophus hipposideros Myotis mystacinus Myotis brandtii M. mystacinus seu M. brandtii Myotis emarginatus Myotis nattereri Myotis bechsteinii Myotis myotis Myotis oxygnathus Myotis daubentonii Vespertilio murinus Eptesicus nilssonii abb. Rhip Mmys Mbra Mmys/bra Mema Mnat Mbec Mmyo Moxy Mdau Vmur Enil species Eptesicus serotinus Nyctalus noctula Nyctalus leisleri Pipistrellus pipistrellus Pipistrellus pygmaeus P. pipistrellus seu P. pygmaeus Pipistrellus nathusii Barbastella barbastellus Plecotus auritus Plecotus austriacus P. auritus seu P. austriacus abb. Eser Nnoc Nleu Ppip Ppyg Ppip/pyg Pnat Bbar Paur Paus Paur/aus Fig. 1. Map of Moravia including the W-Carpathian area under study. Obr. 1. Mapa Moravy s vyznačením studovaného území Západních Karpat. Explanations / vysvětlivky: A – Bohemian Highlands / Česká vysočina; B – Western Carpathians / Západní Karpaty; a – Outer Carpathian Depressions / Vněkarpatské sníženiny; b – Outer Carpathians / Vnější Karpaty; c – Inner Carpathian Depressions / Vnitrokarpatské sníženiny; thick line / silná čára – state border / stární hranice; middle line / středně silná čára – geomorphological system borders / hranice geomorfologických provincií; thin line / tenká čára – geomorphological subsystem border / hranice geomorfologických podprovincií; chequered area / kostkovaná plocha – study area / studované území. 203 Material A data base has been developed from all available sources since the twentieth of the 20th century until the year 2006. It has 1447 items with records of 20 bat species. The species, including abbreviations of their names, are listed in Table 1. Records with unspecified location or timing were excluded from the evaluation of bats’ distribution. Mostly they concern old data (Remeš 1927, Gaisler 1956, Hanák 1960). In addition to previous publication (Řehák 1998), the data base was completed by records published in the papers by Lučan (2000), Lučan & Svačina (2001), Šafář & Rumler (2001), Wagner (2001), Jahelková (2003) and, in certain species, missing data were assumed from Hanák & Anděra (2005). Unpublished own data were added too. Three categories of data are recognized. The first category is represented by old records before the year 1956 when the first thorough faunistic review of bats in the then Czechoslovakia was published (Gaisler 1956). The records made since 1956 were divided into winter and summer ones. The second category (winter records) are those made in hibernacula and concerning torpid bats. In most cases, winter season has formally been fixed to 15 October – 30 April (Gaisler et al. 1988, cf. Hanák & Anděra 2005). Irrespectively of this time span, findigs of torpid bats in May in hibernacula situated at high elevations were considered winter records as well. The third category (summer records) comprises mostly the findings made from 1 May to 14 October but, as in the case of winter records, this delimitation was not absolutely observed. Finds and observations outside a shelter, e.g., made at the beginning of April or at the end of October, were ranged to this category. Records of colonies were specified and evaluated separately from the rest of summer records. Except in Vespertilio murinus, they concerned maternity colonies. All records Fig. 2. Geographical map of the area under study. Obr. 2. Geografická mapa studovaného území. 204 were assigned to the orographic units, quadrats of the mapping grid and their elevation was determined (see Řehák 1998). Methods Various methods were used to get the data. Winter records were made either by occasional checks in house cellars and vaste subterranean vaults of castles (Hanák 1960, Vlašín et al. 1993, 1995), or by regular winter monitoring in former mine galleries (Baroň & Řehák 1997) and natural pseudo-karstic clefts (ibid., Wagner 2001). All winter records of Vespertilio murinus came from above ground rooms of buildings (Řehák 1998, Kašpar in litt.). Summer records were made by searching in lofts and under roofs of buildings, namely churches and castles, by netting at the entrances to underground or on potential bat hunting grounds, namely above brooks and on banks of water reservoirs. Important data have also been obtained by the collection and analysis of owl pellets. Except in some cases, the detection of echolocation signals yielded important data. Acoustic determination was often limited in cases where species identification based on external morphology was difficult as well. In sibling species such as M. mystacinus and M. brandtii, or P. auritus and P. austriacus, visual identification from a distance is problematic (cf. Bar tonička 2004), in Pipistrellus bats it is nearly impossible vithout manipulation of them. In P. pipistrellus and P. pygmaeus their identification based on morphology is problematic in any case (Řehák et al. 2005) but they can be recognized by detecting their ultrasound signals. Acoustically based determination of the two Pipistrellus species mentioned above, however, has been done only since 2000, some earlier records therefore have to be assigned to Pipistrellus pipistrellus sensu lato. The relatively recently discovered species P. pygmaeus (Jones & van Parijs 1993, Barratt et al. 1997) was acousticly detected only in three places at lower elevations on the periphery of the territory studied (own unpublished data). Therefore, records prior to 2000 made at higher elevations are assigned to P. pipistrellus sensu stricto. Areal distribution of records belonging to the respective category was visualized with the aid of the mapping grid (cf. Řehák et al. 2003). Relative representation of each species is then calculated as a percent of quadrats where it was recorded out of the total of all quadrats of the area. Altitudinal distribution was evaluated after the variance of elevations of the localities, divided again according to summer records, winter records and records of maternity colonies. Non-parametric tests were applied, the Kruskal-Wallis H test to the comparison among species and the Mann-Whitney U test to compare species pairs. All statistical operations were made using the software Statistica for Windows. Results Bat fauna in particular orographical units The number of localities with records of bats in particular orographic units, supplemented by the number of recorded species, are given in Table 2. It is evident from the table that the Podbeskydská pahorkatina Upland and the Hostýnsko-vsetínská hornatina Mts have the most rich bat fauna. Both orographic units are situated in the north-western part of the territory which gradually fades away into the lowlands of Outer Carpathians. There the number of both hibernacula and localities with summer occurrence of bats is the highest which is reflected in the highest number of bat species in winter as well as in summer. In contrast, Slezské Beskydy Mts and Jablunkovská vrchovina Highland in the northern part of the territory have the lowest number of both localities and bat species. In most orographic units, the number of localities with summer records is higher than that with winter records (Table 2). All localities arranged according to their situation in individual orographical units and quadrats are listed in Appendix 1. 205 Table 2. The total number of bat species and localities with records of bats in particular orographical units Tab. 2. Celkový počet druhů netopýrů a lokalit s nálezy netopýrů v jednotlivých orografických celcích Explanations / vysvětlivky: NoL – number of localities / počet lokalit; NoS – number of bat species / počet druhů netopýrů; W – winter season – records of torpid bats at hibernacula and all winter records of V. murinus / zimní období – nálezy letargujících netopýrů na zimovištích a všechny zimní nálezy V. murinus; G – growing season – other records of bats, especially from summer / mimohibernační období – ostatní nálezy netopýrů, především letní orographical unit / orografická jednotka W NoL G W NoS G W+G Slezské Beskydy Mts Jablunkovská vrchovina Highland Moravskoslezské Beskydy Mts Podbeskydská pahorkatina Upland Rožnovská brázda Furrow Javorníky Mts Hostýnsko-vsetínská hornatina Mts Vizovická vrchovina Highland Bílé Karpaty Mts 0 1 6 10 1 1 9 9 2 2 0 20 53 25 12 41 30 34 0 3 9 10 1 4 11 8 4 3 0 14 17 15 14 18 13 13 3 3 14 17 15 14 18 13 14 Areal distribution The area studied covers altogether 60 quadrats of the mapping grid but 19 of them by less than 50% only. Some of the peripheral quadrats belong to the area by a very small territory. There are no records of bats in 19 quadrats, 15 of them are the same as above. Bats were recently found in all quadrats with old records prior to 1956. Winter records cover 19 quadrats (31.6%), in one of them, 6478, only hibernating bats were recorded (Na Gírové cave in the Jablunkovská vrchovina Highland). There are 6 quadrats (10%) with five and more hibernacula in each or with an important hibernaculum where 10 or more individuals of one bat species were recorded during a single check. The most important natural hibernacula include the pseudo-karstic cleft Fig. 3. Distribution maps of individual bat species or pairs of sibling species. Fig. 3. Mapy rozšíření jednotlivých druhů nebo dvojic podvojných druhů netopýrů. Explanations / vysvětlivky: empty square / prázdný čtverec – records only before 1956 / jen nálezy před rokem 1956; black square or semisquare / černý čtverec nebo půlčtverec – records of nursery colonies / nálezy letních reprodukčních kolonií; grey square / šedý čtverec – records of male colonies (only V. murinus) / nálezy samčích kolonií (jen V. murinus); little empty circle or semicircle / malý prázdný kruh nebo půlkruh – records of less than 10 individuals in 1–4 hibernacula / nálezy méně než 10 jedinců na 1–4 zimovištích; big empty circle or semicircle / velký prázdný kruh nebo půlkruh – 5 or more hibernacula or records of 10 or more individuals in one hibernaculum during one check at least / 5 a více zimovišť nebo nálezy 10 a více jedinců nejméně na 1 zimovišti během jedné kontroly; little full circle or semicircle / malý plný kruh nebo půlkruh – less than 5 sites of summer records / méně než 5 lokalit s letními nálezy; big full circle or semicircle / velký plný kruh – 5 or more sites of records of bats or records of 10 or more individuals (excl. colonies) at a site per one survey during growing season / 5 a více lokalit s nálezy netopýrů nebo nálezy 10 a více jedinců (vyjma kolonií) na lokalitě při jedné akci během mimohibernačního období. 1 – Rhinolophus hipposideros, 2 – Myotis mystacinus, 3 – Myotis brandtii, 4 – Myotis mystacinus seu M. brandtii, 5 – Myotis emarginatus, 6 – Myotis nattereri. 206 1 2 3 4 5 6 207 Table 3. Areal distribution of particular bat species or species groups Tab. 3. Plošné rozšíření jednotlivých druhů netopýrů či jejich skupin Explanations / vysvětlivky: NoQ – number of quadrates where bats were recorded (Myotis mystacinus group includes M. mystacinus and M. brandtii, Pipistrellus pipistrellus group includes P. pispistrellus and P. pygmaeus and Plecotus group includes P. auritus and P. austriacus) / počet kvadrátů s nálezy netopýrů (Myotis mystacinus group zahrnuje M. mystacinus a M. brandtii, Pipistrellus pipistrellus group zahrnuje P. pipistrellus a P. pygmaeus a Plecotus group zahrnuje P. auritus a P. austriacus) species Rhinolophus hipposideros M. mystacinus group Myotis emarginatus Myotis nattereri Myotis bechsteinii Myotis myotis Myotis oxygnathus Myotis daubentonii Vespertilio murinus Eptesicus nilssonii Eptesicus serotinus Nyctalus noctula Nyctalus leisleri P. pipistrellus group Pipistrellus nathusii Barbastella barbastellus Plecotus group winter season NoQ % 15 8 6 3 4 10 0 7 2 2 3 0 0 0 0 3 14 25.0 13.3 10.0 5.0 6.7 16.7 0 11.7 3.3 3.3 5.0 0 0 0 0 5.0 23.3 growing season NoQ % 19 16 10 9 9 26 1 22 6 13 18 15 9 21 5 10 24 31.7 26.7 16.7 15.0 15.0 43.3 1.7 36.7 10.0 21.7 30.0 25.0 15.0 35.0 8.3 16.7 40.0 total NoQ % 20 18 13 9 9 28 1 24 6 13 19 15 9 21 5 10 30 33.3 30.0 21.7 15.0 15.0 46.7 1.7 40.0 10.0 21.7 31.7 25.0 15.0 35.0 8.3 16.7 50.0 caves in the Moravskoslezské Beskydy Mts (Cyrilka, Kněhyňská jeskyně and Ondrášovy díry caves), Javorníky Mts (Pod kazatelnou I Cave) and Vizovická vrchovina Highland (three caves in the hill Kopce near Lidečko). Important artificial hibernacula include the Sintrová Gallery near Liptál (Vizovická vrchovina Highland), the subterranean vault of the Holešov Castle and the cellars of buildings in Lukov. Except the Kněhyňská and Ondrášovy díry caves dominated by Myotis myotis, all important hibernacula harbour Rhinolophus hipposideros as the most abundant species in winter. Summer records cover altogether 40 quadrats (66.7%); maternity colonies were found in 24 of them (40.0%). Summer male colonies of Vespertilio murinus were recorded behind window shutters at two localities in the Hostýnsko-vsetínská hornatina Mts (quadrats 6572, 6672). In addition to the quadrats with records of bat colonies, there are 6 quadrats with at least five localities in each or with an important summer locality where 10 or more individuals of one bat species were recorded during a single check. Numerous bats were netted in late summer at the entrances to caves serving also as hibernacula in three of the quadrats (6575, 6672, 6774). Bats Fig. 3. Distribution maps of individual bat species or pairs of sibling species (continued). Fig. 3. Mapy rozšíření jednotlivých druhů nebo dvojic podvojných druhů netopýrů (pokračování). 7 – Myotis bechsteinii, 8 – Myotis myotis, 9 – Myotis oxygnathus, 10 – Myotis daubentonii, 11 – Vespertilio murinus, 12 – Eptesicus nilssonii. 208 7 8 9 10 11 12 209 were recorded at more localities in each of the remaining three quadrats. There the records were made by either systematical bat detectoring (quadrats 6377, 6574) or by regular mist netting (6472 – Hranická propast Chasm). The distribution of records of individual bat species according to three categories of data reflecting the time period, number of localities and number of bats is shown in Fig. 3 (1–23). The data on areal distribution of all species are summarized in Table 3. In cases when sibling species were not distinguished from each other or their determination was questionable, they were lumped together in the table. With respect to the absolute and relative number of quadrats with bat records, the species or groups of species with the largest distribution are Plecotus spp., Myotis myotis and M. daubentonii evidenced on at least 40% of the territory. In addition to them, Pipistrellus pipistrellus s. l. and Rhinolophus hipposideros were recorded on one third or a little more of the territory. In contrast, Pipistrellus pygmaeus (3 quadrats and localities) and P. nathusii (5 quadrats and 6 localities) are very rare. Myotis oxygnathus, recorded only once and in one locality, is actually the rarest known bat. In the hibernacula, R. hipposideros and Plecotus spp. have the largest distribution covering ca 25% of the territory. The only other species covering more than 15% of the territory is M. myotis. There are no records of hibernating individuals in Nyctalus and Pipistrellus species. Summer records of M. myotis, Plecotus spp., M. daubentonii and P. pipistrellus s. l. cover more than 1/3 of the territory. Other species common in summer are R. hipposideros and Eptesicus serotinus. In E. nilssonii and Barbastella barbastellus the summer records cover a significantly larger area than the winter ones. Altitudinal distribution Localities situated at the lowest elevations comprise bat hibernacula in house cellars in the Podbeskydská pahorkatina Upland and Vizovická vrchovina Highland and a wine-vault in the slopes of Bílé Karpaty Mts, all under 250 m a. s. l. Single P. austriacus, in one case P. auritus, were recorded in a total of six cellars. In contrast, five pseudo-karstic caves in the massif of highest peaks of the Moravskoslezské Beskydy Mts, i. e. Lysá hora, Kněhyně and Radhošť, represent hibernacula situated at the highest elevations (945 – 1100 m), absolutely highest situation having the Čertova díra Cave, 1100 m a. s. l. However, bats go up to such elevations even in summer. In the Moravskoslezské Beskydy Mts, they were visually observed on the Malá Stolová Mt. and acousticly detected on Lysá hora Mt., both at 900 m a. s. l. or higher and in late summer bats were netted at the entrances of three pseudo-karstic caves (945, 1050 and 1050 m a. s. l). The range of elevations and their median are shown in Fig. 4 (a–c). The variance of elevations is given for individual species as well as for the whole sample. The sample consists of 38 hibernacula and 14 bat species or species groups but M. bechsteinii, V. murinus, E. serotinus and P. auritus seu austriacus were recorded in less than five hibernacula and were excluded from the evaluation. The species M. mystacinus and M. brandtii were lumped together due to the problems of their differentiation. Significant differences among the species were revealed in the distribution of elevations of their hibernacula [Kruskal-Wallis test, H(9, N=112)=33.3, Fig. 3. Distribution maps of individual bat species or pairs of sibling species (continued). Fig. 3. Mapy rozšíření jednotlivých druhů nebo dvojic podvojných druhů netopýrů (pokračování). 13 – Eptesicus serotinus, 14 – Nyctalus noctula, 15 – Nyctalus leisleri, 16 – Pipistrellus pipistrellus, 17 – Pipistrellus pygmaeus, 18 – Pipistrellus pipistrellus seu P. pygmaeus. 210 13 14 15 16 17 18 211 Fig. 4a. Box-plots of altitudinal distribution of hibernacula. Abbreviations – see Table 1. Obr. 4a. Krabicové diagramy výškové distribuce zimovišť. Zkratky – viz tab. 1. p=0.0001]. From among all species, P. austriacus hibernates at the lowest situated localities (median 259 m a. s. l.), being the only species evaluated in which the median of altitude lies below 400 m. The distribution of elevations of hibernacula with P. austriacus differs significantly from that in other 9 species (Mann-Whitney tests: U=3.5–27.0, Z=2.17–4.11, p<0.0301). On the contrary, M. mystacinus seu M. brandtii and M. nattereri hibernate at the highest situated localities (median 872 m a. s. l. and 1050 m a. s. l., respectively). The differences between the elevations of hibernacula of M. nattereri and other species except P. austriacus are insignificant (Mann-Whitney tests) due to the small sample size. The differences between the elevations of hibernacula in the M. mystacinus group and P. austriacus as well as R. hipposideros and B. barbastellus are significant (Mann-Whitney U tests: U=3.5, Z=3.814, p=0.000134, U=53.0, Z=2.532, p=0.0113 and U=9.0, Z=1.967, p=0.049). The medians of elevations of hibernacula range from 400 to 800 m a. s. l. in 7 species, being 400–600 m in R. hipposideros and B. bar bastellus (415 and 440 m, respectively) and 600–800 m in the remaining 5 species (Fig. 4a). Fig. 3. Distribution maps of individual bat species or pairs of sibling species (continued). Fig. 3. Mapy rozšíření jednotlivých druhů nebo dvojic podvojných druhů netopýrů (pokračování). 19 – Pipistrellus nathusii, 20 – Barbastella barbastellus, 21 – Plecotus auritus, 22 – Plecotus austriacus, 23 – Plecotus auritus seu P. austriacus. 212 19 20 22 21 23 213 Fig. 4b. Box-plots of altitudinal distribution sites with occurrence of bats during growing season. Abbreviations – see Table 1. Obr. 4b. Krabicové diagramy výškové distribuce lokalit s výskytem netopýrů v mimohibernačním období. Zkratky – viz tab. 1. In 240 localities 22 species or species groups were recorded in summer. Three couples of sibling species and P. pygmaeus (3 localities only) are excluded from the evaluation. In 11 out of the 18 evaluated species, the maximum median of elevations is 400 m, it exceeds this value in the remaining species (Fig. 4b). Significant differences were found in the distribution of elevations of summer localities among the species [Kruskal-Wallis test: H(17, N=435)=65.6, p<0.0001]. As in the hibernacula, the lowest median of summer elevations concerns P. austriacus (298 m). Again, the distribution of elevations of summer localities with P. austriacus differs significantly from that in most other species (Mann-Whitney tests: U=4.0–338.0, Z=2.264–5.159, p<0.0236), the only exception being P. nathusii (p>0.05). The distribution of elevations of P. nathusii summer localities (median 332 m) differs significantly only when compared with that of M. brandtii (U=5.0, Z=2.286, p=0.0223), M.bechsteinii (U=14.5, Z=2.149, p=0.0317) and E. nilssonii (U=37.0, Z=2.307, p=0.0210). The highest values of medians concern the elevations of localities in M. brandtii (765 m) and M. nattereri ( 665 m). The difference between the data of M. brandtii and that of most other species are significant (Mann-Whitney tests: U=3.5–50.0, Z=2.185–3.780, p<0.0289), exceptions are that of M. nattereri and M. bechsteinii, both insignificant. The distribution of elevations of M. nattereri summer localities differs significantly from that in 9 species (Mann-Whitney tests: U=22.5–195.5, Z=2.239–3.691, p<0.0252), while the differences against the remaining 8 species are insignificant. Following are the results when 214 comparing two related species of the same genus: P. auritus prefers higher situated summer localities than P. austriacus (U=125.0, Z=3.985, p=0.000068), M. brandtii than M. mystacinus (U=30.0, Z=2.308, p=0.0210) and E. nilssonii than E. serotinus (U=363.5, Z=2.849, p=0.00438). Only the differences between elevations of summer localities in N. leisleri and N. noctula are insignificant. No significant differences, probably due to a small sample size, were revealed when comparing the elevations of 43 shelters of summer colonies among 9 bat species (Kruskal-Wallis test). B. barbastellus was not evaluated due to the evidence of only one colony. Maternity colonies at the highest elevations were found in E. nilssonii (n=3, median 500 m a. s. l.), maternity colonies at the lowest elevations in P. austriacus (n=4, median 295.5 m a. s. l.). Discussion So far, 20 bat species were recorded on the studied territory belonging to the province of W Carpathians. At least 13 of them were found hibernating, this number being similar to that of species recorded in hibernacula of near Oderské vrchy Mts where 11 bat species were recorded (Šafář & Rumler 2001, Wagner 2001, 2004). No winter occurrence of Myotis brandtii was recorded Fig. 4c. Box-plots of altitudinal distribution of sites with occurrence of nursery colonies. Abbreviations – see Table 1. Obr. 4c. Krabicové diagramy výškové distribuce lokalit s výskytem reprodukčních kolonií. Zkratky – viz tab. 1. 215 so far but it is probable. When checking torpid bats in hibernacula visually without touching them it is impossible to distinguish safely between M. brandtii and M. mystacinus and, therefore, M. brandtii could be overlooked between bats labelled as M. mystacinus (cf. Bartonička 2004). Most records of M. brandtii concern bats netted in late summer at entrances to caves where the bats later hibernate, including M. mystacinus, which corroborates the hypothesis of M. brandtii hibernation in the area studied. Due to this potential confusion, all winter records of M. mystacinus were considered records of the M. mystacinus group. Another confusion concerning M. brandtii possibly results from the fact that the localities of its netting records are situated at high elevations, M. brandtii thus appearing to be a mountain species. Similar problem of distinguishing in winter concerns the siblings P. auritus a P. austriacus and, accordingly, their winter records were consider records of the P. auritus group. V. murinus, although recorded in winter, was never found in an underground hibernaculum or even in a deep torpidity. Its recods invariably concern single individuals in rooms of buildings above ground or observations of flying bats. Compared to previous data (Řehák 1998) new hibernating species was found, viz., E. serotinus which was repeatedly recorded in castle vaults of Bystřice pod Hostýnem and Holešov (Lučan & Svačina 2001). So far, no winter records of Nyctalus and Pipistrellus bats in the area of study exist. Since bats of the genus Nyctalus do not hibernate underground, they possibly either migrate to warmer regions of southern Europe or they hibernate in large tree cavities. N. noctula, however, is known to hibernate in rock crevices and in various fissures under the roofs and in the outer walls of buildings including the relatively newly built blocks of prefabricated panel houses in towns and cities (Gaisler 1997, Lehotská & Lehotský 2000). Such records are missing in the area so far. Low number of winter records of psychrophilous hibernants, namely B. barbastellus and E. nilssonii, is probably due to the character of pseudo-karstic caves and galleries, all of them being relatively warm in winter with temperatures up to 8 °C (Baroň & Řehák 1997), which are too high to ensure their winter torpor. On the other hand, such hibernacula suit well to species which are dominant in the study area – M. myotis and mainly R. hipposideros. Both species were found to be relatively abundant in all larger hibernacula (Řehák 1998, Wagner 2001). The situation is very different from that in the near Oderské vrchy Mts and Jeseníky Mts with vaste systems of abandoned galleries where winter temperature is close to 0 °C at places and both B. barbastellus and E. nilssonii are very abundant (Rumler 1985, Řehák & Gaisler 1999, Wagner 2001, 2004). At the same time, the hibernating bat assemblages are much larger in that two mountain systems and include high numbers of both eurytherm M. myotis and thermophile R. hipposideros (Buřič & Šefrová 2001, Řehák & Gaisler 2001, Šafář & Rumler 2001). The pseudo-karstic cleft caves of the Moravskoslezské Beskydy Mts are among the highest situated bat hibernacula in the Czech Republic. In four of them, their elevation is above 1000 m. There is only one further hibernaculum in the Czech Republic situated at such elevation, in the Jeseníky Mts (Souček 1969). All 20 bat species evidenced in the area of study were recorded in the summer season. Compared to previous data on bats in the Czech Carpathians (Řehák 1998), P. pygmaeus is a new species which can be reliably determined in the field by the detection of its ultrasound signals (Jones & van Parijs 1993). So far, this species was recorded in only three localities at low elevations, invariably close to water streams or reservoirs (cf. Řehák et al. 2004, 2005). Its sibling P. pipistrellus, in contrast, is abundant from lowlands to mountains over 900 m a. s. l. (Řehák ibid.). As the probability of former recording P. pygmaeus at medium and higher elevated localities is negligible, most older records are considered to represent the species P. pipistrellus. Reproduction was evidenced in 10 bat species in which maternity colonies were 216 recorded. Although some of the colonies disappeared during the research the number of reproducing bat species is still high with respect to the relief of the area studied. Most maternity colonies were recorded at low elevations, only three colonies of two species (E. nilssonii and M. myotis) were discovered in the lofts of buildings at elevations higher than 500 m. In the mounatins, maternity colonies were typically coupled to human settlements in river valleys. The largest number of maternities (20) was found in Myotis myotis, at the same time being the largest bat assemblages in the region (Řehák 1998). As elsewhere, large lofts of churches and castles shelter maternity summer colonies of M. myotis. Large maternity colony in the nearly inaccessible (to humans) Rotunda Cave belonging to the system of Hranická propast Chasm is the only exception. It is the only maternity colony of M. myotis known from a cave in the Czech Republic (Baroň & Řehák 2001, Řehák & Baroň 2002). This M. myotis summer colony is unique in many respects, e.g. concerning its size: the maximum number of pregnant females (young excluded) was estimated to 1,044 individuals, that of lactating females plus young to more than 2,000 individuals (Řehák 2006, unpublished). With respect to the landscape of the area studied, the total of 20 bat species is high and ranges well to the lowland and middle elevated regions where bats have been studied for a long time. There were only a few mountain systems in the Czech Republic the chiropteran fauna of which was studied systematically, namely the Šumava Mts (17 species, Anděra & Červený 1994), Žďárské vrchy Mts (15 species, Řehák et al. 2000) and Železné hory Mts (13 species, Benda et al. 1997). Nineteen bat species were reported from the Krkonoše Mts (Anděra et al. 1974) but the records of two of them (N. leisleri, R. ferrumequinum) are very old and doubtful. Recently, faunistic data on bats were published from eastern Bohemia (Lemberk 2004) and southwestern Moravia (Reiter et al. 2003). In the former region, 21 bat species and 14 species with maternity colonies were recorded. In the latter region, 19 bat species were evidenced of which in 17 the reproduction was proved. The only large region with a more rich chiropterofauna is represented by southern Moravian lowlands in which 22 or 21 species (if one excludes the problematic N. lasiopterus) were recorded and in 11 species reproduction was evidenced (Gaisler et al. 1990, 2002, Řehák et al. 2003). Concerning the lowlands of northern Moravia which, similarly to southern Moravian lowlands, are situated close to the Carpathians, there are faunistic data on bats from the Poodří and Litovelské Pomoraví. In the lowlands of Poodří, 17 species including P. pygmaeus (Řehák & Bryja 1998, unpubl. data) and in that of Litovelské Pomoraví as many as 18 species were recorded (Bartonička et al. 2002). It can be concluded that the regions are mutually similar concerning the numbers of their bat species. In the orographic units, the representation of localities and bat species is uneven. There may be double reasons of it, the differences in their geomorphology, mainly in the elevation, and the differences in research activity concerning the bat faunistics. Important bat hibernacula in the Moravskoslezské Beskydy Mts, Hostýnsko-vsetínská hornatina Mts and Vizovická vrchovina Highland were systematically studied, while hibernacula suitable to such a study are missing in the southern part including the Bílé Karpaty Mts. Summer records reflect mainly the search of bats in large lofts of churches and castles which also are unevenly distributed, this fact again may be responsible of the differences between the orographic units concerning their bat fauna. The rank of species most common from the point of view of areal distribution is similar as in other regions where the mapping grid was applied. The Czech Carpathians (60 quadrats in total) can be compared with the Šumava Mts (50 quadrats, Anděra & Červený 1994), eastern Bohemia (68 quadrats, Lemberk 2004) and southwestern Moravia (38 quadrats, Reiter et al. 2003). Generally, the quadrats of Carpathians are covered less by the records of bats than that 217 in other regions mentioned above what can be due to both worse living conditions for bats and lesser activity of researchers in the Carpathians (Řehák 1998, 2001). Plecotus species were recorded on 50% of the Carpathian territory but their abundance has to be estimated with caution since the taxon comprises two species. Further bats common in the Carpathians are M. myotis and M. daubentonii. The situation is similar in eastern Bohemia and southwestern Moravia. In eastern Bohemia, P. auritus and M. myotis are the most distributed species while P. austriacus ranges fifth; in southwestern Moravia, P. austriacus ranges first followed by M. myotis while P. auritus together with M. daubentonii occupy the third and fourth places. The differences in abundance of P. austriacus and P. auritus in eastern Bohemia and southwestern Moravia probably reflect different geomorphological situation and climatic conditions between these two regions. While P. austriacus prefers warmer open lowland, P. auritus is more common in colder woodland of highlands and mountains. As shown by this paper, in the Carpathians the former species is more common in the warmer south at low elevation and few woods, the latter is more common at higher elevated woodland of the northeast. M. daubentonii followed by P. auritus represent the two most common species in the Šumava Mts where P. austriacus, by the number of quadrats where it was recorded, ranges as far as sixth. This can easily be explained by the lack of warm Pannonian lowlands in the region of Šumava and its foothills. In southern Moravia, in contrast, the impact of warm lowland habitats on the chiropteran fauna is evident (Reiter et al. 2003). When evaluating the altitudinal distribution, uneven representation of most bat species is striking. Often bats were recorded in lowland habitats and then relatively high in the mountains. This in part reflects the situation of most important and regularly checked hibernacula at higher elevations. Therefore the average elevation of winter bat records is higher than that of summer records. Nevertheless, certain species were recorded at high elevations even in summer as in the case of acoustic detection of bats foraging on the massif of the Lysá hora Mt., more than 900 m a. s. l. (Řehák & Bartonička, unpubl.). Summer netting at the entrances to high situated caves of the Beskydy Mts then raised the upper limit of summer occurrence in 10 bat species up to 1050 m a. s. l. which influenced the hypsometric distribution mainly of rare species difficult to monitor by methods other than netting (M. brandtii, M. nattereri). In contrast, summer maternity colonies were significantly more often found at lower elevations. In any study of the distribution of bats over a large territory, the impact of the respective scientist or scientists can not be excluded: the number of records is not only a function of bats’ occurrence but also of human activity. The number of records reflects the extent of monitoring in both space and time. Acoustic bat-detectoring, in spite of certain limits (difficult or even impossible determination of some species), substantially increases the capacity of field work with bats enabling to monitor even less attractive habitats and localities neglected so far. Souhrn V práci jsou shrnuty dostupné údaje o výskytu netopýrů pocházejících z území Vnějších Karpat ve východní části České republiky. Nálezy jsou rozděleny na zimní, letní a na nálezy letních kolonií a poté hodnoceny podle jejich příslušnosti k orografickým jednotkám, kvadrátům síťové mapy a podle nadmořské výšky. Celkem bylo v 9 orografických jednotkách a 41 kvadrátu, pokrývajících studované území, evidováno 20 druhů netopýrů a dvě dvojice podvojných druhů. Dosud bylo na zimovištích v českých Karpatech zjištěno 13 druhů netopýrů a u 10 druhů byly v létě nalezeny reprodukční kolonie samic. 218 Plošnou distribuci jednotlivých druhů znázorňují síťové mapy. Za nejrozšířenější druhy lze považovat netopýry rodu Plecotus (50,0 % kvadrátů pokrývajících studované území), Myotis myotis (46,7 %) a M. daubentonii (40,0 %). Výšková distribuce lokalit, kde byli nalezeni netopýři, je vyjádřena mediány a kvantily jejich nadmořských výšek a znázorněna pro jednotlivé druhy krabicovými diagramy podle období nálezů (zimní, letní). Lokality s letními koloniemi jsou hodnoceny samostatně. P. austriacus preferoval průkazně nižší polohy než ostatní hodnocené druhy, a to jak v zimě, tak v létě (medián 259, resp. 298 m n. m.). Naopak, výše položená zimoviště využívali nejvíce netopýři skupiny M. mystacinus a M. nattereri (medián 872 m, resp. 1050 m n. m.). Vysoké hodnoty jsou ale ovlivněny relativně nízkým počtem zimovišť vhodných pro hibernaci těchto těžko nalezitelných druhů. Většinou se jedná o vysoko položené pseudokrasové jeskyně. V mimohibernačním období byli ve vyšších polohách nalézáni M. brandtii a opět M. nattereri (medián 765, resp. 665 m n. m.). Výšková distribuce letních nálezů je u méně častých druhů do jisté míry ovlivněna odchytem před vchody do jeskyní, nalézajícími se ve velkých nadmořských výškách. Zvláště je to patrné u druhů, které se jinými metodami v létě zjišťují obtížně. Rozdíly v nadmořských výškách lokalit s letními koloniemi jsou vzhledem k nízkému počtu lokalit statisticky neprůkazné. Acknowledgement I am oblidged to Prof. Jiří Gaisler for all his comments on the text and its translation to English. I am also grateful to Dr. Tomáš Bartonička for his help with statistical evaluation. The study was supported by the Long-term Research Project of Ministry of Education, Youth and Sports of the Czech Republic No. MSM0021622416. References Anděra M. & Červený J., 1994: Atlas of distribution of the mammals of the Šumava Mts. Region (SW-Bohemia). Acta Sci. Natur. Brno, 28(2–3): 1–111. Anděra M., Hanák V. & Vohralík V., 1974: Savci Krkonoš [Die Säugetiere des Krkonoše-Gebirges]. Opera Corcontica, 11: 131–184 (in Czech, with a summary in German). Baroň I. & Řehák Z., 1997: K problematice zimování vrápence (Rhinolophus hipposideros) na Vsetínsku [Questions of wintering of Lesser Horseshoe Bat, Rhinolophus hipposideros in the Vsetín region]. Zpravodaj OVM Vsetín, 1997: 3–12 (in Czech). Baroň I. & Řehák Z., 2001: Netopýři Hranické propasti [Bats of the Hranická propast Chasm]. Speleo fórum 2001, 20: 44 (in Czech). Barratt E. M., Deaville R., Burlant T. M. & Bruford M. V., 1997: DNA answers the calls of pipistrelle bat species. Nature, 387: 138–139. Bartonička T., 2004: Visual identification of Myotis mystacinus and Myotis brandtii in a hibernaculum: preliminary results. Pp.: 7–12. In: Flousek J. & Bartonička T. (eds.): Bats of the Sudetes. Proceedings of the 2nd Czech-Polish-German Conference. Krkonoše National Park Administration, Vrchlabí, 116 pp (in Czech, with abstracts in English, Polish and German). Bartonička T., Řehák Z., Wolf P. & Bryja J., 2002: Drobní savci CHKO Litovelské Pomoraví. Část 1. Netopýři – Chiroptera [Small mammals of the Litovelské Pomoraví Protected Landscape Area (Czech Rep.). Part 1. Bats – Chiroptera]. Lynx, n. s., 33: 35–46 (in Czech, with an abstract in English). Benda P., Reiter A., Rejl J. & Bárta Z., 1997: Netopýři Železných hor [Bats of the Železné hory Mts (E-Bohemia)]. Vespertilio, 2: 39–50 (in Czech, with an abstract in English). Buřič Z. & Šefrová D., 2001: Zimoviště netopýrů v Jeseníkách a v Králickém Sněžníku a jeho okolí [Bat hibernacula in the Jeseníky Mts and Králický Sněžník Mt. and its surroundings]. Vespertilio, 5: 19–34 (in Czech). Demek J. & Střída M. (eds.), 1971: Geography of Czechoslovakia. Academia, Praha, 330 pp. 219 Gaisler J., 1956: Faunistische Übersicht der tschechoslowakischen Fledermäuse. Ochr. Přír., 11: 161–169 (in Czech, with summaries in German and Russian). Gaisler J., 1997: Nyctalus noctula na sídlištích (s výzvou ke spolupráci) [Nyctalus noctula at housing estates (with an appeal to cooperation)]. Bull. ČESON, 8: 8–10 (in Czech). Gaisler J., Bauerová Z., Vlašín M. & Chytil J., 1988: The bats of S-Moravian lowlands over thirty years: Rhinolophus and large Myotis. Folia Zool., 37: 1–16. Gaisler J., Chytil J. & Vlašín M., 1990: The bats of S-Moravian lowlands (Czechoslovakia) over thirty years. Acta Sci. Natur. Brno, 24(9): 1–50. Gaisler J., Řehák Z. & Bartonička T., 2002: Mammalia: Chiroptera. In: Řehák Z., Gaisler J. & Chytil J. (eds): Vertebrates of the Pálava Biosphere Reserve of UNESCO. Folia Fac. Sci. Nat. Univ. Masaryk. Brun., Biol., 106: 139–149. Hanák V., 1960: [Rozšíření a taxonomie středoevropských druhů netopýrů (Microchiroptera) se zvláštním zřetelem k území Československa [Distribution and taxonomy of C-European bat species (Microchi roptera) with respect to the territory of Czechoslovakia]. Unpublished PhD thesis, Charles University, Praha, 400 pp (in Czech). Hanák V. & Anděra M., 2005: Atlas rozšíření savců v České republice. Předběžná verze V. Letouni (Chiroptera) – část 1. Vrápencovití (Rhinolophidae), netopýrovití (Vespertilionidae – Barbastella barbastellus, Plecotus auritus, Plecotus austriacus) [Atlas of the Mammals of the Czech Republic. A Provisional Version, V. Bats (Chiroptera) – Part 1. Horseshoe bats (Rhinolophidae), vespertilionid bats (Vespertilionidae – Barbastella barbastellus, Plecotus auritus, Plecotus austriacus)]. Národní muzeum, Praha, 118 pp (in Czech, with a summary in English). Jahelková H., 2003: Přehled a srovnání echolokačních a sociálních signálů čtyř evropských druhů netopýrů rodu Pipistrellus (Chiroptera: Vespertilionidae) [Review and comparison of the echolocation and social calls in four European bat species of the genus Pipistrellus (Chiroptera: Vespertilionidae)]. Lynx, n. s., 34: 13–28 (in Czech, with an abstract in English). Jones G. & van Parijs S. M., 1993: Bimodal echolocation in pipistrelle bats: are cryptic species present? Proc. R. Soc. Lond. B, 251: 119–125. Kirchner K. & Řehák Z., 1995: K zimování netopýrů v podzemních prostorech v okolí Vsetína [Contribution to the wintering of bats in the undergound spaces in surroundings of Vsetín]. Speleo, 19: 33–35 (in Czech). Koutný P., Málková I. & Vlašín M., 1998: Mapování savců ve vybraných čtvercích jižní Moravy [Mapping of mammals in selected squares of South Moravia]. Sbor. Přírodověd. Klubu v Uherském Hradišti, 3: 125–138 (in Czech, with an abstract in English). Lehotská B. & Lehotský R., 2000: Skúsenosti z ochrany zimnej kolónie raniaka hrdzavého (Nyctalus noctula) v panelovém dome na bratislavskom sídlisku Dlhé Diely [Experience of protecting the winter colony of Nyctalus noctula in a prefab at the Bratislava housing estate Dlhé Diely (Slovakia)]. Vesper tilio, 4: 105–110 (in Slovak, with an abstract in English). Lemberk V., 2004: Netopýři (Chiroptera) východních Čech [Bats (Chiroptera) of eastern Bohemia (Czech Republic)]. Lynx, n. s., 35: 49–119 (in Czech, with an abstract in English). Lučan R., 2000: Netopýři Hostýnských vrchů [Bats of the Hostýnské vrchy Mts. (C-Moravia)]. Vespertilio, 4: 111–116 (in Czech, with an abstract in English). Lučan R. & Svačina T., 2001: Přehled zimovišť netopýrů v Hostýnských vrších a jejich předhůří [Review of bat hibernacula in the Hostýnské vrchy Mts and their piedmont]. Vespertilio, 5: 173–174 (in Czech). Reiter A., Hanák V., Benda P. & Barčiová L., 2003: Netopýři (Chiroptera) jihozápadní Moravy [Bats (Chiroptera) of South-Western Moravia (Czech Republic)]. Lynx, n. s., 34: 79–180 (in Czech, with an abstract in English). Remeš M., 1927: Ssavci Moravy a Slezska [Mammals of Moravia and Silesia]. Čas. Vlast. Spol. Mus. v Olomouci, 38: 32–52 (in Czech). Rumler Z., 1985: Výsledky chiropterologických průzkumů některých podzemních prostorů Beskyd a Oderských vrchů v letech 1976–1982 (Mammalia: Chiroptera) [The results of the chiropterologi- 220 cal investigations of some underground spaces in the Beskydy and Oderské vrchy Mts. in the years 1976–1982 (Mammalia: Chiroptera)]. Čas. Slez. Muz. (A) (Opava), 34: 75–89 (in Czech). Řehák Z., 1998: Faunistický přehled netopýrů moravskoslezské části Karpat (Česká republika) I [Faunistic review of bats in the Moravian and Silesian part of the Carpathians (Czech Republic) I]. Vespertilio, 3: 111–130 (in Czech, with an abstract in English). Řehák Z., 2001: Letouni (Chiroptera) [Bats (Chiroptera)]. Pp.: 251–254. In: Pavelka J. & Trezner J. (eds.): Příroda Valašska [Nature of Wallachia]. Czech Union for Nature Protection ZO 76/06 Orchidea, Vsetín, 488 pp (in Czech, with a summary in English). Řehák Z., 2006: Aktivita jediné jeskynní reprodukční kolonie netopýra velkého, Myotis myotis, v České republice [Activity of the Sole Reproductive Colony of Greater Long-eared Bat, Myotis myotis in the Czech Republic]. Unpublished repport of a project by CBCT, Praha, 4 pp (in Czech). Řehák Z. & Baroň I., 2002: Netopýři Hranické propasti [The bats of the Hranická propast Chasm]. Pp.: 24–25. In: Orálek M. (ed.): Hranická propast [Hranická propast Chasm]. Český svaz ochránců přírody, Valašské Meziříčí, 39 pp (in Czech). Řehák Z. & Bryja J., 1998: Drobní savci CHKO Poodří a blízkého okolí: II. Chiroptera [Small mammals in the Protected Landscape Area of Poodří and its vicinity. II. Chiroptera]. Čas. Slez. Muz. (A) (Opava), 47: 133–142 (in Czech, with an abstract in English). Řehák Z. & Gaisler J., 1999: Long-term changes in the number of bats in the largest man-made hibernaculum of the Czech Republic. Acta Chiropterol., 1: 113–123. Řehák Z. & Gaisler J., 2001: Netopýři zimující ve štolách pod Jelení cestou u Malé Morávky v Jeseníkách [Bats wintering in the abandoned mines under the Jelení road near Malá Morávka in the Jeseníky Mts (Czech Republic)]. Vespertilio, 5: 265–270 (in Czech, with an abstract in English). Řehák Z., Eleder P. & Gajdošík M., 2000: Příspěvek k poznání fauny netopýrů v CHKO Žďárské vrchy [Contribution to the knowledge of bat fauna in the Žďárské vrchy Protected Landscape Area]. Žďárské vrchy v čase a prostoru – sborník konferenčních příspěvků [Žďárské vrchy Highlands in Time and Space – Proceedings of the Conference Contributions]. Sphagnum – Ecological Society & Administration of the Žďárské vrchy Protected Landscape Area, Žďár nad Sázavou, 179 pp (in Czech). Řehák Z., Chytil J., Bartonička T. & Gaisler J., 2003: Výskyt drobných savců na území Biosférické rezervace Dolní Morava (rozšířená Biosférická rezervace Pálava). Část II. Netopýři – Microchiroptera [Distribution of small mammals in the Biosphere Reserve Lower Morava (extended BR Pálava). Part II. Bats – Microchiroptera]. Lynx, n. s., 34: 181–203 (in Czech, with an abstract in English). Řehák Z., Bartonička T. & Bielik A., 2004: Distributional status of Pipistrellus pipistrellus (Schreber, 1774) and P. pygmaeus in the Czech Republic: results of mapping. Pp.: 102–103. In: Bogdanowicz W., Lina P. H. C., Pilot M. & Rutkowski R. (eds.): Programme and Abstracts for the 13th Internati onal Bat Research Conference, Poland, Mikołajki, 23–27 August 2004. Polish Academy of Sciences, Warszawa, 120 pp. Řehák Z., Bartonička T. & Bielik A., 2005: Distribution of Pipistrellus pipistrellus and P. pygmaeus in the Czech Republic. P.: 26. In: Abstracts of 19th Polish Chiropterological Conference, Pokrzywna, 4–6 November 2005. Opole University, Opole. Souček J., 1969: Nejvýše položené zimoviště netopýrů v Hrubém Jeseníku [The highest hibernating quarter of bats in the Hrubý Jeseník Mts]. Vertebratol. Zpr., 1969(3): 117–120 (in Czech, with a summary in English). Šafář J. & Rumler Z., 2001: Netopýři zimující na vybraných zimovištích severní Moravy [Bats hibernating in selected hibernacula of northern Moravia]. Vespertilio, 5: 271–278 (in Czech). Vlašín M., Eleder P. & Nečasová I., 1993: Rozšíření ochranářsky důležitých druhů savců v jihomoravském regionu (II. část) [Distribution of mammal species important for nature conservation in S-Moravian region. Part II]. Vlast. Sbor. Vysočiny (Jihlava), 11: 273–295 (in Czech). Vlašín M., Eleder P. & Málková I., 1995: Rozšíření ochranářsky důležitých druhů savců v jihomoravském regionu (III. část) [Distribution of mammal species important for nature conservation in S-Moravian region. Part III]. Vlast. Sbor. Vysočiny (Jihlava), 12: 205–241 (in Czech). 221 Wagner J., 2001: Zimoviště netopýrů v Nízkém a Hrubém Jeseníku, Oderských vrších a Moravskoslezských Beskydách [Bat hibernacula in the Jeseníky Mts, Oderské vrchy Hills and Moravskoslezské Beskydy Mts]. Vespertilio, 5: 287–302 (in Czech). Wagner J., 2004: Zimoviště netopýrů v Oderských vrších [Bat hibernacula in the Oderské vrchy Hills (N-Moravia, Czech Republic)]. Pp.: 91–106. In: Flousek J. & Bartonička T. (eds.): Bats of the Sudetes. Proceedings of the 2nd Czech-Polish-German Conference. Krkonoše National Park Administration, Vrchlabí, 116 pp (in Czech, with abstracts in English, Polish and German). 222 Appendix The list of localities with records of bats / seznam lokalit s nálezy netopýrů Explanations / vysvětlivky: SN – quadrate number of the Czech mapping grid / číslo kvadrátu české mapovací sítě; SPN – number of bat species / počet druhů netopýrů; S – season / období; G – growing season – “summer” records of bats / mimohibernační období – “letní” nálezy netopýrů; W – winter season – records of torpid bats at hibernacula and all winter records of V. murinus / zimní období – nálezy letargujících netopýrů na zimovištích a všechny zimní nálezy V. murinus. SN site / lokalita habitat / stanoviště SPN Slezské Beskydy Mts 6378 Nýdek cottage 1 in the open air 3 Jablunkovská vrchovina Highland 6478 Girová, Na Girové cave cave 3 Moravskoslezské Beskydy Mts 6377 Řeka in the open air 2 Smilovice above a stream 1 Komorní Lhotka above a stream in a village centre 1 near a stream at the end of a village 3 6475 Frenštát p. R. church loft 1 Malá Stolová, Leopoldka in the open air 1 6476 Čeladná cottage 1 Lukšinec, Ondrášovy díry cave cave 6 cave entrance 8 Lukšinec mountain forest 1 Lysá hora, Malenovice canyon mountain forest 7 Lysá hora, Pod Ivančenou mountain forest and meadow 6 Ostravice above a stream in a village 1 Ostravice, Mazák above a stream 1 6477 Morávka cottage 1 6575 Čertův mlýn, Čertova díra cave cave 7 Dolní Bečva, Kamenné in the open air 1 cottage celllar 1 Kněhyně, Kněhyňská cave cave 7 cave entrance 10 Kněhyně, Kyklop cave cave 2 Pustevny, Cyrilka cave cave 8 cave and cave entrance 9 Vasko cave 2 6576 Staré Hamry church loft 1 house roof 1 Podbeskydská pahorkatina Upland 6277 Ropice above a stream 4 Třanovice above a stream 1 6375 Hukvaldy cottige roof 1 a game preserve 1 6376 Baška dam 1 Dobrá unknown 1 Frýdek-Místek free on a pavement 1 S G G W G G G G G G G W G G G G G G G W G G W G W W G W G G G G G G G G G 223 6377 Hnojník Střítež 6471 Helfštýn Lipník nad Bečvou 6472 Černotín Hranice Hranice, Hůrka Lipník nad Bečvou Teplice nad Bečvou 6473 Hustopeče nad Bečvou Hustopeče nad Bečvou, Na Valše Lešná u Valašského Meziříčí Perná-Bučí Poruba Starý Jičín 6474 Hodslavice Nový Jičín Štramberk Štramberk, Bílá hora Štramberk, Kotouč Štramberk, Na Horečkách Štramberk, Šipka Štramberk, Trúba Veřovice-Padolí 6475 Kunčice p. O. Kunčice p. O., Skalka 6572 Kelč 6573 Branky na Moravě Poličná Poličná-Junákov Poličná-Juřinka Valašské Meziříčí 224 above a stream above a stream, under a bridge in the open air above a river gallery in a quarry above a river above a river above a street-lamp near a autocamp chasm cellar above a river boiler room in a spa building cave cave castle loft, pellets of T. alba above a pond castle cellar and corridor castle loft church loft house roof anf loft house castle church loft, pellets of T. alba cellar free in a castle park and a city above a pond above a swimming pool in the open air in a park garden in an abandoned quarry in the open air in the Kamenárka quarry gallery in a quarry galleries cave and cave entrance near a cottage above a stream near a cottage above a pond house cottage, garden church and castle lofts, pellets of T. alba castle loft house loft farm loft stream bank - pellets of S. aluco pellets of S. aluco buildings, in the open air, pellets of T. alba buildings and in the open air castle cellars castle loft castle park house loft 1 3 1 3 7 5 4 2 14 1 2 1 2 1 4 2 2 5 3 5 1 2 3 1 1 2 1 3 3 1 2 2 4 1 1 1 1 1 2 3 1 1 1 1 1 4 1 1 2 2 2 G G G G W G G G G W G G W G G G W G G G G G G W G G G G G G W W G G G G G G G G G G G G G G W W G G G 6671 Holešov house loft church loft castle cellar house cellars Němčice house cellar Rožnovská brázda Furrow 6574 Hrachovec above a river Rožnov pod Radhoštěm bulding buildings owl box, pellets of S. aluco park and above a river Rožnov p. R., Dolní Paseky beech wood Rožnov p. R., Hradisko pellets of S. aluco Střítež in the open air Velká Lhota in the open air Veselá in the open air Vidče church loft in the open air Zašová in the open air church loft, owl pellets 6574 Zubří church loft, owl pellets above a stream village and in the open air Zubří-Hamry tree hole, owl pellets pond banks and in the open air 6575 Dolní Bečva river bank in the open air Horní Bečva church loft in the open air above a dam Rožnov p. R., Rysová owl box, pellets of S. aluco Vigantice in the open air Javorníky Mts 6674 Hovězí in the open air Huslenky church tower above a river 6675 Velké Karlovice village 6676 Malé Karlovice-Tísňavy school loft, garden Velké Karlovice-Podťaté house 6774 Francova Lhota cottage loft Lužná in the open air Pulčín-Hradisko rocks Pulčín-Hradisko, Pod Kazatelnou cave cave entrance 6874 Francova Lhota in the open air Zděchov church loft Hostýnsko-vsetínská hornatina Mts 6572 Bystřice p. Hostýnem castle cellars castle park church loft 1 1 3 2 1 G G G G W 1 1 2 1 6 1 1 2 1 3 1 3 6 1 2 1 2 4 6 1 2 1 3 2 1 1 G W G G G G G G G G G G G G G G G G G G G G G G G G 2 1 1 3 3 1 1 2 1 4 10 4 1 G G G G G G G G G W G G 5 2 1 W G G 225 6573 Bystřička school loft 1 Podhradní lhota cadaver on the road 1 Polomsko cottage 1 Rajnochovice buildings 1 church loft 1 Rajnochovice, Juhyně valley above a stream 2 6574 Bystřička-Zadrhlov gallery 4 Malá Lhota, Páleniska house loft 1 6671 Dobrotice-Lysina cottage 1 6672 Chvalčov road 1 village 3 Chvalčov, Bystřička valley above a stream 8 clearing 1 Chvalčov, Smrdutá cave 3 cave entrance 10 Rajnochovice, Košový building 1 Rajnochovice, Uhliska above a stream 1 Rusava Protestant church, loft 1 Catholic church - loft 1 cadaver on the road 1 Rusava, Ráztoka banks of a water reservoir 1 Rusava, Čecheřinka cave cave 3 cave entrane 8 6673 Jablůnka river bank 1 Kateřinice school loft 1 Křížový, Zbojnická cave cave 3 Pržno church loft 1 Růžďka loft 1 Vsetín town, above a river 4 cellar 1 loft 1 6674 Halenkov railway station, in the open air 1 Janová in the open air 5 Vsetín-Hluboké in the open air 1 6675 Karolinka in the open air 1 Nový Hrozenkov church loft and tower 2 Velké Karlovice, Jezerné above a pond 7 6676 Velké Karlovice, Miloňov loft of a hotel 1 6772 Kašava church loft 1 Lukov house cellars 1 Lukov, castle castle cellars 3 6 Lukov - Hradisko c. cave 3 cave entrance 2 6773 Liptál Protestant church, loft 2 Catholic church, loft 1 school loft 1 Vizovická vrchovina Highland 6771 Zlín in the open air 1 6772 Lešná u Zlína cellar 1 Slušovice church loft 1 226 G G G G G G W G G G G G W G G G G G G G W G G G W G G G W G G G G G G G G G W W G W G G G G W G 6773 Bratřejov church loft 3 Jasenná church loft 1 Liptál, Malá gallery mine 2 Liptál, Sintrová gallery mine 5 Pozděchov Protestant church, loft 3 Catholic church, loft and tower 4 Vizovice castle cellar 2 castle loft 2 church loft 1 hospital loft 1 6774 Kopce u Lidečka, Naděje cave cave, cave entrance 8 Kopce u Lidečka, Ďáblova díra c. cave 2 Kopce u Lidečka, Kolonie cave cave 5 Kopce u Lidečka 3 cave entrances 9 Leskovec in the open air 1 Lidečko church loft 2 in the open air 3 6872 Březnice church loft 1 Doubravy in the open air 1 Horní Lhota church loft 1 Kaňovice chapel turret 1 Luhačovice castle cellar 2 castle loft 3 Pozlovice church loft 1 Velký Ořechov church loft 1 6873 Újezd church loft 1 6874 Horní Lideč in the open air 3 Lačnov church loft 1 above a pond 1 6971 Uherský Brod house cellar 1 church loft 1 7070 Blatnice church loft 1 7071 Blatnička church loft 1 above a pond and a stream 1 7072 Bánov school loft 1 Nivnice church tower 1 Bílé Karpaty Mts 6874 Nedašov church loft 1 Poteč barn 1 Valašské Klobouky house loft 1 church loft 1 mill loft 1 town hall loft 1 in the open air 2 6972 Bojkovice castle cellar 2 castle loft 1 Nezdenice church loft 1 Rudice house loft 1 Záhorovice house roof 1 6973 Slavičín castle loft 2 church loft 1 G W W G G W G G G W W W G G G G G G G G W G G G G G G G W G G G G G G G G G G G G G W G G G G G G 227 Štítná nad Vláří Štítná nad Vláří, Popov 6974 Brumov 7070 Lipov 7071 Boršice u Blatnice Horní Němčí 7072 Březová Komňa Strání 7073 Starý Hrozenkov, Rasová quarry 7170 Hrubá Vrbka Kněždub Kútky Kuželov Měsíční údolí Radějov Tvarožná Lhota, Horní mlýn 228 church loft church loft old mill house loft cottage wine cellar church loft and tower below house roof church tower church tower above a pond and around street lamps above a pond church loft and tower church loft cadaver on the road chaple loft above a pond 13 wooden hides church, external wall, loft and tower wood hide church tower above a swimming pool 1 1 1 2 1 1 1 1 1 1 4 1 2 1 1 2 2 2 2 1 1 4 G G G G G W G G G G G G G G G G G G G G G G Lynx (Praha), n. s., 37: 229–240 (2006). ISSN 0024–7774 Dynamics of the Pleistocene bat fauna from the Matuzka Paleolithic site (Northern Caucasus, Russia) (Chiroptera) Dynamika netopýří fauny paleolitického stanoviště Matuzka (severní Kavkaz, Rusko) (Chiroptera) Valentina V. Rossina1, Gennadiy F. Baryshnikov2 & Bronislaw W. Woloszyn3 Paleontological Institute, Russian Academy of Sciences, Moskva, Russia; ros@paleo.ru Zoological Institute, Russian Academy of Sciences, 199 034 Sankt Petersburg, Russia 3 Institute of Systematics and Evolution of Animals PAS, Sławkowska 17, PL–31-016 Kraków, Poland 1 2 received on 31 May 2006 Abstract. The upper Pleistocene sedimentary series in Matuzka cave (northern Caucasus) covering a period from MIS6 to MIS2 provided remains of 18 species of bats. The bat record is particularly rich in the layers corresponding to Eemian and early Vistulian. It is characterized with appearance of thermophilous elements Rhinolophus ferrumeqiunum and Miniopterus schreibersii and a broad spectrum of taxa including dendrophilous and demanding elements such as Plecotus auritus, Myotis brandtii, M. emarginatus, M. nattereri and M. blythii. The lithophilous forms Eptesicus serotinus, Vespertilio murinus and Nyctalus noctula appear continuously in all layers and represent a dominant component of the assemblage. In Eemian layers they are supplemented also with Hypsugo savii and Pipistrellus cf. kuhlii which absent from the upper layers while Pipistrellus pipistrellus appear as late as in the Early Holocene. Introduction During the last decades, many Paleolithic sites have been discovered on the northwestern Caucasus. The rich bone material from these sites includes numerous records of bats. The Caucasus is a unique region particularly in respect to its climatic specificities and biogeographical role. The Caucasian mammal fauna is characterized by considerable species richness and high degree of endemism. This holds true also for bats: the northern Caucasus is presently inhabited by one of the richest bat fauna of Russia. Unfortunately, until now almost no information was available on the history if the Caucasian bat fauna except for rather episodical and isolated records (comp. Vereščagin 1959). To gain a better understanding of the development of bat faunas, the study of new paleontological data on this group is required. The present paper describes the material of Late Pleistocene bats collected by G. F. Bary shnikov during archeological excavations of the Matuzka Paleolithic site. This study provides insight into the major stages of the formation of Caucasian bat communities during the essential part of the last glacial cycle, Eeminan and Vistulian. 229 Material and Methods The cave site Matuzka (42° 26’ N, 39° 45’ E) is located at 720 m above sea level, at the northern margin of the Lagonakskoe Plateau on the right bank of the Matuzka River, a right tributary of the Pshekha River, 27 km south-southeast of Apsheronsk (Fig. 1). The cave cavity is in Upper Jurassic limestones, has a dome-shaped roof and karstic niches in the walls and roof. The cave is up to 35 m wide and about 40 m deep, the entrance faces southwest and is 20 m high. According to geomorphologic analysis performed by Nesmeyanov (1999), the primary cave cavity, which is most likely of karstic-erosive origin, was formed about 150–130 thousand years ago. The analysis of limestone pack deformation has shown that the cave initially had two, or possibly, three layers. The three-floor primary cavity of Matuzka cave, with the total height up to 30–35 m, suggests a long time of the cavity formation, with the participation of lateral river erosion. This process apparently resulted in the large final size of Matuzka cave (Nesmeyanov 1999). The cave is presently a large grotto. The site was excavated from 1985 to 1988. The 6-m-deep sequence of Pleistocene-Holocene deposits was exposed in the site. The section of excavation comprises 8 major lithologic beds (Table 1; Baryshni kov & Golovanova 1989). The sequence is subdivided by lithologic features into four units. Three lower units (beds 7–3a) contain a Pleistocene mammal fauna and Mousterian artifacts (Golovanova et al. 1995). The first lower unit is 0.6 m thick (beds 8a–7b) includes the deluvial deposits composed of slightly inclined laminated loam with insertion of gruss. Apparently, it was formed under conditions of low-activity washout. The second and third units (beds 7a–3a) are composed of rubbly-clumpy material with varying content of loamy-gruss-rubbly filler. The second unit (beds 7a–5) is 3 m thick, of land-rockslide genesis, composed of yellow loam. The third unit (beds 4d–3a) is formed by gray loam. Bed 4b is 0.1 m thick, contains charcoals of fire spots (Nesmeyanov 1999), with the absolute date 34200±1410 BP (Golovanova 1996). There are no absolute dates for the others layers. However, based on rodent fauna (Nadachowski & Baryshnikov 1991) beds 7–6 are dated to terminal Middle Pleistocene, beds 5–3 are assigned to the time of the last glaciation, and beds 2–1 are dated Upper Pleistocene-Holocene boundary (Zaitsev & Osipova 2004). The fourth unit (beds 2–1) is 0.6 m thick, composed of lumpy-rubbly loam and contains of ceramics fragments and burnt bones of domestic animals (Nesmeyanov 1999). Fig. 1. The region of investigation with a location of the Matuzka cave (asterisk). Obr. 1. Oblast výzkumu s lokalisací jeskyně Matuzka (hvězdička). 230 In addition to the main sites, excavations were performed in further trench assigned as trench 3. Based on the mammal fauna and bone fossilization pattern, the layers of trench 3 were dated to Pleistocene-Holocene boundary, i.e. they correspond roughly to beds 3 and 1–2. A total of 217 skull fragments and isolated teeth of bats were examined (Table 1). The material was examined by binocular microscope, identified and compared with the samples of Recent species. The parataxonomic categories were applied in few cases in which considerable fragmentation and poor preservation of specimens did not allow exact identification: “small-sized Myotis” means of the same size as M. brandtii, “medium-sized Myotis” means of the M. emarginatus size. Results Structure of the oryctecoenoses The complete list of species found in individual samples from the Late Quaternary beds of Matuzka cave is in Table 1. In total, 17 species were recorded: Eptesicus serotinus (Schreber, 1774), incl. cf. E. serotinus; Vespertilio murinus Linnaeus, 1758 (cf. V. murinus); Nyctalus noctula (Schreber, 1774) (incl. cf. N. noctula); Nyctalus leisleri (Kuhl, 1817); Myotis blythii (Tomes, 1857) (M. cf. blythii); Barbastella barbastellus (Schreber, 1774); Plecotus auritus (Linnaeus, 1758); Rhinolophus ferrumequinum (Schreber, 1774); Miniopterus schreibersii (Kuhl, 1817); Pipistrellus pipistrellus (Schreber, 1774); P. nathusii (Keyserling et Blasius, 1839); P. cf. kuhlii (Kuhl, 1817); Hypsugo savii (Bonaparte, 1837); Myotis brandtii (Eversmann, 1845) (Myotis cf. brandtii); M. nattereri (Kuhl, 1817) (M. cf. nattereri); M. bechsteinii (Kuhl, 1817) (M. cf. bechsteinii); M. emarginatus (Geoffroy, 1806) (M. cf. emarginatus). The particular remains were directly compared to the samples of Recent specimens and apparently fall in variation of the respective Recent species except for several Pleistocene items which appear to be somewhat larger than reported for the Recent Caucasian material. The most abundant species of the oryctocenosis were E. serotinus and V. murinus (30% and 26% of all remains, respectively). N. noctula, Plecotus auritus (both about 7%), B. barbastellus and R. ferrumequinum (about 5%) were recedent elements while Myotis blythii (with somewhat more than 2%) and all remaining species (with less than 1.9%) represent the subrecedent elements of the sample. As concerns the structural characteristics of the oryctcenoses the whole set clearly splits into two markedly different units: (I) The faunal association of the bed 7 (accumulated during the Mikulino Interglacial = Eemian in the Western Europe) that is characterized by considerable species richness and includes thermophilous elements such as R. ferrumeqiunum, M. schrei bersii, P. kuhlii, H. savii. Its essential characteristics are partly retained in the overlaying bed 6, where Miniopterus or H. savii are absent, of course (Fig. 2). The distribution of these species in the section marks the boundary between the Mikulino Interglacial and the onset of the Valdai Glacial Period (= Würm). (II) Beds 5–3a (accumulated during the Valdai glaciation) yielded the bat fauna that is slightly poorer both in species richness and abundance. It also differs from (I) by presence of M. nattereri, M. emarginatus and M. bechsteinii and well-pronounced instability in the proportions of particular taxa (Fig. 2). It is dominated with the mesophilous elements, among other Plecotus auritus and B. barbastellus, and Nyctalus noctula which reaches peak of its abundance in the middle Valdai horizons. As demonstrated in Fig. 3, there is a clear correlation between the contribution of bats to the total mammalian community of a layer and contributions of the rodents inhabiting either forest or open-ground habitats: a positive correlation exists with the proportion of rodents inhabiting mountain forest formations and negative correlation with the proportion of rodents living in 231 Table 1. Chronological position of Matuzka cave strata and numbers of Chiroptera records Tab. 1. Chronologická posice vrstev jeskyně Matuzka a počty v nich nalezených netopýrů time MA O-isotope mammal age stage Eastern Europe Caucasus Matuzka cave layer Chiroptera (217) 0.025 2 Sungilian Akhtyr 1–2 + shaft 3 P. pipistrellus (3), P. nathusii (1) E. serotinus (13) V. murinus (4), P. auritus (1) M. nattereri (1) M. bechsteinii (2) M. emarginatus (1) 3 Chasovali 3 E. serotinus (3), N. leisleri (1) B. barbastellus (2) M. brandtii (1) medium-sized Myotis (2) 0.073 4 Shkurlatian 4 E. serotinus (19) V. murinus (6), P. auritus (7) B. barbastellus (8) medium-sized Myotis (1) small-sized Myotis (1) M. nattereri (1) M. bechsteinii (2) M. emarginatus (2) N. noctula (12) 0.116 5a–5d 5–5b E. serotinus (2) V. murinus (3), P. auritus (1) M. nattereri (1) M. emarginatus (1) M. blythii (1), N. noctula (1) 0.128 5e Binagady 6 P. cf. kuhlii (1), N. leisleri (1) E. serotinus (13) V. murinus (21), P. auritus (1) M. nattereri (3), M. blythii (1) M. emarginatus (1) R. ferrumequinum (1) 0.195 6 Khazarian Kvaisi 7 P. cf. kuhlii (1), H. savii (1) E. serotinus (20) V. murinus (23), P. auritus (4) B. barbastellus (2) M. brandtii (2), M. blythii (3) R. ferrumequinum (10) N. noctula (2) M. schreibersii (2) open landscapes, such as mountain-steppe and shrub-and-grassland habitats. This suggests considerable retreat of bat populations in periods of the most pronounced glacial conditions (indicated by percentage of open-ground elements). 232 Fig. 2. Percentual distribution of a total number of individuals of particular bats species in sequence of layers of the Matuzka cave. Obr. 2. Procentuální rozložení celkového počtu jedinců jednotlivých druhů netopýrů ve sledu vrstev jeskyně Matuzka. Fig. 3. Percentages of the bats in the total mammalian samples of particular layers (bold line) compared to those of forest and open-ground rodents. Obr. 3. Procentuální zastoupení netopýrů v celkovém vzorku savců (silná čára) ve srovnání se zastoupením lesních druhů hlodavců a hlodavců otevřené krajiny. 233 Taphonomy Holocene bones from bed 1 have natural white colour. Most of the Pleistocene bone specimens are light brown, some are colored with manganese. Some bones (mostly teeth) were damaged by the digestive juice (enzymes): bed 4b (N. noctula), trench 3 (bed 3, V. murinus), bed 6 (3 specimens of E. serotinus (2) and P. auritus (1)), bed 7 (10 specimens; 4 V. murinus, 5 E. serotinus and 1 M. cf. brandtii). This suggests that a significant part of bat sample may represent a taphocenosis. The predominating representation of the lithophilous elements in the samples (Eptesicus serotinus, Vespertilio murinus, Nyctalus spp. Pipistrellus spp., Hypsugo savii) suggests at the same time that a considerable part of the material may originated from autochtonous sources, supposedly winter colonies of the respective species roosting in the ceiling fissures in the cave entrance and/or in the rocky walls surrounding the cave. Discussion Out of 23 bat species presently inhabiting the northern Caucasus, 17 have been recorded in the fosil assemblages under study. The species: Rhinolophus hipposideros, R. euryale, Myotis daubentonii, M. aurascens, M. mystacinus and Nyctalus lasiopterus are absent from the Matuzka orictocenosis. It is very important to recognize the reasons for their absence for analysis of the species diversity in the Pleistocene bat community. At present, Rhinolophus hipposideros is a common bat species in Caucasia which range is restricted to the forest zone (Gazaryan 2002). Though it frequently occurs in cave roosts it does not form large colonies but prefers smaller caverns for roosting which would indicate its possible incidence in Matuzka cave. In respect to the positive prediction on its appearance in fossil record, its absence may suggest that the pattern of abundance and geographical range of this species differed from those in the present time. In the case of R. euryale, a strict cave-dweller, extremely rare in the area under study (only one record by Bobrinskoj et al. 1965), its incidence in the site under study seems to be quite improbable, similarly as in the case of the migratory bat Nyctalus lasiopterus, a tree-dweller closely associated with forest vegetation. Also the remaining species of the Recent list which absents in the site are not too probable to appear there. The group of small-sized Myotis (M. brandtii, M. daubentonii, M. aurascens and M. mysta cinus) is scarce in the oryctocenosis, it is represented by ten specimens only. It is noteworthy that all the listed species are considered abundant in the Recent of Caucasia (Gazaryan 2002, 2003). The total abundance of Myotis daubentonii, extent of its range and degree of synanthropy substantially increased during past decades (Gazaryan 2003). The natural habitats of this species are confined to the foothills and river valleys. This species seems to avoid the altitudes above 1000 m above sea level (Gazaryan 2003). The sibling species M. aurascens and M. mystaci nus were only recently distinguished as the separate species based on detailed morphological analysis (Benda & Tsytsulina 2000). At present, M. aurascens inhabits lowlands and plains of Western Caucasus; it has not been recorded higher than 400 m above sea level (Gazaryan 2002). M. mystacinus and M. brandtii are very similar ecologically, both inhabiting mountain forests near water bodies. They coexist in the region under study (Gazaryan 2002). In case of all these species, their habitat preferences and abundances do not suggest particularly high probability of their incidence in Matuzka cave – therefore, their absence in the fossil sample need not be of a serious indication value. 234 Fig. 4. Percentual contributions of particular bat species to a community structure of individual layers of the Matuzka cave. Obr. 4. Procentuální zastoupení jednotlivých druhů netopýrů ve sledu vrstev jeskyně Matuzka. Single records of P. pipistrellus and P. nathusii in the early Holocene sediments of Matuzka cave are of the particular significance. They suggest the spread of these species in that period (especially in comparison to their absence in the earlier strata) which corresponds well to the scenario presented for their spread in central and Western Europe by Horáček & Jahelková (2005). There are more factors which supposedly might considerably influenced structure of the record and actual dynamics of bat communities recorded in Matuzka cave. Here we will discuss briefly three of them: (1) activity of the Paleolithic man; (2) aspects of bone accumulation; and (3) the climatic and environmental dynamics of the period under study. (1) The reconstruction of bat fauna history in the northwestern Altai region from the Pleistocene to the present time demonstrated that the historical development of cave-dwelling bat community was considerebly affected by cave-dwelling effort of the Paleolithic humans (Agadjanian & Rossina 2001, Rossina 2004). Occupation of caves by the humans apparently reduced the population of the majority of bat species because of the smoke produced by human fires. Thus, humans were a limitative factor for the cave-dwelling bat community since the Pleistocene (Rossina 2005). The Mousterian man did not produce tools in Matuzka cave, but brought there artifacts ready for use (Nesmeyanov 1999). However, Paleolithic humans regularly visited Matuzka cave for many ten thousand years, which is evident from the regular distribution of tools in beds 7–3 (Golovanova et al. 1995). The analysis of large mammal fauna leads to the same conclusion; the Mousterian man used Matuzka cave as a short-term shelter for humans during hunting (Baryshnikov & Golovanova 1989). Thus, in the case under consideration, human activity exerted a little effect on the bat community. (2) It is commonly accepted that the main source of fossil bats in cave deposits is dying bats from seasonal cave-dwelling colonies (Ovodov 1974, Tiunov 1997, Filippov & Tiunov 235 1999). But it is doubtful in the case under consideration. During the past 120 thousand years, Matuzka cave was a large grotto (Potapova 1992, Nesmeyanov 1999). Thus it seems largerly improbable that it may serve a roost for large winter colonies of cave-dwelling bat species, such as Myotis emarginatus or Miniopterus schreibersii, though occasional appearance of these bats during vegetation period (which used the cave e.g. as a night roost) or in transient period cannot be excluded, of course. On contrary, the regular appearance of the species inhabiting rocky fissures is here quite a probable, and their considerable representation in the sample fits to that possibility quite a well. The considerable fragmentation of all specimens and signs of treatment by the digestive juice indicate that pellets of birds of prey may were possibly a significant source of fossil material from Matuzka cave. Such a possibility would be indirectly supported also by the conclusion resulting of the study of fossil birds from Matuzka cave which demonstrated that a degree and character of damage of birds bones as well as the species composition of the sample indicate that fossils come from owl pellets (Potapova 1992). In these connection, the specificities of the two most abundant bat species, E. serotinus and V. murinus are worth mentioning. At present, they are synanthropic species which roost preferably in human buildings while the records from their natural roosts are generally rare. Correspondingly, only few winter records of E. serotinus and V. murinus are known from the western Caucasus (Kuzjakin 1950, Gazaryan 2002). V. murinus similarly like Nyctalus noctula and N. leisleri are seasonal migrants. Particularly worth of attention is that the large seasonal colonies of these bats may become a hunting objects of owls and the frequency of these species in owl diet may thus locally grow quite a high (Schmidt & Topal 1971, Ruprecht 1990, 2005, Obuch 1989, original data). It should be mentioned that also in several teeth of P. auritus and M. cf. brandtii the enamel shows clear signs of corrosion by the digestive juice. Note that Plecotus along with E. serotinus and N. noctula is the most favorite object of owls hunting (Kowalski 1995). Unfortunately, the material is too small to allow a detailed statistical analysis of possible taphonomic effects. In any case, it is clear that the fauna is to be considered a true oryctocenosis contributed both by taphocenoses and thanatocenoses. Despite the, of course, the stratigraphic setting of the site and the particular records prove that the records under study come from a well defined stratigraphical context and bear the respective stratigraphical and paleoecological information. The comparison of the absolute amount of bat records and percentages of habitat specialists in rodents (Fig. 3) is particularly significant in these connections. An increase of bat percentage in layers with increased frequency of woodland elements conforms well to a genral expectations on the paleoclimatic significance of chiropteran record. The general dynamics of the bat fauna in study thus apparently reflects changes in landscape and climatic conditions in the source area. In this sense the analysis of the structure of fossil bat assemblages not only provides additional information on environmental changes, but, in some cases, it promises to regard some aspects not accessible from other evidence (thermal balance of rocky massif as a key factor for hubernation of fissure-dwelling bats etc.). In addition, the analysis of faunal structure of Pleistocene bats provides important stratigraphic results. (3) The greatest proportion and taxonomic diversity of Chiroptera (about 30% of all specimens) is recorded in beds 7–5b. The absolute maximum of bats is in bed 7.1 (Figs. 2, 4). The oryctocenosis contains forest species of Pipistrellus and Nyctalus and the forest-steppe Myotis blythii (Gazaryan 2002). Apparently, during the accumulation of beds 7–6, a vast area was occupied by forests. The small mammal fauna includes forest species of the genera Pitymys and Dryomys. However, the proportions of mountain-steppe and shrub-and-grass rodent species are 236 also high (Baryshnikov & Golovanova 1989, Baryshnikov et al. 1995). The climate was warm; this is evident from the highest diversity of the bat assemblage from these beds (14 bat species out of 17 are recorded) and the presence of migratory species of Pipistrellus and thermophilous species, such as Rhinolophus ferrumequinum and Miniopterus schreibersii (more then 40% of all bat specimens; Fig. 4). At the boundary between beds 6 and 5b, the bat populations sharply decreased in number, with the absolute minimum in bed 5a (Figs. 2, 3). Apparently, at that time, the area of open landscapes increased and a more mosaic landscape structure was formed. This is supported by an increase in the number and proportion of mountain-steppe rodents (Fig. 3), such as Spermophillus cf. musicus, Spalax microphtalmus and Cricetulus migratorius guamensis. The climate became somewhat cooler (Baryshnikov & Golovanova 1989). During the accumulation of beds 5–4d–4c, the abundance and proportion of bats gradually increased (Figs. 2, 3). The proportion of forest rodent species increased, while rodents of open landscapes decreased in number (Fig. 3). This suggests an increase in the area of forest landscapes and, as follows from palynological data, the mountain pine forests spread, although they were more xerophilous than at the present time (Baryshnikov et al. 1995). Bed 4b shows a decrease in the proportion and abundance of bat remains, while the proportions of forest and steppe rodent species are approximately equal (Fig. 3). According to palynological data, the forest-steppe or steppe with mixed broad-leaved and pine forests were widespread at that time. However, grasslands dominated. The climate was relatively arid and temperate (Baryshnikov & Golovanova 1989, Baryshnikov et al. 1995). Against the background of a general increase in the proportion of rodents in bed 4a, the diversity and abundance of bats also increased (more then 16% of all specimens) (Figs. 2, 3). Apparently, the landscapes showed a mosaic pattern of distribution, so that mountain forests alternated with steppes and grasslands. The pollen spectra suggest the presence of mixed pine-birch forests with alder-tree and grass meadows (Baryshnikov et al. 1995). Bed 3c yielded only Barbastella barbastellus and medium-sized Myotis (Fig. 2). In bed 3b, the proportion of bat remains is equal to that of forest rodents (Fig. 3). At that time, the proportion of Chionomys increased and steppe rodent taxa predominated (Baryshnikov & Golovanova 1989). Palynological data suggest that the subalpine grasslands and mesophilic motley grasses dominated (Baryshnikov et al. 1995). The finds of bats and forest rodents are sporadic in bed 3a but steppe rodents predominate here as well (more than 90%; Fig. 3). Apparently, open landscapes dominated at that time. Floral samples contain underdeveloped pollen of trees and traces of turf-uncovered slopes, and Woodsia alpina. This suggests that at that time the cave was in the Alpine belt, close to the upper boundary of forests (Baryshnikov et al. 1995). As demonstrated above, the bat assemblage from Matuzka cave is divisible into two faunal associations, which describe different climatic periods of the terminal Pleistocene. Particular elements of the bat fauna can be used as biostratigraphic markers of time boundaries in Late Quaternary sediments of the Caucasus. Conclusions (1) Out of 23 bat species presently inhabiting the western Caucasus, 17 have been found in the fossil record. Absence of some species in Matuzka oryctocenosis (Rhinolophus hipposide ros, R. euryale, Myotis daubentonii, M. aurascens, M. mystacinus and Nyctalus lasiopterus) is most likely caused by taphonomic factors. Thus, by the end of the Middle Pleistocene, the 237 general appearance of the bat fauna had already been formed and remained almost constant to the present time. (2) Considerable part of the fauna is formed by lithophilous forms Eptesicus serotinus, Vespertilio murinus, Nyctalus noctula, supposedly inhabiting rocky fissures in the site. At least part of the sample apparently represents a taphocenosis coming from bird pellets. (3) The general dynamics of the number and structure of Pleistocene bat communities from Matuzka cave are in accordance with those of rodents inhabiting different landscapes and, hence, indirectly reflect environmental changes in the area of Matuzka cave. The bat fauna apparently decreased in time of spread of open-ground habitats. (4) In the Eemian Interglacial, the fauna of bats was the richest and included thermophilic R. ferrumeqiunum and Miniopterus schreibersii, besides of the records of Hypsugo savii and Pipistrellus kuhlii, which demonstrate in the Eemian the ranges corresponding to their Recent distribution. (5) The Valdai glaciation (the time of beds 6–3a accumulation) is characterized by a slightly poorer and less numerous bat fauna, which includes Myotis nattereri, M. emarginatus and M. bechsteinii, and is distinguished by well-pronounced fluctuations of the proportions of taxa. The proportion of psychrophilic faunal elements, such as Plecotus airutus and B. barbastellus, noticeably increased. SOUHRN Svrchnopleistocenní sled vrstev sedimentů jeskyně Matuzka, která se nachází na severním Kavkaze (Rusko), postihuje údobí od MIS6 po MIS2. V těchto vrstvách byly nalezeny zbytky nejméně 217 jedinců 18 druhů netopýrů. Faunový nález netopýrů je nebývale bohatý ve vrstvách odpovídajících Eemu a spodnímu Vistulianu. Ten je typický přítomností thermofilních prvků (Rhinolophus ferrumeqiunum a Miniopterus schreibersii) a širokým spektrem taxonů včetně dendrofilních a vzácných prvků (Plecotus auritus, Myotis brandtii, M. emarginatus, M. nattereri a M. blythii). Lithofilní formy (Eptesicus serotinus, Vespertilio murinus a Nyctalus noctula) se objevují nepřetržitě ve všech vrstvách a představují dominující složku společenstva. V eemských vrstvách jsou ale navíc druhy Hypsugo savii a Pipistrellus cf. kuhlii, které chybějí ve svrchních vrstvách, zatímco Pipistrellus pipistrellus se objevuje nejdříve až ve spodním holocenu. Acknowledgments We are heartily grateful to Professor A. K. Agadjanian (PIN RAS) for an all-round support in our research. We would also like to thank G. S. Rautian (PIN RAS) for valuable comments on the manuscript. This study was supported by the Presidium of the Russian Academy of Sciences (Program no. 25, “Biosphere Origin and Evolution”) and “Historical Dynamics of Bioresources As a Prerequisite for Their Conservation and Harmonious Exploitation,” the Russian Foundation for Basic Research (project nos. 04-05-64805 and 05-04-48493), and the Russian State Program for Support of Leading Scientific Schools (project no. NSh-6228.2006.4). 238 References Agadjanian А. K. & Rossina V. V., 2001: [Bats from Pleistocene deposites of the Denisova cave]. The Problems of Archeology, Ethnography, Anthropology of Siberia and Contiguous Territories (Novosibirsk), 7: 33–36 (in Russian, with a summary in English). Baryshnikov G. F. & Golovanova L. V., 1989: [Mammals of the Mousterian site Matuzka in the Kuban Caucasus]. Proc. Zool. Inst. AS USSR, 189: 3–55 (in Russian, with a summary in English). Benda P. & Tsytsulina K. A., 2000: Taxonomic revision of Myotis mystacinus group (Mammalia: Chiroptera) in Western Palearctic. Acta Soc. Zool. Bohem., 64: 331–398. Bobrinskij N. A., Kuznecov B. А. & Kuzjakin A. P., 1965: [Key to Identification of Mammals of the USSR] Prosvjaščenie, Moskva, 384 pp (in Russian). Filippov A. G. & Tiunov M. P., 1999: [Bat remains in caves of Irkutsk region]. Plecotus et al., 2: 100–108 (in Russian, with a summary in English). Gazaryan S. V., 2002: [The Ecological and Faunal Analyses of the Bat Populations of the Western Cau casus]. Unpublished PhD Thesis, Moskva, 216 pp (in Russian). Gazaryan S. V., 2003: [Current faunal status of the Daubenton’s bat Myotis daubentonii (Chiroptera, Vespertilionidae) in the Caucasus]. Plecotus et al., 6: 37–48 (in Russian, with a summary in English). Golovanova L., 1996: Paleolithic of the Northern Caucasus. Pp.: 260–261. In: The Workshops and the Posters of the XIII International Congress of Prehistoric and Prohistoric Scienses. Forli (Italy), 8–14 September 1996. Golovanova L. V., Baryshnikov G. F., Levkovskaya G. M. & Nesmeyanov S. A., 1995: [Multilayer Mousterian site Matuzka in the Northern Caucasus (results of research 1985–1989)]. Russian Archeology, 3: 105–118; 4: 77–86 (in Russian, with summaries in English). Horáček I. & Jahelková H., 2005: History of the Pipistrellus pipistrellus group in Central Europe in light of its fossil record. Acta Chiropterol., 7: 189–204. Kowalski K., 1995: Taphonomy of bats (Chiroptera). Geobios, M. S., 18: 251–256 Kuzjakin A. P., 1950: [Bats]. Sovetskaja Nauka, Moskva, 443 pp (in Russian). Nadachowski A. & Baryshnikov G., 1991: Pleistocene snow voles (Chionomys Miller, 1908) (Rodentia, Mammalia) from Northern Caucasus (USSR). Acta Zool. Cracov., 34: 437–451. Nesmeyanov S. A., 1999: [Geomorphological Aspects of Paleolithic Palaeoecology of the Western Cau casus]. Scientific World, Moskva, 392 pp (in Russian, with a summary in English). Obuch J., 1989: Chiropteran thanatocenoses in rocky fissures. P.: 453. In: Hanák V., Horáček I. & Gailser J. (eds): European Bat Research 1987. Charles University Press, Praha, 718 pp. Ovodov N. D., 1974: [The subfossil bat records in the caves of Siberia and the Far East]. Pp.: 84–90. In: Kuzjakin A. P. & Strelkov P. P. (eds.): [First All-Union Conference on Bats]. Zoological Institute AN USSR, Leningrad (in Russian). Potapova O. R., 1992: [Birds from the Mousterian site Matuzka in the North-Western Caucasus]. Proc. Zool. Inst. RAS, 246: 141–159 (in Russian, with a summary in English). Rossina V. V., 2002: [Odontological diversity of mouse-eared bats (Myotis) from Palaearctic]. Plecotus et al., Pars Spec.: 27–29 (in Russian, with a summary in English). Rossina V. V., 2004: [The dynamics of bat fauna (Chiroptera, Mammalia) of Northwest Altay in Pleistocene and Holocene]. Pp.: 208–218. In: [Ecological Mechanisms of Dynamics and Stability of Biota]. Istitute of Plants and Animals Ecology UrO RAS, Ekaterinburg (in Russian). Rossina V. V., 2005: [The influence of the anthropogenous factor on bats community in Pleistocene of Altai]. Pp.: 78–80. In: [Problems of Paleontology and Archeology of the South of Russia and Neigh bouring Territories. Materials of International Conference, Azov, May 18–20, 2005]. Rostov-na-Done (in Russian). Ruprecht A. L., 1990: [Bats (Chiroptera) in the food of owls in the Nadnotecka Forest]. Przegl. Zool., 34: 349–358 (in Polish, with a summary in English). Ruprecht A. L., 2005: [Some aspects of my research on bats (Chiroptera) of Poland in 1964–1990]. Leśne Prace Badawcze, 2005(2): 107–119. 239 Schmidt E. & Topal G., 1971: Fledermäuse in Eulengewöllen aus Ungarn. Vertebrata Hungar., 12: 93–102. (in Hungarian, with a summary in German). Tiunov M. P., 1997: [The Bats of the Far East of Russia]. Dal’nauka, Vladivostok, 134 pp (in Russian, with a summary in English). Vereščagin H. K., 1959: [Mammals of the Caucasus. History of the Fauna Forming]. Zoological Institute AN USSR, Moskva and Leningrad, 704 pp (in Russian). Zaitsev M. V. & Osipova V. A., 2004: [Insectivorous mammals (Insectivora) of the Late Pleistocene in the Northern Caucasus]. Zool. Žurnal, 83: 851–868 (in Russian, with a summary in English). 240 Lynx (Praha), n. s., 37: 241–246 (2006). ISSN 0024–7774 Plecotus macrobullaris – new bat species for Albanian fauna (Chiroptera: Vespertilionidae) Plecotus macrobullaris – nowy gatunek w faunie nietoperzy Albanii (Chiroptera: Vespertilionidae) Konrad SACHANOWICZ1 & Mateusz CIECHANOWSKI2 Department of Animal Ecology, Nicolaus Copernicus University, ul. Gagarina 9, PL–87-100 Toruń, Poland; chassan@poczta.onet.pl 2 Department of Vertebrate Ecology and Zoology, University of Gdańsk, al. Legionów 9, PL–80-441 Gdańsk, Poland; matciech@kki.net.pl 1 received on 27 July 2006 Abstract. Plecotus macrobullaris was recorded in the mountains of northern Albania (Pukë district) on 9–10 August 2003. This represents the new species of bat in the poorly known mammal fauna of Albania. At the reported locality, P. macrobullaris occurred in syntopy with P. auritus. Introduction The bat fauna of Albania includes ca. 24 species and belongs to the least studied among Eu ropean countries, with ca. 60% of species reported from single localities (Uhrin et al. 1996). Moreover, the presence of several species in the country is doubtful after recent changes in taxonomy of Myotis mystacinus and Pipistrellus pipistrellus complexes, as well as the genus Plecotus (Benda & Tsytsulina 2000, Jones & Barratt 1999, Kiefer & Veith 2001, Spitzen berger et al. 2001, 2003). Actually, four species of Plecotus are recognized in the continental Europe and all of them can be identified on the basis of external characters (Kiefer & von Helversen 2004, Tvrtković et al. 2005, Spitzenberger et al. 2006). Their geographical ranges overlap largely over the Balkan Peninsula. Previous records of Plecotus auritus (Linnaeus, 1758) and P. austriacus (Fischer, 1829) pub lished for Albania (Hanák et al. 1961, Hanák 1964, Lamani 1970) have not been documented sufficiently to reject the bat misidentification for their newly defined sibling species P. macrobul laris Kuzyakin, 1965 and P. kolombatovici Ðulić, 1980. The presence of P. auritus – a female collected in 1914 in Vermosha (Malësi e Madhe district) – has been confirmed recently in the result of molecular investigation (Spitzenberger et al. 2001). The occurrence of P. austriacus in Albania, although very probable, demands confirmation. Herein, we report the first record of P. macrobullaris from Albania and comment its identifi cation in the field. 241 Records On 9–10 August 2003, 7 individuals of long-eared bats (4 females, 3 males) identified as P. macrobullaris were mist-netted in the entrance of a small adit (42° 06’ N, 20° 07’ E), 787 m a.s.l., east of Qafëmal village, by the road Shkodra – Kukës (N Albania, Pukë district). The adit, cut in serpentinite rocks, has been used as a local water intake, containing a spring and a small pond in the entrance, which is located on the rocky mountain slope, sparsely overgrown with low black pines Pinus nigra and bushes (Fig. 1). There have been a small settlement and cultivations in the area nearby, although the dominating habitat in the surrounding was heavily overlogged mountain forest of the black pine and the beech Fagus sylvatica. Two adult females of P. auritus were caught at the same site, what allowed comparison of characters of both species held in a hand. After being identified, sexed, measured and photographed all bats were released. For the species identification in the field we have used characters given by Spitzenberger et al. (2002), Mucedda et al. (2002) and later confirmed by Tvrtković et al. (2005). Fig. 1. Entrance of the small adit near the Qafëmal village (K. Sachanowicz). Rys. 1. Wejście do sztolni w pobliżu miejscowości Qafëmal (K. Sachanowicz). 242 Table 1. External measurements and weight of P. macrobullaris individuals mist-netted in the entrance of the adit near Qafëmal (N Albania) on 9–10 August 2003 (F – female, M – male, ad. – adult, juv. – juvenile, lact. – lactating) Tab. 1. Wymiary zewnętrzne oraz ciężar osobników P. macrobullaris odłowionych przy otworze sztolni w pobliżu Qafëmal (północna Albania) 9–10 sierpnia 2003 (F – samica, M – samiec, ad. – dorosły, juv. – młody, lact. – karmiąca) No sex age 1 2 3 4 5 6 7 F F F F M M M ad., lact. juv. juv. ad. juv. juv. ad. mean±SD forearm length [mm] weight [g] thumb length [mm] claw length [mm] 41.6 41.0 41.4 41.0 38.5 40.8 39.4 9.0 7.0 7.0 7.5 7.0 7.5 7.0 7.2 7.0 7.1 – 7.0 7.0 7.0 2.6 2.8 1.9 – 2.9 2.7 2.9 40.53±1.14 7.43±0.73 7.05±0.08 2.63±0.38 Morphologically, individuals of P. macrobullaris resembled rather P. auritus than P. austriacus, but by the overall, contrasting coloration they were more similar to the latter species. Fur on dorsal and ventral sides was very dense, woolly and unusually long (ca. 10 mm), in which character the species differed markedly from P. auritus. The base of hairs on both sides was blackish. Dorsal fur was dark greyish-brown, strongly contrasting with almost pure white (whitish in some adults) on a ventral side. Juveniles had more greyish dorsal fur and brighter ventral side. The head and muzzle were greyish-brown, slightly darker than dorsal pelage. Protuberances above eyes were of the size (ca. 1.0–1.5 mm in diameter) similar to their equivalents in P. auritus. The triangular pad on a lower lip was obvious in all individuals of P. macrobullaris. It was pale, lip coloured, in adults and darker in juveniles (Fig. 2). The base of the tragus was pale pinkish, while its upper half was dark greyish. The tragus width in two adult females was 5.3 and 5.9 mm. The size of a hind foot was almost the same as in P. auritus; feet were also covered with well visible hairs. Penis was broad (ca. 2.2 mm), cylindrical and parallel sided at almost whole length with a pointed tip. A thumb of P. macrobullaris was slightly longer than in two P. auritus (6.3 and 6.0 mm), while a claw was of similar size (2.6 and 2.5 mm, cf. Table 1). External measurements and weight of P. macrobullaris are given in Table 1. comments P. macrobullaris occurs in mountain regions from central Pyrenees to Bosnia and Herzegovina and from central Greece to the Caucasus, Syria and Iran, with a single records from Corsica and Crete (Garin et al. 2003, Juste et al. 2004, Kiefer & von Helversen 2004, Spitzenberger et al. 2006). Astonishingly, in the Balkans the species is known only from the western part of the Peninsula (Slovenia, Croatia, Bosnia and Herzegovina, Greece), while no localities were found in its eastern regions e.g. in Bulgaria (Benda & Ivanova 2003). The Albanian record confirms continuous distribution of P. macrobullaris in the Dinarian Mts, as well as its syntopic occur rence with P. auritus. However, the species apparently is not restricted to higher (above 800 m) altitudes, at least in the Dinarian Mts, as it has been suggested (Kiefer & Veith 2002, Kiefer & von Helversen 2004). In Croatia, it was recorded within altitudinal range from the sea level up to 1800 m a.s.l., with most localities – like the only Albanian – located below 800 m a. s. 1. (Pavlinić & Tvrtković 2004). In contrast to other members of the genus, P. macrobullaris in 243 Table 2. Comparison of the forearm length of P. macrobullaris from different parts of its range. Data source: Caucasus and Turkey – Spitzenberger et al. (2003); Greece, Liechtenstein and French Alps (holo type) – Kiefer & Veith (2001), P. Benda, unpubl.; Austria – Spitzenberger et al. (2002); Pyrenees – Garin et al. (2003), P. Benda, unpubl.; Iran, Syria and Switzerland – P. Benda, unpubl.; Croatia – Tvrtković et al. (2005); Albania – this paper Tab. 2. Porównanie długości przedramienia osobników P. macrobullaris z różnych części zasięgu. Źródła danych: Kaukaz i Turcja – Spitzenberger et al. (2003); Grecja, Liechtenstein i Alpy Francuskie (holotyp) – Kiefer & Veith (2001), P. Benda, unpubl.; Austria – Spitzenberger et al. (2002); Pireneje – Garin et al. (2003), P. Benda, unpubl.; Iran, Syria i Szwajcaria – P. Benda, unpubl.; Chorwacja – Tvrtković et al. (2005); Albania – niniejszy artykuł region mean Caucasus Iran Syria Turkey Greece Albania Croatia Austria Liechtenstein Switzerland French Alps Pyrenees 42.77 41.73 41.30 – 39.85 39.57 39.95 40.59 – 41.55 – 42.00 males SD min 0.48 1.63 0.96 – 0.21 1.16 1.57 0.59 – 1.20 – 1.13 42.1 39.6 40.2 – 39.7 38.5 37.3 39.6 – 40.7 – 41.3 max n mean 43.2 6 43.2 6 42.0 3 40.9 1 40.0 2 40.8 3 42.5 25 41.5 7 – – 42.4 2 40.5 1 43.3 3 43.10 42.80 42.68 42.67 – 41.25 41.22 41.91 – 40.67 – 42.55 females SD min 1.45 1.32 1.44 1.23 – 0.30 1.26 0.86 – 0.60 – 0.66 40.7 41.8 39.8 40.5 – 41.0 39.0 40.5 – 40.1 – 42.1 max n 44.2 5 45.1 5 44.6 11 44.2 9 – – 41.6 4 43.5 30 43.5 11 39.7 1 41.7 6 39.6 1 43.5 4 Croatia is restricted to karstic areas, where its coexistence with P. auritus was observed at three sites (Tvrtković et al. 2005). The forearm length of Albanian P. macrobullaris includes within the variation range of spe cimens from the Alps and Croatia (Kiefer & Veith 2001, Spitzenberger et al. 2002, Tvrtković et al. 2005). Individuals from the Pyrenees, Caucasus and the Middle East seem to have longer forearms (Spitzenberger et al. 2006, Table 2). From the other hand, bats from Albania have signi ficantly longer thumb than the Pyrenean individuals of P. macrobullaris (Mann-Whitney U-test, U=0, p<0.005, n=12; data from Garin et al. 2003 and Table 1). Our data, as well as those from Spain (Garin et al. 2003) seem to contradict that thumb and claw of P. macrobullaris are shorter than in P. auritus (Spitzenberger et al. 2006). The coloration of the tragus, facial mask as well as contrasting greyish-brown and white pelage of Albanian bats correspond with characters of specimens from the Alps and Croatia (Kiefer & Veith 2001, Tvrtković et al. 2005) but differ from the Pyrenean and other populations (Garin et al. 2003, Spitzenberger et al. 2006). Finally, measurements and external characters of Albanian bats may suggest their affiliation to smaller subspecies P. m. alpinus distributed from the Alps to Bosnia and Herzegovina (Spitzenberger et al. 2003, 2006, Kiefer & von Helversen 2004). The Pyrenean population shows measurements and characters more similar to larger P. m. macrobullaris ranging from the north-eastern Italy to the Caucasus, Syria and Iran (Kiefer & von Helversen 2004, Spitzenberger et al. 2006, Table 2), but the results of genetic analysis have placed that population within the western subclade (Spitzenberger et al. 2006). Therefore, the zone at least from the eastern Alps to the southern Dinarian Mts should be inhabited by sympatric western and eastern populations. Moreover, this 244 Fig. 2. Plecotus macrobullaris from the area of Qafëmal, northern Albania (K. Sachanowicz). Rys. 2. Plecotus macrobullaris z okolic Qafëmal w północnej Albanii (K. Sachanowicz). may contradict the statement that in P. macrobullaris the forearm length decreases towards the west (Spitzenberger et al. 2003), suggesting rather different pattern of geographical variation – with smaller individuals in central part of the range and larger in its marginal parts. However, larger samples for morphometric and molecular studies are required to solve the problem of external variation and differences in skull measurements (Spitzenberger et al. 2003, 2006) among populations of P. macrobullaris from different parts of its range. Streszczenie Nowy gatunek dla fauny nietoperzy Albanii – Plecotus macrobullaris – został stwierdzony w górach północnej części kraju (region Pukë) 9–10 sierpnia 2003. Na opisanym stanowisku gacek alpejski współwystępował z gackiem brunatnym Plecotus auritus. Acknowledgements We thank Petr Benda for unpublished morphological data, comments on the manuscript and help in obtaining two rare publications. References Benda P. & Ivanova T., 2003: Long-eared bats, genus Plecotus (Mammalia: Chiroptera), in Bulgaria: a revision of systematic and distributional status. J. Natl. Mus., Natur. Hist. Ser., 172: 157–172. 245 Benda P. & Tsytsulina K., 2000: Taxonomic revision of Myotis mystacinus group (Mammalia: Chiroptera) in the western Palearctic. Acta Soc. Zool. Bohem., 64: 331–398. Garin I., García-Mudarra J. L., Aihartza J. R., Goiti U. & Juste J., 2003: Presence of Plecotus macro bullaris (Chiroptera: Vespertilionidea) in the Pyrenees. Acta Chiropterol., 5: 243–250. Hanák V., 1964: Zur Kenntnis der Fledermäusfauna Albaniens. Věst. Čs. Společ. Zool., 28: 68–88. Hanák V., Lamani F. & Muraj X., 1961: Të dhëna nga përhapja e lakuriqëve të natës (Ordo Chiroptera) në Shqipëri [The results of the research work carried out on the bats in our country]. Bul. Univ. Shtetëror Tiranës, Ser. Shk. Natyrore, 3: 124–156 (in Albanian, with a summary in English). Jones G. & Barratt E. M., 1999: Vespertilio pipistrellus Schreber, 1774 and V. pygmaeus Leach, 1825 (currently Pipistrellus pipistrellus and P. pygmaeus; Mammalia, Chiroptera): proposed designation of neotypes. Bull. Zool. Nomencl., 56: 182–186. Juste J., Ibáñez C., Muñoz J., Trujillo D., Benda P., Karataş A. & Ruedi M., 2004. Mitochondrial phylo geography of the long-eared bats (Plecotus) in the Mediterranean and Atlantic Islands. Mol. Phylogenet. Evol., 31: 1114–1126 Kiefer A. & von Helversen O., 2004: Plecotus macrobullaris (Kuzjakin, 1965) – Alpenlangohr. Pp.: 1051–1058. In: Krapp F. (ed.): Handbuch der Säugetiere Europas. Band 4: Fledertiere. Teil II: Chiro ptera II. Vespertilionidae 2, Molossidae, Nycteridae. Aula-Verlag, Wiebelsheim, x+528 pp. Kiefer A. & von Helversen O., 2004: Bestimmungsschlüssel und Kurzbeschreibung der europäischen Langohren. Pp.: 947–952. In: Krapp F. (ed.): Handbuch der Säugetiere Europas. Band 4: Fledertiere. Teil II: Chiroptera II. Vespertilionidae 2, Molossidae, Nycteridae. Aula-Verlag, Wiebelsheim, 528 pp. Kiefer A. & Veith M., 2001: A new species of long-eared bat from Europe (Chiroptera: Vespertilionidae). Myotis, 39: 5–16. Lamani F., 1970: Lloje të reja lakuriqesh nate në vëndin tonë [Nouvelles especes de chauves-souris dans notre pays]. Bul. Shk. Natyrore, 2: 143–150 (in Albanian, with a summary in French). Mucceda M., Kiefer A., Pidinchedda E. & Veith M., 2002: A new species of long-eared bat (Chiroptera, Vespertilionidae) from Sardinia (Italy). Acta Chiropterol., 4: 121–136. Pavlinić I. & Tvrtković N., 2004: Altitudinal distribution of four Plecotus species (Mammalia, Vesperti lionidae) occurring in Croatia. Natura Croat., 13: 395–401. Spitzenberger F., Piálek J. & Haring E., 2001: Systematics of the genus Plecotus (Mammalia, Vespertili onidea) in Austria based on morphometric and molecular investigations. Folia Zool., 50: 161–172. Spitzenberger F., Haring E. & Tvrtković N., 2002: Plecotus microdontus (Mammalia, Vespertilionidea), a new bat species from Austria. Natura Croat., 11: 1–18. Spitzenberger F., Strelkov P. P. & Haring E., 2003: Morphology and mitochondrial DNA sequences show that Plecotus alpinus Kiefer & Veith, 2002 and Plecotus microdontus Spitzenberger, 2002 are synonyms of Plecotus macrobullaris Kuzjakin, 1965. Natura Croat., 12: 39–53. Spitzenberger F., Strelkov P. P., Winkler H. & Haring E., 2006: A preliminary revision of the genus Plecotus (Chiroptera, Vespertilionidae) based on genetic and morphological results. Zool. Scripta, 35: 187–230. Tvrtković N., Pavlinić I. & Haring E., 2005: Four species of long-eared bats (Plecotus, Geoffroy, 1818; Mammalia, Vespertilionidae) in Croatia: field identification and distribution. Folia Zool., 54: 75–88. Uhrin M., Horáček I., Šíbl J. & Bego F., 1996: On the bats (Mammalia: Chiroptera) of Albania: survey of the recent records. Acta Soc. Zool. Bohem., 60: 63–71. 246 Lynx (Praha), n. s., 37: 247–254 (2006). ISSN 0024–7774 Supplementary notes on the distribution of Pipistrellus pipistrellus complex in the Balkans: first records of P. pygmaeus in Albania and in Bosnia and Herzegovina (Chiroptera: Vespertilionidae) Uzupełnienia do rozmieszczenia nietoperzy z kompleksu Pipistrellus pipistrellus na Bałkanach: pierwsze stwierdzenia P. pygmaeus w Albanii oraz w Bośni i Hercegowinie (Chiroptera: Vespertilionidae) Konrad Sachanowicz1, Mateusz Ciechanowski2 & Alek Rachwald3 1 Department of Animal Ecology, Nicolaus Copernicus University, ul. Gagarina 9, PL–87-100 Toruń, Poland; chassan@poczta.onet.pl 2 Department of Vertebrate Ecology and Zoology, University of Gdańsk, al. Legionów 9, PL–80-441 Gdańsk, Poland; matciech@kki.net.pl 3 Department of Forest Ecology and Wildlife Management, Forest Research Institute, ul. Bitwy Warszawskiej 3, PL–00-973 Warszawa, Poland; merlin@las.ibl.bialowieza.pl received on 28 April 2006 Abstract. Two males Pipistrellus pygmaeus were captured in August 2003 near Qafëmal (northern Al bania) and identified on the basis of morphological characters. Hunting individuals of this species were observed and detected also in a riparian forest of the Vjosës valley near Tepelenë (southern Albania) in April 2004. They represent the first records of this species from Albania. P. pygmaeus is reported for the first time also from Bosnia and Herzegovina, where echolocation calls of the species (lowest frequencies 52–57 kHz) were recorded near Miljevina and in Dobro Polje. Introduction Since the time of recognition of the pygmy (soprano) pipistrelle Pipistrellus pygmaeus (Le ach, 1825) within the P. pipistrellus complex (Barratt et al. 1997, Jones & Barratt 1999), its distribution has been intensively surveyed across Europe. Soon, it was discovered that the species ranges widely from Portugal to western part of Russian Federation and the Caucasus, and from Great Britain, southern Scandinavia and Estonia to Greece and Turkey (Horáček et al. 2000, Limpens 2000, Mayer & von Helversen 2001, Benda et al. 2003a, Hulva et al. 2004, Wermundsen & Siivonen 2004, Vierhaus & Krapp 2004). Although widespread, the range of this species in the Balkans is known only fragmentary due to the low intensity of bat surveys there in general. Actually, its presence has been confirmed for most of the Balkan countries (Limpens 2000, 2001, Presetnik et al. 2001, Dietz et al. 2002, Benda et al. 2003b, Decu 2003, Hulva et al. 2004). As P. pygmaeus was found at numerous localities in Greece (Hanák et al. 2001), including its northern parts, it was expected to occur also in Albania and in the countries of former Yugoslavia, where it has not been recorded yet. Hence, taking into account the present state of knowledge, the range of pygmy pipistrelle in southern Balkans, depicted by von Helversen & Holderied (2003) and reprinted by Vierhaus 247 Fig. 1. Male of Pipistrellus pygmaeus captured on 10 August 2003 near Qafëmal, northern Albania (K. Sachanowicz). Rys. 1. Samiec Pipistrellus pygmaeus odłowiony 10 sierpnia 2003 w pobliżu Qafëmal w północnej Albanii (K. Sachanowicz). & Krapp (2004) is misleading as giving an impression of its confirmed occurrence in southern half of Albania and in Macedonia. This tentative range also covers a belt of southern Bulgaria althought only one record was known by then from southernmost part of the country (Dietz et al. 2002). Single old records of bats belonging to small Pipistrellus complex are known from Macedo nia, Montenegro as well as from Bosnia and Herzegovina (e.g. Bolkay 1926, Đulić & Mirić 1967, Kryštufek et al. 1992, Jones 1999). The only record for Albania, published recently without any details – a male captured in a flat in Tirana – has to be assigned to P. pipistrellus sensu lato (Uhrin et al. 1996). In addition to the present knowledge of P. pygmaeus distribution, herein we report first data on the species presence in Albania and in Bosnia and Herzegovina. Material and methods Data were collected in the Balkans during the trips in 2003–2004. Hunting bats were detected using a heterodyne detector Pettersson D100 and recorded with broadband Pettersson D980 detector and SONY 248 WM-D6C tape recorder. Recordings were analysed using BatSound 3.1 computer software (Pettersson Elektronik AB, Sweden). For identification of the time-expanded calls we relied on characters given by Zingg (1990), Russo & Jones (2002) and Obrist et al. (2004), while for determination of captured indivi duals – we followed Häussler et al. (1999), Sendor et al. (2002), von Helversen & Holderied (2003), and Dietz & von Helversen (2004). ReCORDS P. pygmaeus was recently recorded at two localities in Albania (1–2) and at two sites in Bosnia and Her zegovina (3–4): (1) On 10 August 2003, two adult, sexualy active males were mist-netted over a small artificial pool (42° 06’ N, 20° 06’ E), 731 m a. s. l., in a bed of a mountain stream, east of the Qafëmal village, along the road Shkodra–Kukës (northern Albania, Pukë district). The habitat around the site comprised a small settlement, cultivations and overlogged mountain forest of the black pine Pinus nigra and the beech Fagus sylvatica. More individuals of P. pygmaeus were observed and detected (Pettersson D100) at dusk when hunting over the stream and sparse vegetation covering surrounding mountain slopes. Measurements of two males were as follow: forearm length – 31.0 and 29.6 mm, fifth finger length – 39.0 and 38.1 mm and weight – 5.5 and 5.0 g, respectively. Their pelage was light brown with yellowish tinge on the back and with weak contrast between dorsal and ventral side, which was whitish-grey with yellowish tinge. The muzzle and ears (7.9 mm) were relatively short (Fig. 1), light brown but slightly Fig. 2. The sonogram and power spectrum graph of time-expanded echolocation calls of P. pygmaeus recorded near Tepelenë, southern Albania, on 19 April 2004. Rys. 2. Sonogram i wykres widma mocy sygnałów echolokacyjnych P. pygmaeus nagranych w okolicy Tepeleny w południowej Albanii, 19 kwietnia 2004. 249 darker than the pelage; the skin was lighter, pinkish particularly around the eyes and in the basal part of the ears. Both individuals had pronounced small ridge between the nostrils. Penis was yellowish without paler medial stripe. Glandular bumps inside of mouth were of similar yellowish colour. Basal part of the uropatagium was densely covered with fur on dorsal side, a condition usually seen in P. nathusii (Keyser ling et Blasius, 1839). After being identified, sexed, measured and photographed the bats were released. (2) P. pygmaeus was also recorded in the Vjosës valley, ca 1 km south of Tepelenë (40° 17’ N, 20° 02’ E), 114 m a. s. l. Soon before the dusk (19.10 p.m.), on 19 April 2004, several individuals were observed and detected with Pettersson D980 (Fig. 2), while hunting above a small branch of the river and near the tree crowns in an old stand of oriental planes Platanus orientalis and poplars Populus sp. Apparently, these bats were shortly after leaving their roost, which could be located in a tree hole or crevice, since no other type of shelters was available nearby. The mean parameters of recorded calls were: frequency of maximum intensity: 57.2 kHz (range 55–60 kHz, n=12), highest frequency: 78.8 kHz (range 67–99 kHz, n=8), lowest frequency: 55.4 kHz (range 54–57 kHz, n=9), pulse length: 5.5 ms (range 4–7 ms, n=8), interval length: 74.4 ms (range 61–78 ms, n=10). Fig. 3. The sequence of time-expanded search calls of P. pygmaeus (upper row of signals), recorded near Miljevina, Bosnia and Herzegovina, on 2 May 2004. Frequency of maximum intensity shown in a power spectrum graph is marked with a cross. Lower row of signals illustrates echolocation calls of Hypsugo savii recorded simultaneously. High background noise level caused by turbulent water. Rys. 3. Sekwencja sygnałów echolokacyjnych P. pygmaeus, nagranych w okolicach Miljewiny, w Bośni i Hercegowinie, 2 maja 2004 (górny szereg). Krzyżyk na wykresie widma mocy wskazuje częstotliwość maksymalnego natężenia dźwięku. Poniżej, na sonogramie, nagrane jednocześnie sygnały Hypsugo savii. Wysoki poziom szumów tła powodowany był przez płynącą wodę. 250 (3) Miljevina (43° 31’ N, 18° 39’ E), 599 m a. s. l., approximately 2.5 km south of the village, in a rocky canyon of Bistrica river (100–150 m in depth) on the Dinarian Mts, eastern Bosnia. The site was surroun ded by almost vertical limestone walls and steep slopes, partially overgrown with sparse mixed forest (spruces and beeches). At sunset on 2 May 2004, a search calls of hunting individuals of P. pygmaeus were detected and recorded with Pettersson D980 bat detector and the tape recorder. The signals were FM-qcf calls (Fig. 3) of the following mean parameters: frequency of maximum in tensity: 57.2 kHz (range 55–62 kHz, n=12), highest frequency: 72.4 kHz (range 64–85 kHz, n=9), lowest frequency: 54.2 kHz (range 52–57 kHz, n=9), pulse length: 5.8 ms (range 5–7 ms, n=9), interval length: 76 ms (range 70–90 ms, n=11). (4) Later, in the same night as above, echolocation and social calls of P. pygmaeus (composed of three elements at frequency of 18 kHz) were recorded near Dobro Polje (43° 35’ N, 18° 31’ E), 1008 m a. s. l., ca. 0.5 km west of the village, at a bridge over Bistrica river, eastern Bosnia. Discussion Individuals captured near Qafëmal fitted well into the range of morphological variation of P. pygmaeus, given by the most recent authors (Häussler et al. 1999, von Helversen & Holde ried 2003, Dietz & von Helversen 2004), thus their identification, in this case also supported by detector observation, appeared unambiguous. More attention should be paid to the species determination based on recorded echolocation calls. Although in the northern Europe, acoustic identification of P. pygmaeus is one of the easiest among vespertilionid bats, it is much more complicated in the Meditterranean zone, where the species occurs sympatrically with Miniopterus schreibersii (Kuhl, 1817) and both these species often share the same hunting habitats (Russo & Jones 2003). Noteworthy, although M. schreibersii is taxonomically distant and belongs to a different family (Miniopteridae), it reveals echolocation calls suprisingly similar to those emitted by P. pygmaeus. Its structure (FM-qcf) strongly resembles typical pipistrelle signals. Mean value of its frequency of maximum intensity (53.9 kHz) is slightly lower than that recorded in P. pygmaeus (56.2 kHz), similarly as the lowest frequency (47.4 and 51.5 kHz, respectively – Obrist et al. 2004). However, ranges of these parameters overlapp (frequency of maximum intensity: M. schreibersii 49–62 kHz, P. pygmaeus 53–63 kHz; lowest frequency: 47–55 kHz and 53–60 kHz, respectively – Russo & Jones 2002), thus large portion of their calls from sou thern Europe can be determined only with multivariate statistical methods, e.g. discriminant function analysis (Russo & Jones 2002) or synergetic pattern recognition (Obrist et al. 2004). Identification of pygmy pipistrelle with narrowband heterodyne detectors is widely practised in Europe (e.g. Limpens 2000, 2001, Wermundsen & Siivonen 2004), but in the Mediterranean it might be reliable only when supported with visual determination (flight and hunting style, size of the bat and its wings shape – short and moderately broad in P. pygmaeus vs. long and narrow in M. schreibersii). The lowest frequencies of some P. pygmaeus signals recorded in Albania and in Bosnia were higher than the upper range of this parameter in M. schreibersii (see also Zingg 1990), so the determination may be regarded as unambiguous. In the second Bosnian locality (Dobro Polje), the pygmy pipistrelle was also confirmed by recorded social calls with their typical, three-element structure (Barlow & Jones 1997). The range of P. pygmaeus covers most likely the whole territory of the Balkan Peninsula. Its relatively numerous localities are known only from Slovenia (Presetnik et al. 2001, Kryštufek & Donev 2005) and from Greece (Hanák et al. 2001, Mayer & von Helversen 2001), that both belong to fairly well studied countries. For other Balkan states, mostly single records are avai lable, mainly due to the insufficient recording and low intensity of acoustic surveys. Therefore, 251 several localities, scattered over southern and western regions, were published for Romania (Limpens 2000, Decu 2003, Hulva et al. 2004). Single records are known also for Croatia and Serbia (Limpens 2001), Bulgaria (Dietz et al. 2002, 2005, Benda et al. 2003b) and for the European part of Turkey (Benda et al. 2003a). Supplementing the picture of P. pygmaeus distribution in this part of Europe, we found the species in the least surveyed countries: Albania and Bosnia and Herzegovina (this paper). Recently, the species was recorded also in Montenegro, where two foraging bats were observed and detected on 10 July 2004 near the old fortress of Budva, located on the Adriatic coast (C. Dietz – pers. comm.). At present, only Macedonia has no presence of this species confirmed on its territory, for which only records of P. pipistrellus (Schreber, 1774) s. str. are known (Kryštufek et al. 1992, B. Kryštufek – pers. comm.). Data on habitat use by the species in the Balkans are also scarce. In Slovenia, P. pygmaeus was observed hunting mainly in riparian habitats, over ponds, lakes and rivers, below 650 m a. s. l. (Presetnik et al. 2001). In all cases described above, P. pygmaeus was recorded in the vicinity of water, over small pool and in forested river valleys, what is in accordance with its habitat preferences in other parts of Europe (Russo & Jones 2003, Bartonička & Řehák 2004). Finally, the presence of both small pipistrelles has been confirmed for Albania, as P. pipist rellus s. str. was recorded recently for the first time at several localities (authors’ unpublished data). It is obvious that both these Pipistrellus species live in sympatry also in other Balkan countries, as it was reported for Slovenia, Croatia, Serbia, Romania, Bulgaria and for Greece (Limpens 2000, 2001, Hanák et al. 2001, Mayer & von Helversen 2001, Presetnik et al. 2001, Benda et al. 2003, Decu 2003, Kryštufek & Donev 2005). However, relevant data on P. pygmaeus abundance are available only for Slovenia, where it appears less frequent (44 localities) than P. pipistrellus (99 sites) (Kryštufek & Donev 2005). On the other hand, Hanák et al. (2001) suggest that in Greece the former species is more abundant, but this conclusion is based on small number of records (15 and 6, respectively), hence it may be misleading due to the insufficient recording. Streszczenie Dwa samce Pipistrellus pygmaeus odłowiono w sierpniu 2003 w pobliżu miejscowości Qafëmal (północna Albania). Żerujące osobniki tego gatunku obserwowano w nadrzecznym łęgu w dolinie Vjosy w okoli cach Tepeleny (południowa Albania) w kwietniu 2004. Są to pierwsze stwierdzenia karlika drobnego w Albanii. P. pygmaeus został także po raz pierwszy stwierdzony w Bośni i Hercegowinie, gdzie w maju 2004, w okolicach Miljewiny i Dobrego Polja nagrano sygnały echolokacyjne tego gatunku. Acknowledgements Authors express cordial thanks to Boris Kryštufek for valuable information on P. pipistrellus distribution in Macedonia, giving an access to some hardly available publications and comments on the manuscript, while to Christian Dietz for his own unpublished observation from Montenegro. References Barlow K. E. & Jones G., 1997: Differences in songflight calls and social calls between two phonic types of the vespertilionid bat Pipistrellus pipistrellus. J. Zool., London, 241: 315–324. Barratt E. M., Deaville R., Burland T. M., Bruford M. W., Jones G., Racey P. A. & Wayne R. K., 1997: DNA answers the call of pipistrelle bat species. Nature, 387: 138–139. 252 Bartonička T. & Řehák Z., 2004: Flight activity and habitat use of Pipistrellus pygmaeus in a floodplain forest. Mammalia, 68: 365–375. Benda P., Hulva P., Andreas M. & Uhrin M., 2003a: Notes on the distribution of Pipistrellus pipistrellus complex in the Eastern Mediterranean: First records of P. pipistrellus for Syria and of P. pygmaeus for Turkey. Vespertilio, 7: 87–95. Benda P., Ivanova T., Horáček I., Hanák V., Červený J., Gaisler J., Gueorguieva A., Petrov B. & Vo hralík V., 2003b: Bats (Mammalia: Chiroptera) of the Eastern Mediterranean. Part 3. Review of bat distribution in Bulgaria. Acta Soc. Zool. Bohem., 67: 245–357. Bolkay S. J., 1926: Additions to the Mammalian Fauna of the Balkan Peninsula. Glasn. Zem. Muz. Bosni i Hercegovini (Sarajevo), 38: 159–179. Decu V., 2003: Atlasul chiropterolor din România. Pp.: 201–352. In: Decu V., Murariu D. & Gheorghiu V. (eds.): Chiroptere din România. Ghid instructiv şi educativ. Institutul de Speologie “Emil Racoviţă” al Academiei Române, Muzeul Naţional de Istorie Naturală “Grigore Antipa”, Bucureşti, 522 pp. Dietz C., Schunger I., Nill D., Siemers B. M. & Ivanova T., 2002: First record of Pipistrellus pygmaeus (Leach, 1825) (Chiroptera: Vespertilionidae) for Bulgaria. Hist. Natur. Bulg., 14: 117–121. Dietz C., Schunger I., Keşapli-Didrickson Ö., Karataş A. & Mayer F., 2005: First record of Pipistrellus pygmaeus (Chiroptera: Vespertilionidae) in Anatolia. Zool. Middle East, 34: 5–10. Dietz C. & von Helversen O., 2004: Illustrated Identification Key to the Bats of Europe. Electronic publication. Version 1.0, Tuebingen & Erlangen, 72 pp. Đulić B. & Mirić Đ., 1967: Catalogus Faunae Jugoslaviae. IV/4 Mammalia. Acad. Sci. Art. Slov., Ljubljana, 46 pp. Hanák V., Benda P., Ruedi M., Horáček I. & Sofianidou T. S., 2001: Bats (Mammalia: Chiroptera) of the Eastern Mediterranean. Part 2. New records and review of distribution of bats in Greece. Acta Soc. Zool. Bohem., 65: 169–279. Häussler U., Nagel A., Braun M. & Arnold A., 1999: External characters discriminating sibling species of European pipistrelles, Pipistrellus pipistrellus and P. pygmaeus. Myotis, 37: 27–40. von Helversen O. & Holderied M., 2003: Zur Unterscheidung von Zwergfledermause (Pipistrellus pi pistrellus) und Mückenfledermause (Pipistrellus mediterraneus / pygmaeus) im Feld. Nyctalus (N. F.), 8: 420–426. Horáček I., Hanák V. & Gaisler J., 2000: Bats of the Palearctic Region: a taxonomic and biogeographic review. Pp.: 11–157. In: Wołoszyn B. W. (ed.): Proceedings of the VIIIth EBRS. Vol. 1. Approaches to bi ogeography and ecology of bats. Chiropterological Information Center, ISEZ PAN, Kraków, 278 pp. Hulva P., Horáček I., Strelkov P. P. & Benda P., 2004: Molecular architecture of Pipistrellus pipistrellus / Pipistrellus pygmaeus complex (Chiroptera: Vespertilionidae): further cryptic species and Mediterra nean origin of the divergence. Mol. Phylogenet. Evol., 32: 1023–1035. Jones G., 1999: Pipistrellus pipistrellus (Schreber, 1774). Pp.: 126–127. In: Mitchell-Jones A. J., Amori G., Bogdanowicz W., Kryštufek B., Reinjders P. J. H., Spitzenberger F., Stubbe M., Thissen J. B. M., Vohralík V. & Zima J. (eds.): The Atlas of European Mammals. Academic Press, London, 484 pp. Jones G. & Barratt E. M., 1999: Vespertilio pipistrellus Schreber, 1774 and V. pygmaeus Leach, 1825 (currently Pipistrellus pipistrellus and P. pygmaeus; Mammalia, Chiroptera): proposed designation of neotypes. Bull. Zool. Nomencl., 56: 182–186. Kryštufek B. & Donev R. N., 2005: The atlas of Slovenian bats (Chiroptera). Scopolia, 55: 1–92. Kryštufek B., Vohralík V., Flousek J. & Petkowski S., 1992: Bats (Mammalia: Chiroptera) of Macedonia, Yugoslavia. Pp.: 93–111. In: Horáček I. & Vohralík V. (eds.): Prague Studies in Mammalogy. Charles University Press, Praha, 246 pp. Limpens H. J. G. A., 2000: Report on the program of bat detector training workshops in Bulgaria and Croatia in 1999 and in Ukraine, Georgia, Slovenia, Romania and Moldova in 2000. Unpublished report. Eco Consult & Project Management, Wageningen, 41 pp. Limpens H. J. G. A., 2001: Report on the program of bat detector training workshops in Lithuania/Baltic region, Slovakia and Yugoslavia in 2001. Unpublished report. Eco Consult & Project Management, Wageningen, 28 pp. 253 Mayer F. & von Helversen O., 2001: Sympatric distribution of two cryptic bat species across Europe. Biol. J. Linn. Soc., 74: 365–374. Obrist M., Boesch R. & Flückiger P. F., 2004: Variability in echolocation call design of 26 Swiss bat species: consequences, limits and options for automated field identification with a synergetic pattern recognition approach. Mammalia, 68: 307–322. Presetnik P., Koselj K. & Zagmajster M., 2001: First records of Pipistrellus pygmaeus (Leach, 1825) in Slovenia. Myotis, 39: 29–32. Russo D. & Jones G., 2002: Identification of twenty-two bat species (Mammalia: Chiroptera) from Italy by analysis of their echolocation calls. J. Zool., London, 258: 91–103. Russo D. & Jones G., 2003: Use of foraging habitats by bats in a Mediterranean area determined by acoustic surveys: conservation implications. Ecography, 26: 197–209. Sendor T., Roedenbeck I., Hampl S., Ferreri M. & Simon M., 2002: Revision of morphological identifica tion of pipistrelle bat phonic types (Pipistrellus pipistrellus Schreber, 1774). Myotis, 40: 11–17. Uhrin M., Horáček I., Šíbl J. & Bego F., 1996: On the bats (Mammalia: Chiroptera) of Albania: survey of the recent records. Acta Soc. Zool. Bohem., 60: 63–71. Wermundsen T. & Siivonen Y., 2004: Distribution of Pipistrellus species in Finland. Myotis, 41–42: 93–98. Vierhaus H. & Krapp F., 2004: Pipistrellus mediterraneus (Cabrera, 1904) oder P. pygmaeus (Leach, 1825) – Mückenfledermaus. Pp.: 815–823. In: Krapp F. (ed.): Handbuch der Säugetiere Europas. Band 4: Fledertiere. Teil II: Chiroptera II. Vespertilionidae 2, Molossidae, Nycteridae. Aula-Verlag, Wiebelsheim, x+582 pp. Zingg P. E., 1990: Akustiche Artidentifikation von Fledermäusen (Mammalia: Chiroptera) in der Schweiz. Rev. Suisse Zool., 97: 263–294. 254 Lynx (Praha), n. s., 37: 255–262 (2006). ISSN 0024–7774 Small mammal fauna of the Kraków metropolitan area (southern Poland) – problem of synurbisation (Insectivora, Chiroptera, Rodentia) Drobne ssaki dawnego województwa krakowskiego – problem synurbizacji (Insectivora, Chiroptera, Rodentia) Katarzyna Stanik & Bronisław W. Wołoszyn Institute of Systematics and Evolution of Animals, Polish Academy of Sciences, ul. Sławkowska 17, PL–31-016 Kraków, Poland; maukasiek@o2.pl; woloszbr@isez.pan.krakow.pl received on 6 June 2006 Abstract. Current knowledge regarding the occurrence of small mammal species in the Kraków agglomeration and adjacent areas is presented. Special attention is given to urbanised species and their population characteristics. INTRODUCTION Studies in urban ecology have been carried out in Poland for at least 30 years. Their aim is an assessment of the current state and future prospects regarding the natural environment in urban and suburban areas, and its role in creating optimal living conditions for the urban population of people. Therefore the research is focused on three major aspects: soil science, flora and fauna (Karolewski 1981). The occurrence, abundance and structure of animal assemblages in urbicenosis are not intentionally shaped by man, in contrast to floristic ones (Andrzejewski 1975). However, such factors as climate, technical infrastructure, flora, human population and environmental pollution have a deep impact – termed “urbanisation pressure” – on urban fauna and its distribution (Markowski 1997). Observation of its effects has led to creation of new terms in urban ecology. One of these is “antropopressure” defined as a mode of human impact on natural environment. Another term is “synanthropisation”, covering a whole range of changes in flora and fauna. Organisms that are well adapted to new conditions are named “synanthrophic species” (Pietraszewska 1999). “Synurbisation” is defined as an adaptation to populate core urban areas. Synurbisation requires a wide spectrum of ecological tolerance (eurytopic and polytopic species) and considerable species adaptability. From the ecological perspective, the enrichment of relatively homogeneous urban green areas with new species is favourable, because it makes the ecosystem more complicated and thus more stable and self-sufficient (Gliwicz 1980). The first studies of small mammal fauna in Kraków were carried out already in the 1940s (Kowalski 1950). Nevertheless, the state of knowledge regarding the urban fauna is still insufficient (Karolewski 1981). So far, neither a detailed assessment of changes in the fauna structure caused by urban anthropogenic factors, nor a holistic description of the current state of urban fauna have been made. 255 Table 1. Climate parameters of Kraków Tab. 1. Parametry klimatyczne Krakowa month January February March April May June July August September October November December temperature [°C] precipitation [mm] –2.9 –1.4 2.6 8.6 14.1 17.5 19.3 18.4 14.4 8.8 3.8 –0.2 34 34 35 42 57 86 95 83 56 46 42 34 annual average temperature 8.6 annual precipitation sum 644 This paper is aimed at the description of species composition of small mammal assemblages in natural and anthropogenic biocenoses of the Kraków metropolitan area. The topographic and climatic differentiation of neighbouring areas has a great impact on the urban environment, which makes it an interesting object to study mammal habitat preferences. Despite the fact that synurbisation of fauna is a dynamic and intensifying process, it is rarely investigated. Therefore, the presented data may be helpful in planning similar studies in other areas. MATERIAL AND METHODS Area under study The Kraków agglomeration lies at the place of contact between several physiographic units: Małopolska Uppland in the north, Krakowsko-Wieluńska Uppland in the northwest (from where the Atlantic air masses flow in), Beskid Zachodni Mts. in the south, and Sandomierz Lowlands in the east (where continental climate dominates, Table 1). Boundaries of the area overlap with the boundaries of the former Kraków voivodship that existed between 1975 and 1998. Faunal data sources Information on assemblages of small mammal species in the area comes from a large number of publications listed in Table 2. The Jaccard’s index (Q), describing similarity between assemblages, was calculated according to the formula: Q = (c / a+b–c)×100, where Q = Jaccard’s index, a – number of species in assemblage 1 (here: centre of the Kraków agglomeration), b – number of species in assemblage 2 (here: natural areas and semi-natural suburban areas), c – number of species common for both assemblages. RESULTS 56 species of small mammals occur in Poland, which constitutes 62% of the total number of mammal species (90) of the country. In the study area, 43 small mammal species have been 256 Table 2. List of small mammal species recorded in the former Kraków voivodship. Tab. 2. Lista gatunków Micromammalia zasiedlających województwo krakowskie. A – agglomeration / aglomeracija; V – voivodship / województwo species \ occurrence in Kraków: A V reference Talpa europaea (Linnaeus, 1758) + + Sorex araneus (Linnaeus, 1758) + + Sorex minutus (Linnaeus, 1766) – + Neomys fodiens (Pendant, 1771) – + Neomys anomalus (Cabrera, 1907) – + Crocidura leucodon (Hermann, 1780) – + Crocidura suaveolens (Pallas, 1811) + + Rhinolophus hipposideros (Bechstein, 1800) + + Rhinolophus ferrumequinum (Schreber, 1774) – + Myotis myotis (Borkhausen, 1797) + + Myotis nattereri (Kuhl, 1817) – + Myotis emarginatus (Geoffroy, 1806) + + Myotis mystacinus (Kuhl, 1817) – + Myotis brandtii (Eversmann, 1845) – + Myotis dasycneme (Boie, 1825) – + Myotis daubentonii (Kuhl, 1817) – + Vespertilio murinus (Linnaeus, 1758) + + Eptesicus nilssonii (Keyserling et Blasius, 1839) – + Eptesicus serotinus (Schreber, 1774) – + Pipistrellus pipistrellus (Schreber, 1774) – + Nyctalus noctula (Schreber, 1774) – + Nyctalus leisleri (Kuhl, 1818) – + Plecotus auritus (Linnaeus, 1758) – + Plecotus austriacus (Fischer, 1829) + + Barbastella barbastellus (Schreber, 1774) + + Sciurus vulgaris (Linnaeus, 1758) + + Cricetus cricetus (Linnaeus, 1758) + + Clethrionomys glareolus (Schreber, 1780) + + Arvicola terrestris (Linnaeus, 1758) + + Pitymys subterraneus + + (de Sélys-Longchamps, 1836) Microtus oeconomus (Pallas, 1776) – + Microtus agrestis (Linnaeus, 1761) + + Microtus arvalis (Pallas, 1779) + + Mus musculus (Linnaeus, 1758) + + Rattus norvegicus (Berkenhout, 1769) + + Micromys minutus (Pallas, 1771) – + Apodemus agrarius (Pallas, 1771) + + Apodemus microps (Kratochvíl et Rosický, 1952) – + Apodemus sylvaticus (Linnaeus, 1758) + + Apodemus flavicollis (Melchior, 1834) + + Dryomys nitedula (Pallas, 1779) – + Glis glis (Linnaeus, 1766) – + Muscardinus avellanarius (Linnaeus, 1758) – + Kowalski 1950, Rzebik-Kowalska 1972 Skiba 2005, Haitlinger & Szyszka 1977 Grodziński et al. 1958 Ruprecht 1976 Skuratowicz & Warchalewski 1954 Sałata-Piłacińska 1977 Simm 1952, Ruprecht 1976 Harmata 2000 Nowak et al. 2001 Harmata 1994 Harmata 1960, 1962 Harmata 1969, Harmata & Wojtusiak 1963 Ruprecht 1974 Ruprecht 1974 Biała cave Kowalski 1953 Kowalski 1957 Ojców Kowalski 1953, Harmata 1960 Harmata 1960 Harmata 1962, Markowski & Suskiewicz 1981 Harmata 1960 Harmata 1969 Ruprecht 1971, Harmata 1969 Kowalski et al. 1957 Kowalski 1950, Udziela 1925 Kowalski 1950, Skiba 2005 Skiba 2005, Skuratowicz & Warchalewski 1954 Skiba 2005, Kowalski 1960 Skiba 2005, Haitlinger & Szyszka 1977 Skiba 2005, Haitlinger & Szyszka 1977 Skiba 2005, Haitlinger & Szyszka 1977 Skuratowicz & Warchalewski 1954 Skuratowicz & Warchalewski 1954 Skiba 2005, Haitlinger & Szyszka 1977 Skiba 2005, Haitlinger & Szyszka 1977 Skiba 2005, Haitlinger & Szyszka 1977 Grodziński 1959 Kowalski 1950 Kowalski 1950 257 recorded in the last 50 years or so. Most of them represent 4 families: Soricidae, Vespertilionidae, Arvicolidae, and Muridae. Four of the recorded species are considered synurbic: Sciurus vulgaris, Apodemus agrarius, Mus musculus, and Rattus norvegicus. Among Insectivora, three out of seven species are common for both compared areas, within Rodentia the proportion of common species is ⅔ of the total species number and in Chiroptera only ⅓. In total, 21 out of 43 small mammal species inhabit both subareas: city centre and its surroundings. The Jaccard’s index, reflecting a degree of similarity of small mammal assemblages between (a) core urban and (b) suburban and natural areas, equals 48.84. DISCUSSION Studies on structural changes in small mammal assemblages in urban areas have two practical aspects. Firstly, mammals as animals living in human settlements are a potential source of zoonoses. Secondly, small mammals may be useful as specific bioindicators of anthropogenic environmental pollution. Another interesting problem is species composition of fauna inhabiting buildings and various urban facilities, dominated by such species as Mus musculus, Rattus rattus and Rattus norvegicus (Andrzejewski 1977). The urban fauna is dominated by species of wide geographic ranges. Xerophilous and thermophilous species, mostly those characteristic of warmer climatic zones, occur inside buildings (Markowski 1997). For the majority of city dwelling species communal waste is an abundant food source. Urban areas offer free ecological niches but also impose many threats, e.g. in- Figs. 1, 2. Percentage of small mammal species of the former Kraków voivodship in the whole small mammal fauna of Poland (3, left) and the whole mammal fauna of Poland (4, right). Rys. 1, 2. Liczba gatunków Micromammalia dawnego województwa krakowskiego na tle ogółu krajowej fauny drobnych ssaków (3, nalevo) i na tle ogółu krajowej fauny ssaków (4, napravo), wyrażona w procentach. 258 Table 3. Number of small mammal species in the Kraków centre and in the remaining area of the former Kraków voivodship Tab. 3. Liczba gatunków Micromammalia w centrum Krakowa i na obszarze całego województwa krakowskiego order insectivores rodents bats total / ogółem Kraków agglomeriation Kraków voivodship Σ 3 12 6 21 7 18 18 43 7 18 18 43 creased predation risk (presence of cats, dogs and birds of prey) and many barriers limiting migration (highways, walls, fences). Also a high level of pollution reduces species richness of urban biocenoses (Pisarski & Trojan 1976). The Kraków agglomeration and adjacent areas are situated in the Wisła river valley. Its relatively warm climate is under a strong influence of adjacent physiographic regions. This karstic region is characterised by high faunistic diversity, enriched with several mountain species whose northern distribution limits reach the area. The Wisła river and its tributaries flowing through the Kraków agglomeration contribute to relatively high air humidity and provide feeding grounds and shelters in riparian habitats. The Jaccard’s index (Q) for the city centre and suburbs is 48.84, reflecting high faunal similarity of the two areas, probably caused by the presence of a narrow belt of parkland surrounding the old town. Observations in the Las Wolski, a wood situated on the western periphery of Kraków, have brought interesting information on many mammal species from the following families: Arvicolidae, Muridae, Soricidae, and Gliridae (Glis glis, Muscardinus avellanarius), and a very rare bat species Nyctalus leisleri (Harmata 1960). It is known that synanthropic species show cyclical changes in population. In the years of abundance the range of population is wedge-shaped, narrowing towards the city centre. Currently, Apodemus agrarius is regarded as a synurbic species and it is one of the most abundant species in the Kraków centre. Despite the fact that the small mammal assemblage in the suburbs is relatively rich in species, very often, only Apodemus agrarius is able to populate the core urban area (Gliwicz 1980). In contrast, Arvicolidae species have been observed more and more rarely in the urban agglomeration, which is due to the small area of open habitats inside the city (Kasprzak & Banaszak 1978). One may assume that the Kraków common grounds (Błonia Krakowskie) and banks of the Wisła river are densely populated by Arvicolidae voles, however, no data is available. Studies carried out so far have shown that the number of species connected with forest, shrub and open habitats has been increasing in the urban areas where they enter a mixture of ‘ecotone biocenoses’ (Ziomek 1998). Preference of small mammals towards urban areas depends on season. In spring synanthropic rodent species (Mus musculus, Rattus rattus and Rattus norvegicus) migrate from the city into surrounding areas and in autumn they are forced to come back due to adverse weather conditions and lack of food. Also Microtus arvalis migrates into agrocenoses in spring (Chudoba & Humiński 1963). Contrary to them, some Chiroptera hibernating outside the urban area (in caves or tree hollows) tend to establish summer colonies in human settlements (in attics of old houses and churches, crevices in buildings etc.) (Wołoszyn 1981). 259 STRESZCZENIE Składem gatunkowym drobnych ssaków Krakowa zainteresowano się już w latach czterdziestych ubiegłego stulecia (Kowalski 1950). Brak jak dotąd nie tylko dokładnej oceny zmian w strukturze fauny pod wpływem miejskich czynników antropogenicznych, ale także całościowego obrazu stanu fauny terenów zurbanizowanych (Kasprzak & Banaszak 1978). Celem niniejszej pracy było określenie składu zespołów drobnych ssaków w zbiorowiskach naturalnych, oraz antropogenicznych obszaru metropolitalnego Krakowa. Teren badań wybrano ze względu na zróżnicowanie topograficzno – klimatyczne obszarów z nim graniczących i mających wpływ na tutejsze środowisko i czyni go dobrym do analizowania preferencji siedliskowych gatunków. Proces synurbizacji fauny jest zjawiskiem dynamicznym i nasilającym się, a mimo to relatywnie rzadko badanym. Drobne ssaki w liczbie 56 gatunków stanowią 62,2% spośród 90 gatunków krajowej teriofauny. W ciągu ostatnich około 50 lat na badanym obszarze zanotowano obecność 43 gatunków drobnych ssaków. Wśród nich dominują gatunki, należące do czterech Rodzin: Soricidae, Vespertilionidae, Arvicolidae i Muridae. Cztery spośród zaobserwowanych gatunków, są uważane za synurbijne. Są to Sciurus vulgaris (Linnaeus, 1758), Apodemus agrarius (Pallas, 1771), Mus musculus (Linnaeus, 1758) i Rattus norvegicus (Berkenhout, 1769). Ogółem 21 spośród 43 gatunków drobnych ssaków zasiedlających badany obszar, zasiedla zarówno centrum aglomeracji, jak i tereny otaczające. REFERENCES Andrzejewski R., 1975: Problemy ekologicznego kształtowania środowiska w mieście [Problems of ecological forming of the environment in the urban areas]. Wiadom. Ekol., 21: 175–186 (in Polish). Andrzejewski R., 1977: Stan badań nad ssakami na terenach miast w Polsce [Current research on small mammals of urban areas in Poland]. Wiadom. Ekol., 23: 407–409 (in Polish). Bujalska G., 1970: Reproductin stabilizing elements in an island of C. glareolus (Schreber, 1780). Acta Theriol., 15: 381–412. Chudoba S. & Humiński S., 1963: Owadożerne i gryzonie w osiedlu ludzkim podczas jesieni i zimy [Insectivores and Rodents in inhabited place in autumn and winter]. Zeszyty Naukowe Wyższej Szkoły Rolniczej we Wrocławiu, Zootechnika, 11(52): 11–23 (in Polish, wit a summary in English). Gliwicz J., 1980: Ekologiczny aspekt synurbizacji myszy polnej Apodemus agrarius (Pall.) [The ecological aspect of synurbisation of striped field mouse Apodemus agrarius (Pall.)]. Wiadom. Ekol., 26: 117–124 (in Polish, with a summary in English). Grodziński W., 1958: The succession of small Mamlian communities on the overgrown clearing and landslip in the Western Carpathians. Bull. Acad. Pol. Sci., Cl. II, Ser. Sci. Biol., Warszawa, 6: 431–439. Grodziński W., 1959: Sukcesja zespołów drobnych ssaków na zarastającym zrębie i zsuwie górskim w Beskidzie Średnim (Karpaty Zachodnie) [The succession of small mammalian communities on the overgrown clearing and landslip in the Beskid Średni (Western Carpathians)]. Ekol. Pol. A, 7(4): 83–143 (in Polish, with a summary in English). Haitlinger R. & Szyszka K., 1977: Drobne ssaki Gorców, Beskidu Wyspowego, Pasma Pradziejowej i niektórych obszarów sąsiednich [Small mammals of the Gorce Mts., Beskid Wyspowy Mts., Radziejowa Range and some neighbouring areas]. Przegl. Zool., 21(2): 155–170 (in Polish, with a summary in English). Harmata W., 1960: Obserwacje etologiczne i ekologiczne nad nietoperzami (Chiroptera) z Lasu Wolskiego pod Krakowem [Ethological and ecological observations made on the bats (Chiroptera) of the Wolski forest near Cracow]. Przegl. Zool., 5: 163 – 203 (in Polish, with a summary in English). Harmata W., 1962: Sezonowa rytmika obyczajów i ekologia nietoperzy (Chiroptera) przebywających w niektórych zabytkowych budowlach województwa krakowskiego [Seasonal rhytmicity of behaviour and the ecology of bats (Chiroptera) living in some old buildings in the district of Kraków]. Przegl. Zool., 7: 149–179 (in Polish, with a summary in English). Harmata W., 1969: The thermopreferendum of some species of bats (Chiroptera). Acta Theriol., 14: 49–62. 260 Harmata W., 1994: Winterschlaf des Mausohrs, Myotis myotis (Borkhausen, 1797) in den Festungsanlangen in der Umgebung von Kraków (Südpolen). Folia Zool., 43: 325–330. Harmata W., 2000: Phenological dynamic of hibernating lasser horseshoe bats Rhinolophus hipposideros (Bechst.) (Chiroptera, Rhinolophidae) in a limestone cavern. Studia Chiropterol., 1: 13–28. Harmata W. & Wojtusiak J., 1963: Podkowiec duży (Rhinolophus ferrumequinum, Schreber) nowym ssakiem dla fauny Polski [Greater horseshoe bat Rhinolophus ferrumequinum Schreber (Chiroptera), a species new to the fauna of Mammals in Poland]. Przegl. Zool., 7: 154–157 (in Polish, with a summary in English). Karolewski M. A., 1981: Specyfika i status ekologiczny miasta [Peculiarity and ecological status of a town]. Wiadom. Ekol., 17: 3–35 (in Polish). Kasprzak K. & Banaszak J., 1978: Fauna terenów zurbanizowanych [Fauna of the urban areas]. Wszech świat, 12: 305–307 (in Polish). Kowalski K., 1950: Badania nad ekologią drobnych ssaków leśnych okolocy Krakowa [Ecological research of small mammals in the forests of the Cracow surroundings]. Docum. Physiograph. Polon., 22 (in Polish). Kowalski K., 1953: Materiały do rozmieszczenia i ekologii nietoperzy jaskiniowych w Polsce [Material relating to the distribution and ecology of cave bats in Poland]. Fragm. Faun., 6: 541–567 (in Polish, with a summary in English). Kowalski K., 1957: Microtus nivalis (Martins, 1842) (Rodentia) w Karpatach [Snow vole Microtus nivalis (Martins, 1842) (Rodentia) in the Carpathian Mountains]. Acta Theriol., 1: 159–182 (in Polish, with a summary in English). Kowalski K., 1960: Pitymys Mc. Murtrie 1831 (Microtidae, Rodentia) in the Northern Carpathians. Acta Theriol., 4: 81–91. Kowalski K., Krzanowski A. & Wojtusiak R., 1957: Sprawozdanie z akcji obrączkowania nietoperzy w Polsce w latach 1939–1953 [Report from the ringing action of bats (1939–1953)]. Acta Theriol., 1: 109–158 (in Polish, with a summary i English). Markowski J., 1997: Specyfika synurbijnych populacji zwierząt [Specifics of synurbistic populations of animals]. PWN, Warszawa, 169 pp (in Polish). Markowski J. & Suskiewicz B., 1981: Ssaki (Mammalia) doliny rzeki Pilicy [Mammals of the Pilica River valley]. Acta Univ. Lodz. Folia Zool. Anthropol. (Łódź), 1: 67–94 (in Polish). Nowak J., Kozakiewicz K. & Grzywiński W., 2001: Podkowiec duży Rhinolophus ferrumequinum (Schreber, 1774), nowy gatunek dla fauny Ojcowskiego Parku Narodowego [Greater horseshoe bat Rhinolophus ferrumequinum (Schreber, 1774), a new species for the Ojców National Park fauna]. Studia Chiropterol., 2: 100 (in Polish, with a summary in English). Petrusewicz K., 1978: Osobnik, populacja, gatunek [An individual, population, species]. PWN, Warszawa, 384 pp (in Polish). Pietraszewska T., 1999: Miasto jako układ ekologiczny [Town as an ecological system]. Pp.: 278–310. In: Strzałko J. (ed.): Kompendium wiedzy o ekologii [Compendium of the Ecological Science]. PWN, Warszawa (in Polish). Pisarski B. & Trojan P., 1976: Zoocenozy obszarów zurbanizowanych [Zoocenoses of urban areas]. Wiadom. Ekol., 22: 4 (in Polish). Pucek Z. & Raczyński J., 1983: Atlas of Polish Mammals. PWN, Warszawa, 188 pp (in Polish). Ruprecht A. L., 1971: Distribution of Myotis myotis (Borkhausen, 1797) and representatives of the genus Plecotus Geoffroy, 1818 in Poland. Acta Theriol., 16: 95–104. Ruprecht A. L., 1974: The occurrence of Myotis brandtii (Eversmann, 1845) in Poland. Acta Theriol., 19: 81–90. Ruprecht A. L., 1976: Über die Verbreitung der Rauhhautfledermaus, Pipistrellus nathusii in Polen. Myotis, 14: 25–29. Rzebik-Kowalska B., 1972: Badania nad pokarmem ssaków drapieżnych w Polsce [Research on the diet of Carnivores in Poland]. Acta Zool. Cracov., 17: 415–506 (in Polish). 261 Sałata-Piłacińska B., 1977: Ssaki w pokarmie płomykówki (Tyto alba guattata, Brehm) z terenu Polski, ze szczególnym uwzględnieniem zachodniej części kraju [Mammals in the diet of Tyto alba guattata (Brehm) in Poland with special consideration of the Western Poland]. Bad. Fizjogr. Pol. Zach., Poznań C, 30: 7–27 (in Polish). Simm K., 1952: Zębiełek karliczek (Crocidura mimula, Miller) w Polsce [Lesser white-toothed shrew (Cro cidura mimula, Miller) in Poland]. Chrońmy Przyrodę Ojczystą (Kraków), 6(3–4): 52–53 (in Polish). Skiba J., 2005: Skład gatunkowy drobnych ssaków wybranych obszarów Krakowa na podstawie analizy wypluwek [Small mammal communities of some areas in Kraków, based on the Long-eared owl Asio otus (Linnaeus, 1758) and Tawny owl Strix aluco (Linnaeus, 1758) pellets analysis]. MSc. Thesis. Zakład Ekologii Zwierząt Uniwersytetu Jagiellońskiego, Kraków, 60 pp (in Polish). Skuratowicz W. & Warchalewski E., 1954: Przyczynek do fauny drobnych ssaków Podkarpacia [Contribution to the knowledge of the small mammals fauna in the Carpathian Foothills (Podkarpacie; South-East Poland)]. Poznań. Tow. Przyj. Nauk, Pr. Kom. Biol., 15: 119–130 (in Polish, with a summary in English). Udziela W., 1924: Odmiany geograficzne wiewiórki w Polsce, jej rozsiedlenie oraz znaczenie gospodarcze [Geographical forms of red squirrel Sciurus vulgaris (Linnaeus, 1758)in Poland – occurrence and economic significance]. Rocz. Nauk Rol. Leś. Poznań, 12: 348–373 (in Polish). Wołoszyn B. W., 1981: Nietoperze i cywilizacja [Bats and civilization]. Rocz. Muz. Okręg. Częstochowie, Przyr., 2: 97–108 (in Polish, with a summary in English). Ziomek J., 1998: Drobne ssaki (Micromammalia) Roztocza. Część I. Micromammalia wybranych biotopów Roztocza Środkowego [Small mammals of Roztocze (Eastern Poland), Part I. Micromammals in some biotopes in the Central Roztocze]. Fragm. Faun., 41(8): 93–123 (in Polish, with a summary in English). 262 Lynx (Praha), n. s., 37: 263–273 (2006). ISSN 0024–7774 Population ecology of Apodemus flavicollis in two lowland forest habitats (Rodentia: Muridae) Populačná ekológia Apodemus flavicollis v dvoch typoch nížinného lesa (Rodentia: Muridae) Kristína Trubenová, Dávid Žiak & Peter Miklós Department of Zoology, Faculty of Natural Sciences, Comenius University, Mlynská dolina B-1, SK–842 15 Bratislava, Slovakia; trubenova@fns.uniba.sk, ziak@fns.uniba.sk, miklos@fns.uniba.sk received on 12 June 2006 Abstract. Several population characteristics were studied and compared between populations of Apode mus flavicollis inhabiting two different lowland forest habitats. Small mammals were live-trapped during 2000–2002 on two 1.44 ha plots in alder forest and oak-elm forest (Nature Reserve Jurský Šúr, Slovakia). The density of A. flavicollis was 0.0–25.3 individuals per ha in alder forest and 3.1–28.0 individuals per ha in oak-elm forest. Throughout the study, A. flavicollis was the most numerous species in oak-elm forest (72–100%), while in alder forest its proportion was much lower (0–50%). Higher reproductive activity in alder forest was probably caused by higher food and shelter availability in this area during breeding season. Animals of both sexes were heavier in oak-elm forest which suggests a better fitness of individuals in this habitat. While the degree of male residency was significantly higher in oak-elm forest, we did not find any difference in resident rate of females. Different occurrence and residency of individuals on the study plots are probably caused by higher heterogeneity and larger total area of alder forest together with temporary winter insufficiency of resources on the local study plot. Introduction The structure and dynamics of a population represent the outcome of the interactions between life histories of its individuals and the spatial and temporal variations in its environment (Zeng & Brown 1987). Local populations of widely distributed species living in different environmental conditions may exhibit different demographic parameters in relation to ecological characteristics of habitats (Montgomery & Gurnell 1985). The yellow-necked mouse, Apodemus flavicollis (Melchior, 1834), as one of the most ubiquitous and abundant small mammals in the western Palaearctic, utilises a variety of habitat types throughout its geographical range. It is primarily a woodland rodent preferring mature successional stages of deciduous forests with an open ground layer. However, it can also inhabit field-forest ecotones, grassy areas and brushwood sets (Gurnell 1985, Jensen 1984). Its occurrence in coniferous forests, subalpine and alpine zone is mainly seasonal (Kratochvíl & Rosický 1952, Zima et al. 1984), depending on local seed supply and immigration from nearby stable populations with higher densities (Martínková et al. 2001). Populations of A. flavicollis are able to persist in different habitat types because of their broad flexibility in demography and habitat selection as well as high adaptability to different environmental conditions (Douglass et al. 1992, Montgomery & Gurnell 1985). 263 In the present study we compared several demographic parameters (abundance, residency, age composition breeding activity and body mass) between two populations of A. flavicollis inhabiting different lowland forest habitats in National Nature Reserve Jurský šúr. MATERIAL AND METHODS Study area The study was carried out in the Jurský Šúr National Nature Reserve, 12 km north-east of Bratislava (128–132 m a. s. l., DFS 7769c). Study sites were situated in two different forest habitat types, an alder forest (study plot A) and an oak-elm forest (plot B). The alder forest forms an isolated complex of swampy boggy forest in the centre of wet meadows. It spreads over an area of 334.7 ha and forms the greatest part of the Jurský Šúr National Nature Reserve. The alder forest belongs phytocoenologically to the Carici elongatae-Alnetum medioeuropaeum associati on. The tree layer of study plot A is formed by the following species: Alnus glutinosa, Quercus robur and Ulmus laevis. The shrub layer develops mainly in the sites with loose canopy and is formed by the species Alnus glutinosa, Salix cinerea, Prunus spinosa and Rosa spp. The herb layer is formed by the species Iris pseudacorus, Urtica kiovenensis, Alisma plantago-aquatica and Carex spp. This plot is characterised by the presence of surface floods (XI–II/III) when the water level reaches 20–50 cm (Kupcová 1980). The oak-elm forest is a remnant of old thermophilous mixed forests of the Pannonian basin, partly it keeps its forest-steppe character. It spreads over an area of 42.38 ha. The tree layer of study plot B is represented by the species Quercus robur, Q. cerris, Ulmus laevis, Acer campestre, and Carpinus betulus. Shrub layer is formed by the species Prunus spinosa, Rosa canina and Crataegus spp. The herb layer is formed mainly by grasses (Poales). Besides them, the species Viola sp., Geum urbanum, Alliaria petiolata, Allium ursinum, Lamium maculatum, and Galium aparine occur most frequently (Kupcová1980, Miklos 1999). The study sites were separated by approximately 1 km wide corn field. Methods Small mammals were live-trapped every six weeks between February 2000 and November 2002 on two 1.44 ha plots. Live-traps were set 15 m apart in a 9×9 matrix on each grid. A single Chmela-type live trap baited with oats was placed at each station. Traps were set during six consecutive nights and checked twice a day, mornings and evenings. All captured small mammals were individually marked by toe-clipping and released at trapping points immediately after data collection. Upon capture the following parameters were noted for each individual animal: capture point co-ordinates, species, individual number, sex, weight and status of external sexual characters to assess reproductive individuals (males: scrotal testes; females: enlarged nipples, perforated vagina or pregnancy). Monthly population abundance was estimated by direct enumeration of mice captured on the study plot, and population densities were obtained by dividing abundance by the trapping area, increased by one trap distance per side. Abundance variation was compared between habitats using the chi-square goodness of fit test. All the individuals caught were divided into three groups according to their residential status. The first group (long time residents) comprised animals present throughout at least two series of trapping, the second group (short time residents) included individuals captured several times during the sampling session. The third group (transients) encompassed those individuals that were caught only once and no more. Resident rate was expressed as the proportion of long time residents excluding mice captured in the last trapping period of a year. The age of each individual was estimated on the basis of body weight according to Gliwicz (1988) and all individuals were divided into adult (m>25 g) and young (subadult and juvenile; m<25 g) age classes. The following cha racteristics were analysed for each trapping session, individual years and also jointly for the three years of study: sex ratio, age structure (proportion of young), breeding intensity (proportion of reproductively active females) and resident rate. These characteristics were compared between habitats using chi-square analysis of contingency tables. The body weights of sexually active and adult individuals of both sexes 264 were compared between habitats using the Mann-Whitney test. We tested the spatial distribution of either sex as well as that of both sexes together to find possible departures from random (Poisson) distribution using the χ2-test. When we found non-random spatial distribution, we used Lloyd (1967) index to decide whether the distribution was clumped or even. RESULTS The total material consisted of 1,927 captures of 617 individuals of Apodemus flavicollis (Mel chior, 1834), comprising 914 captures of 301 individuals in alder forest and 1,013 captures of 316 individuals in oak-elm forest. Besides that, the following species were recorded: in the alder forest Clethrionomys glareolus (Schreber, 1780), Sorex araneus Linnaeus, 1758, Neomys fodiens (Pennant, 1771), Micromys minutus (Pallas, 1771) and Mustela nivalis Linnaeus, 1766, and in the oak-elm forest Clethrionomys glareolus, Sorex araneus, Crocidura leucodon (Hermann, 1780), Micromys minutus and Mustela nivalis. The density of A. flavicollis was 0–25.33 individuals per ha on plot A and 3.11–28 individuals per ha on plot B. During the reproduction season (III/IV–IX/X) we did not find any differences in abundance variation between the study sites. Statistically significant difference (χ2=105.8139, p<0.01, df=19) in variation of abundance found, when comparing the whole study period, was caused by much lower minimum numbers and even absence of individuals on plot A during winter months (II–IV 2000, II 2001, III 2002). Throughout the study, the proportion of A. flavicollis in small mammal community was diff erent between the two study sites. In the oak-elm forest, A. flavicollis was the most numerous Fig. 1. Resident rate of males in alder forest (pooled data for 2-months intervals). Obr. 1. Rezidencia samcov v jelšovom lese (zhrnuté dáta pre dvojmesačné obdobia). 265 Fig. 2. Resident rate of males in oak-elm forest (pooled data for 2-months intervals). Obr. 2. Rezidencia samcov v dubovo-brestovom lese (zhrnuté dáta pre dvojmesačné obdobia). species, its proportion in small mammal community reaching 72 to 100%. In the alder forest, the proportion of A. flavicollis ranged from 0 to 50%. Small mammal abundance was always higher in the alder forest (on the average 3.5-times higher, during peak abundance 2.8–4.3 times). We found higher resident rate on plot B when comparing pooled data for the whole study period for all individuals (χ2=8.37, p<0.05) and for males (χ2=13.27, p<0.05) (Figs. 1, 2). Higher resident rate of males on plot B was statistically significant also for pooled data for 2000 (χ2=6.42, p<0.05), 2001 (χ2=4.01, p<0.05), and in 2002 it was close to the limit of statistical significance (χ2=3.73, p=0.054). A similar trend can be observed also in the individual trapping series, but the difference was statistically significant only in IX 2000 (χ2=8.31, p<0.05), VIII 2001 (χ2=4.82, p<0.05) and VIII 2002 (χ2=9.72, p<0.05). We did not find any difference in resident rate of females. We did not find any difference in timing of reproduction (sexually active individuals and juveniles were recorded during the same trapping series on both plots). Reproduction period started in III/IV and ended in IX/X. In the alder forest we found a higher proportion of sexually active females, this difference being statistically significant for the pooled data for the whole study period (χ2=5.7, p<0.05), and for the pooled data for the year 2001 (χ2=8.14, p<0.05) (Fig. 3a, b). In alder forest we found also a higher proportion of young (subadult and juvenile) individu als, this difference being statistically significant for the pooled data for the whole study period (χ2=4.17, p<0.05) (Fig. 4a, b). A higher proportion of young individuals in the alder forest was observed also in individual trapping series, the difference being statistically significant in VIII 2001 (χ2=4.37, p<0.05) and X 2002 (χ2=5.2, p<0.05). 266 Table 1. Mean body weights and standard deviations for sexually active and adult individuals of Apode mus flavicollis Tab. 1. Priemerné hmotnosti pohlavne aktívnych a adultných jedincov Apodemus flavicollis females males alder forest (m±SD) [g] active adult 29.98±3.68 31.96±5.00 28.78±4.68 30.60±6.23 oak-elm forest (m±SD) [g] active adult 31.45±4.21 32.25±4.99 31.79±4.59 33.32±5.95 Average weights of sexually active and adult individuals were higher in the oak-elm forest, this differences being statistically significant only for females (sexually active females: w=5,340, p<0.01, adult females: w=9,433, p<0.01) (Table 1). Spatial distribution of long-time residents on plot A was random in most trapping series, with the exception for females in November 2000 (χ2=6.50, p<0.05, L=1.22) and August 2001 (χ2=6.10, p<0.05, L=2.24) and for both sexes together in October 2001 (χ2=9.63, p<0.01, L=1.68), when the spatial distribution was aggregated. Spatial distribution of males was random in all trapping series. On plot B we found aggregated spatial distribution for males in February (χ2=20.01, p<0.001, L=3.83), April (χ2=18.12, p<0.001, L=4.13), June 2001 (χ2=12.66, p<0.01, L=2.88) and in August 2002 (χ2=6.57, p<0.05, L=2.88). For both sexes together we found aggregated distribution in September (χ2=53.59, p<0.001, L=1.97) and November 2000 (χ2=10.44, Fig. 3a. Proportion of sexually active females in alder forest (pooled data for 2-months intervals). Obr. 3a. Zastúpenie pohlavne aktívnych samíc v jelšovom lese (zhrnuté dáta pre dvojmesačné obdobia). 267 Fig. 3b. Proportion of sexually active females in oak-elm forest (pooled data for 2-months intervals). Obr. 3b. Zastúpenie pohlavne aktívnych samíc v dubovo-brestovom lese (zhrnuté dáta pre dvojmesačné obdobia). Fig. 4a. Proportion of young individuals in alder forest (pooled data for 2-months intervals). Obr. 4a. Zastúpenie mladých jedincov v jelšovom lese (zhrnuté dáta pre dvojmesačné obdobia). 268 p<0.05, L=1.84), February (χ2=6.05, p<0.05, L=2.43), April (χ2=8.05, p<0.05, L=2.45) and July 2001 (χ2=15.22, p<0.01, L=1.79) and February (χ2=11.69, p<0.001, L=3.31) and August 2002 (χ2=9.32, p<0.01, L=2.80). Spatial distribution of females was random in all trapping series. DISCUSSION Populations under study revealed typical seasonal trend of abundance fluctuation known for this species (Flowerdew 1985, Gurnell 1985, Montgomery 1979, Pucek et al. 1993) with peak numbers in late summer/early autumn and decrease during late autumn and winter months. Maximal densities achieved on both plots are consistent with published data for A. flavicollis in Central Europe (Mazurkiewicz & Rajska-Jurgiel 1998, Wojcik & Wolk 1985). The most part of the study plot A was flooded and frozen during winter (XI–II/III), so as regards abundance and availability of food, which is the most important factor governing local distribution and abundance of A. flavicollis (Angelstam et al. 1987, Montgomery 1979), as well as regarding the availability of shelters, it represents an unfavourable environment for this species (Montgomery 1978). The animals retreat from this study plot during winter. However, after a temporary absence during winter we repeatedly captured the same individuals on plot A as in the previous year, and in quite high numbers (Trubenová et al. 2004). The animals inhabiting study plot A are able to survive winter, despite the fact that they are forced to seek out more favourable refugees for overwintering, and probably the refugees inhabited during winter are within easy reach for this species. In contrast to study plot in alder forest, the continuous Fig. 4b. Proportion of young individuals in oak-elm forest (pooled data for 2-months intervals). Obr. 4b. Zastúpenie mladých jedincov v dubovo-brestovom lese (zhrnuté dáta pre dvojmesačné obdobia). 269 occurrence of individuals of A. flavicollis on the study plot in oak-elm forest suggests that this habitat is suitable for A. flavicollis all year over, also during winter months (Trubenová et al. 2004). This seems consistent with the findings of Castien & Gosalbez (1994) and Montgomery (1979), according to whom A. flavicollis is quite rare in many forest types during winter and the populations are restricted to one habitat type characterised by the presence of a number of seed-bearing tree species and by a dense entanglement of branches above ground level. This way could be characterised the study plot in the oak-elm forest as well as the oak-elm forest as a whole. Distinct difference in the proportion of A. flavicollis in the small mammal community between these two habitats is caused by the much higher numbers of small mammals in the alder forest (especially Clethrionomys glareolus), which suggests higher productivity and/or higher hetero geneity of this habitat (M’Closkey 1976, Van Horne 1983). Almost monospecific occupancy of plot B by A. flavicollis is in contrast with the situation in alder forest, where the numbers of A. flavicollis were similar or lower than the numbers of C. glareolus. However, we do not suppose that the presence of C. glareolus limits the numbers achieved by A. flavicollis population. Where these two species occur together, A. flavicollis is generally considered dominant (Andrzejewski & Olszewski 1963, Bergstedt 1965, Grüm & Bujalska 2000), affecting popu lation dynamics, spatial distribution and/or demographic variables of C. glareolus population, but not conversely. The different residency of males and females is caused by different reproductive strategies between sexes. Females are characterised by lower mobility, higher residency and selection of places providing secure cover for rearing the young, while males are distributed with respect to breeding opportunity (Ostfield 1985). As we have found difference only in the resident rate of males, while the resident rate of females did not differ between the plots, we can suppose that higher residency of males in oak-elm forest was not caused only by higher quality of this habitat. In oak-elm forest we have found also longer residency time of individuals of both sexes, significantly for males (Trubenová et al. 2004). Differences in male residency could be associated with more opportunity to disperse in alder forest, and also with higher heterogeneity of this habitat (Trubenová et al. 2004). The alder forest seems to be a highly seasonal environment in which different habitat patches may be preferred at different seasons. Spatial activity and residency of Apodemus spp. are affected not only by the woodland habitat itself but also by the surrounding landscape, distances to neighbouring woodlands and connectivity between them (Kozakiewicz 1993, Ylönen et al. 1991). In smaller woodlands, where the surrounding area of ancient woodland tends to be lower, there is less opportunity to disperse (Marsh & Harris 2000). Brown (1966) concluded that a substantial proportion of male Apodemus have different areas of activity at different times. She reported that the individuals concerned were the dominant males having very large home-ranges and visiting various parts of these ranges on a regular rotation basis. So the lifetime home ranges of A. flavicollis males in oak-elm forest are probably smaller, the animals have fewer seasonal centres of activity, and they stay longer in the same place. Aggregated distribution of males in oak-elm forest can be connected with their higher residency and restricted spatial activity in this habitat. Balanced sex ratio found in both populations (Trubenová et al. 2004) is a characteristic feature of stable populations living in optimal habitats. Although we have found no difference in timing and duration of reproduction period, the higher proportion of sexually active females and young individuals in alder forest indicate a higher intensity of reproduction in this habitat. As the numbers of females active sexually may be a good indicator of the quality of a given 270 habitat (Mazurkiewicz 1991, 1994), the alder forest seems more suitable for A. flavicollis during breeding season. Aggregated spatial distribution of females in alder forest can be connected with locally and temporally more favourable conditions, which cause concentration of females in this habitat during the time it provides suitable conditions for weaning the litter. Higher numbers of sexually active females in alder forest could have also another reason. In oak-elm forest the individuals remain all year over, as they find suitable conditions for surviving the winter there. So the structure of the local population is stable, with well-defined dominance hierarchy. Social interactions may prevent young animals from entering the reproducing part of population (Mazurkiewicz & Rajska-Jurgiel 1998). In contrast, the study plot in alder forest is not populated by A. flavicollis individuals during winter and the population in spring consists mainly of immigrants. In such environment, where there is no stable structure of the population and where at that time there is no social interaction factor, every spring and early-summer immigrant may mature and breed (Mazurkiewicz & Rajska-Jurgiel 1998, Van Horne 1983). The higher body weight of individuals in the oak-elm forest found, when comparing all in dividuals (Trubenová et al. 2004), can be partly caused by the higher proportion of adults in this habitat. However, higher body weights were found in this habitat also when comparing sexually active individuals (although significantly only for females), and also when comparing adult individuals (significantly for females), which suggest that individuals living in the oak-elm forest are heavier. It is used to present body weight as one of the measures of fitness, which provides information about habitat quality for the studied species, as animals in higher-quality habitats tend to be heavier (Van Horne 1983). Higher body weight of animals of both sexes in oak-elm forest suggests better fitness of individuals in this habitat, which is probably caused by more suitable year-long conditions here in comparison to the alder forest, where the fitness of individuals may be lowered by the cost of winter migration. SÚHRN V práci sú vyhodnotené a porovnané demografické parametre populácií Apodemus flavicollis, obývajú cich dva typy nížinného lesa. Terénny výskum prebiehal v rokoch 2000–2002 v jelšovom lese a dubovo-brestovom lese (NPR Šúr, SR) na odchytových kvadrátoch veľkosti 1,44 ha s použitím CMR metódy. Počas obdobia výskumu bolo v jelšovom lese zaznamenaných 914 odchytov 301 jedincov a v dubovo-brestovom lese 1013 odchytov 316 jedincov A. flavicollis. Denzita druhu dosahovala hodnoty 0,0–25,3 jedincov na hektár v jelšovom lese a 3,1–28,0 jedincov na hektár v dubovo-brestovom lese. Zatiaľ čo v dubovo-brestovom lese bol A. flavicollis najpočetnejším druhom počas celého obdobia (72–100 %), v jelšovom lese bolo jeho zastúpenie výrazne nižšie (0–50 %). Obdobie rozmnožovania, stanovené na základe odchytávania pohlavne aktívnych a juvenilných jedincov trvalo v jednotlivých rokoch v obidvoch prostrediach rovnako. Vyšší podiel pohlavne aktívnych samíc, ako aj juvenilných jedincov v jelšovom lese poukazuje na vyššiu reprodukčnú aktivitu v tomto prostredí. V dubovo-brestovom lese bol zistený vyšší podiel dlhodobo rezidentných samcov, rezidencia samíc sa medzi sledovanými prostrediami nelíšila. U oboch pohlaví bola zistená vyššia hmotnosť jedincov v dubovo-brestovom lese, čo naznačuje lepšiu kondíciu jedincov v tomto prostredí. Rozdiely v rezidencii jedincov medzi sledovanými lokalitami boli pravdepodobne spôsobené väčšou rozlohou a heterogenitou jelšového lesa, súčasne s jeho nevhodnými podmienkami počas zimného obdobia. ACKNOWLEDGEMENTS This work was carried out with the financial support of the grants by VEGA No. 1/7197/20 and 1/0017/03. We also thank Lenka Trebatická for her help with the field-work. 271 REFERENCES Andrzejewski R & Olszewski J., 1963: Social behaviour and interspecific relations in Apodemus flavicollis (Melchior, 1834) and Clethrionomys glareolus (Schreber, 1780). Acta Theriol., 7: 155–168. Angelstam P., Hansson L. & Pehrsson S., 1987: Distribution borders of field mice Apodemus: the importance of seed abundance and landscape composition. Oikos, 50: 123–130. Bergstedt B., 1965: Distribution, reproduction, growth and dynamics of the rodent species Clethrionomys glareolus (Schreber), Apodemus flavicollis (Melchior) and Apodemus sylvaticus (Linné) in southern Sweden. Oikos, 16: 132–160. Brown L. E., 1966: Home range and movements of small mammals. Symp. Zool. Soc. Lond., 18: 111–142. Castien E. & Gosalbez J., 1994: Habitat selection of A. flavicollis in a Fagus sylvatica forest in the Western Pyrenees. Folia Zool., 43: 219–224. Douglas R. J., Douglass K. S. & Rossi L., 1992: Ecological distribution of bank voles and wood mice in disturbed habitats: preliminary results. Acta Theriol., 37: 359–370. Flowerdew J. R., 1985: The population dynamics of wood mice and yellow-necked mice. Symp. Zool. Soc. Lond., 55: 315–338. Gliwicz J., 1988: Seasonal dispersal in non-cyclic populations of Clethrionomys glareolus and Apodemus flavicollis. Acta Theriol., 33: 263–272. Grüm L. & Bujalska G., 2000: Bank voles and yellow necked mice: what are interrelations between them? Pol. J. Ecol. 48, Suppl.: 141–145. Gurnell J., 1985: Woodland rodent communities. Symp. Zool. Soc. Lond., 55: 377–411. Jensen T. S., 1984: Habitat distribution, home range and movements of rodents in mature forest and reforestations. Acta Zool. Fenn., 171: 305–307. Kozakiewicz M., 1993: Habitat isolation and ecological barriers – the effect on small mammal populations and communities. Acta Theriol., 38: 1–30. Kratochvíl J. & Rosický B., 1952: K bionomii a taxonomii myší rodu Apodemus žijících v Československu [Zur Bionomie und Taxonomie in der Tschechoslowakei lebenden Apodemus-Arten]. Zool. Entomol. Listy, 1: 57–71 (in Czech, with a summary in German). Kupcová A., 1980: Avifauna Jurského šúru [Bird fauna of the Jurský šúr]. Ochr. Prír., 1: 71–103 (in Slovak). Lloyd M., 1967: Mean crowding. J. Anim. Ecol., 36: 1–30. M’Closkey R. T., 1976: Community structure in sympatric rodents. Ecology, 57: 728–739. Marsh A. C. W. & Harris S., 2000: Partitioning of woodland habitat resources by two sympatric species of Apodemus: lessons for the conservation of the yellow-necked mouse (A. flavicollis) in Britain. Biol. Cons., 92: 275–283. Martínková N., Žiak D. & Kocian Ľ., 2001: Response of Apodemus flavicollis to conditions at the altitude limit in the Western Tatra Mountains. Mammal. Biol., 66: 185–189. Mazurkiewicz M., 1991: Population dynamics and demography of the bank vole in different tree stands. Acta Theriol., 36: 207–227. Mazurkiewicz M., 1994: Factors influencing the distribution of the bank vole in forest habitats. Acta Theriol., 39: 113–126. Mazurkiewicz M. & Rajska-Jurgiel E., 1998: Spatial behaviour and population dynamics of woodland rodents. Acta Theriol., 43: 137–161. Miklós P., 1999: Selekcia prostredia tromi druhmi drobných zemných cicavcov v podmienkach Panónske ho hája [Microhabitat Selection by Three Small Mammal Species in an Oak-elm Forest]. Unpublished M.Sc. Thesis. Department of Zoology, Comenius University, Bratislava, 48 pp (in Slovak). Montgomery W. I. & Gurnell J., 1985: The behaviour of Apodemus. Symp. Zool. Soc. Lond., 55: 89–115. Montgomery W. I., 1978: Studies on the distributions of Apodemus sylvaticus (L.) and A. flavicollis (Melchior) in Britain. Mammal Rev., 8: 177–184. 272 Montgomery W. I., 1979: Seasonal variation in numbers of Apodemus sylvaticus, A. flavicollis and Cle thrionomys glareolus. J. Zool., Lond., 188: 183–186. Ostfield R. S., 1985: Limiting resources and territoriality in microtine rodents. Am. Natur., 126: 1–15. Pucek Z., Jedrzejewski W., Jedrzejewska B. & Pucek M., 1993: Rodent population dynamics in a primeval deciduous forest (Bialowieza National Park) in relation to weather, seed crop, and predation. Acta Theriol., 38: 199–232. Trubenová K., Žiak D. & Miklós P., 2004: Niektoré populačné charakteristiky Apodemus flavicollis (Melchior, 1834) v dvoch lesných biotopoch [Characteristics of Apodemus flavicollis populations inha biting two forest habitats]. Pp.: 59–66. In: Adamec M. & Urban P. (ed.): Výskum a ochrana cicavcov na Slovensku VI. Zborník referátov z konferencie (Zvolen 10.–11. 10. 2003) [Research and Protection of Mammals in Slovakia VI. Proceedings of the Conference (Zvolen 10–11 October 2003)]. Štátna ochrana prírody SR, Banská Bystrica, 189 pp (in Slovak, with an abstract in English). Van Horne B., 1983: Density as a misleading indicator of habitat quality. J. Wildl. Manag., 47: 893–901. Wojcik J. M. & Wolk K., 1985: The daily activity rhythm of two competitive rodents: Clethrionomys glareolus and Apodemus flavicollis. Acta Theriol., 30: 241–258. Ylönen H., Altner H. J. & Stubbe M., 1991: Seasonal dynamics of small mammals in an isolated woodlot and its agricultural surroundings. Ann. Zool. Fenn., 28: 7–14. Zeng Z. & Brown J. H., 1987: Population ecology of a desert rodent: Dipodomys meriami in the Chihua huan desert. Ecology, 68: 1328–1340. Zima J., Hrabě V., Štěrba O. & Schlamp M., 1984: Drobní savci kotliny Siedmych prameňov v Belianskych Tatrách. Zborn. Prác Tatran. Nár. Parku, 25: 29–46. 273 Lynx (Praha), n. s., 37: 275–284 (2006). ISSN 0024–7774 Karyotype analysis of Murina suilla and Phoniscus atrox from Malaysia (Chiroptera: Murininae, Kerivoulinae) Karyotypová analysa dvou netopýrů z Malajsie: trubkonosa vepřího (Murina suilla) a vlnouška vroubkozubého (Phoniscus atrox) (Chiroptera: Murininae, Kerivoulinae) Marianne Volleth Department of Human Genetics, Leipziger Str. 44, Otto-von-Guericke University, D–39120 Magdeburg, Germany; Marianne.Volleth@Medizin.Uni-Magdeburg.de received on 26 May 2006 Abstract. For the first time, chromosomal data are presented for Murina suilla (2n=44, FN=58) and Phoniscus atrox (2n=40). The karyotype of Murina is distinguished from that of Myotis by the presence of euchromatic short arms on chromosomes 12, 13 and 15. The chromosomal complement of Phoniscus atrox is characterized by extensive addition of heterochromatic short arms and a paracentric inversion in chromosome 8. The reduction in the diploid chromosome number is due to Robertsonian fusion of chro mosomal arms 8+13 and 11+14, respectively. Introduction In recent years, progress in molecular genetic methods yielded fascinating new insights into bat phylogeny (for references see Eick et al. 2005, Teeling et al. 2005) and let, together with mor phological and call frequency differences, to the discovery of cryptic species (e.g. Pipistrellus pygmaeus, Barrat et al. 1997, Mayer & von Helversen 2001; Plecotus macrobullaris, Kiefer & Veith 2001, Kiefer et al. 2002; Myotis alcathoe von Helversen et al. 2001, etc.). The members of the vespertilionid subfamilies Murininae and Kerivoulinae, however, gained less interest than most other vespertilionid genera. This is not only true for molecular genetic investigations but also for cytogenetic studies. Up to now, out of 15 recognized species of Murina Gray, 1842 (13 in Nowak 1994 plus two newly described species: Maeda & Matsumura 1998, Csorba & Bates 2005), chromosomal and fundamental numbers have been reported for six species only. Banded karyotypes have been studied only in Murina ussuriensis Ognev, 1913 and M. hilgendorfi Peters, 1880 (Harada et al. 1987). Even less is known from the subfamily Kerivoulinae. Out of the 23 species recognized (Nowak 1994, Vanitharani et al. 2003, Bates et al. 2004), conventionally stained karyotypes have been presented only from Kerivoula lanosa (Smith, 1847) (Rautenbach et al. 1993) and K. papillosa (Temminck, 1840) (McBee et al. 1986). In this paper we present detailed karyological information of Murina suilla and Phoniscus atrox from Malaysia together with a literature overview. 275 Material and methods Specimens examined Murina suilla, 1 male, 1 female, Templer Park near Kuala Lumpur, SMF 69348, 69349, 1 male, Ulu Gombak Field Studies Centre, Selangor, Malaysia, SMF 69350. Phoniscus atrox, 1 female, Ulu Gombak Field Studies Centre, Selangor, Malaysia, SMF 69316. (SMF = catalogue number of the Senckenberg Museum, Frankfurt am Main, Germany) Chromosome preparations and analysis Cell culture and cytogenetic procedures were done as described in Volleth (1987) and Volleth et al. (2001). Chromosomal arms were numbered according to Bickham’s scheme for American Myotis species (Bickham 1979). Karyotypes were compared with the basic karyotype of Vespertilionidae (Volleth & Heller 1994) Results Murina suilla (Temminck, 1840) The karyotype of Murina suilla shows a diploid chromosome number (2n) of 44 and a funda mental number of autosomal arms (FN) of 58. The G-banded karyotype of a male specimen is shown in Fig. 1. Chromosomes 7, 12, 13 and 15 show a small G-negative short arm. C-banding revealed that these short arms consist of euchromatic material (Fig. 2). Therefore these short arms arose very likely by pericentric inversions from the acrocentric homologues present in the basic karyotype of Vespertilionidae (Volleth & Heller 1994). AgNOR-staining detected active Nucleolus Organizing Regions (NORs) in the distal part of the euchromatic short arms of chro mosomes 7, 12 and 15 and proximal to the centromere in minute short arms of chromosomes 8, 9, 14, 18–20, 22 and 24 or 25. These minute short arms are visible only in silver-stained preparations. The differences in the distribution of active NORs in two specimens are shown in Fig. 3 (35 analyzed cells from specimen 69350 and 20 from specimen 69349). In many cells only one homologue of a chromosomal pair was shown to bear an active NOR. The X is a submetacentric chromosome with a heterochromatic segment in the long arm adjacent to the centromere (see Fig. 2). The subtelocentric Y-chromosome has approximately the same size as chromosome 18 and consists largely of heterochromatic material. Phoniscus atrox Miller, 1905 The karyotype of P. atrox consists of 40 chromosomes and shows a total number of autoso mal arms of 76. However, quite a large number of autosomal arms consist of heterochromatin and therefore the fundamental number is presumed to be approximately 50 only. Fig. 4 shows a comparison of G-banded and C-banded chromosomes of the female studied. The reduction in the diploid chromosome number from the 2n=44 vespertilionid basic kary otype to a 2n=40 in Phoniscus atrox is due to two Robertsonian fusion events, i.e. fusion of chromosomal arms 8+13 and 11+14, respectively. In addition, the karyotype is characterized by a paracentric inversion in arm 8. In the P. atrox karyotype, extensive heterochromatin addition was found: first, thin interstitial heterochromatic bands are present in arm 2 (two bands) and in arm 6; second, heterochromatin 276 Fig. 1. G-banded karyotype of Murina suilla. Obr. 1. Karyotyp trubkonosa vepřího (Murina suilla) barvený G-pruhováním. 277 Fig. 2. C-banded metaphase spread of a Murina suilla male. The arrows point to the euchromatic short arms of chromosomes 7, 12, 13, and 15. Obr. 2. Rozptyl C-pruhováním barvené metafase samce Murina suilla. Šipky ukazují euchromatická krátká ramena chromosomů 7, 12, 13 a 15. Fig. 3. NOR distribution in two specimens of Murina suilla (SMF catalogue number in parentheses). Chromosomal arms of metacentric chromosomes are marked by x. Obr. 3. Distribuce oblastí organisujících jadérko (NOR) u dvou jediců Murina suilla (v závorkách jsou sbírková čísla exemplářů Senckenbergského musea, Frankfurt nad Mohanem, Německo). Chromosomální ramena metacentrických chromosomů jsou označena x. 278 is found in the short arms of chromosomes 7, 9, 10, 12, 15, 18–25. In the specimen studied, the size of these heterochromatic arms differs between the homologues of several pairs. This is especially evident for chromosomes 9 and 18 (Fig. 4). In the heterochromatin blocks, G-positive segments are followed by G-negative bands. According to the staining properties, three different entities can be distinghuished: C-positive, CMA brightly fluorescing, G-negative bands are found only adjacent to centromeres of chromosomes 9, 10, 12, 18, 19, 22 and 24. Second, G-positive, C-positive, and DAPI-positive heterochromatin is found in the proximal parts of the short arms of chromosomes 10, 15, 18, 22 and 24. Applying Distamycin A/DAPI double staining, these regions show a weak but clearly positive fluorescence, rarely found in other bat species. The third class is C-negative non-euchromatin, staining either G-positive or G-negative. It is found in the distal parts of the short arms of chromosome 7, 10, 12, 19–21, 23–25. Silver staining revealed active NORs in the short arms of chromosomes 7, 12, 18, 20 and 23. In the case of chromosomes 12, 18 and 23, however, only one homologue each showed an active NOR. These NOR-bearing chromosomes were clearly distinguishable from the respective homologues by the presence of a secondary constriction in the short arm. The X chromosome is a medium-sized submetacentric with a banding pattern similar to that of Myotis Kaup, 1829. Karyotype comparison of vespertilionid genera have revealed that some of the chromosomal arms exist in two states (I, II) differing by inversions (Volleth & Heller 1994). In Phoniscus, chromosomal arms 11, 13, 15 and the X, all show state I, as in the genus Myotis. The state of chromosome 7 is difficult to determine. The short arm is clearly different from that of Myotis due to the terminal addition of two G-positive bands. According to the banding pattern of the long arm, however, chromosome 7 of Phoniscus Miller, 1905 evolved very likely from state I. Discussion As Myotis, the members of the subfamily Kerivoulinae show the highest tooth number found in Vespertilionidae, i.e. 38. Apart from this primitive character, a wide range of morphological features characterizes this subfamily (see Miller 1907, Koopman 1994). There are 23 recognized species in two genera, Kerivoula Gray, 1842 and Phoniscus. According to Miller (1907), “the greatly increased size and peculiar shape of the upper canine and the four-cusped inner mandibular incisors distinguish this genus (Phoniscus) sufficiently from Kerivoula”. Together with Hill (1965) and Corbet & Hill (1992) we follow Miller (1907) in treating Phoniscus as a separate genus and not as a subgenus of Kerivoula as Koopman (1994). The subfamily Murininae, comprising three genera, Murina, Harpiola Thomas, 1915 (Bhat tacharyya 2002, Kuo et al. 2006) and Harpiocephalus Gray, 1842, is characterized by tubular nostrils. It has been regarded by Miller (1907) “as a specialized offshoot from some low, Myotis-like Vespertilioninae form”. Up to now, only few molecular genetic studies have dealt with phylogenetic relationships of Murininae and Kerivoulinae. Consistently all studies revealed a sister relationship between Myotis and a clade containing Murininae and Kerivoulinae (Van Den Bussche & Hoofer 2001, Hoofer & Van Den Bussche 2003, Stadelman et al. 2004, Kawai et al. 2002). As a result, Hoofer & Van Den Bussche (2003) proposed subfamiliar rank for Myotini. Our cytogenetic results revealed one common feature for Myotis, Murina and probably also Phoniscus, i.e. an euchromatic short arm on chromosome 7 (see Volleth & Heller 1994). From karyological reasons, it cannot be decided at the moment whether this character is ancestral or derived. Remarkably, in the Australian genus Vespadelus Troughton, 1943, belonging to the 279 280 Fig. 4. Karyotype of Phoniscus atrox after G-banding (upper rows) and C-banding (lower rows). Note size polymorphism of heterochromatic short arms. Obr. 4. Karyotyp vlnouška vroubkozubého (Phoniscus atrox) po G-pruhovém barvení (horní řady) a C-pruhovém barvení (dolní řady). Nápadný je polymorfismus velikosti heterochromatických krátkých ramen. Vespertilionini, euchromatic short arms are also present on chromosomes 7 and 13 (Volleth & Tidemann 1989). As in Myotis and Murina, these short arms occurred probably by pericentric inversions. In the case of Vespadelus, however, these characters are clearly derived, as closely related genera lack euchromatic short arms. In the case of Murina and Vespadelus, we are clearly faced with a re-use of identical or similar inversion break-points on two chromosomes, 7 and 13. It could be suspected that these break-points have occurred within relatively short fragile regions, being hot-spots of rearrangements. Recently, this model has been proposed to explain break-point clustering in chromosome evolution by Pevzner & Tesler (2003). The authors concluded that “mammalian genomes are mosaics of fragile regions with high propensity for rearrangements and solid regions with low propensity for rearrangements”. This break-point clustering in fragile regions reveals limitations of the widely accepted random breakage theory. Comparison with published karyological data From the subfamily Kerivoulinae, only two out of 23 species have been studied karyologically up to now. The examined female specimen of Kerivoula papillosa showed a diplod chromoso me number of 38 and a fundamental number (including the X chromosomes) of 52 (McBee et al. 1986). The karyotype of the second species studied, K. lanosa, consists of 28 chromosomes Table 1. Chromosomal data for Murina. Abbreviations: M – metacentric; SM – submetacentric; ST – subtelocentric; A – acrocentric; conv – conventional staining; G – G-banding; C – C-banding; AgNOR – silver-staining of NORs Tab. 1. Chromosomální údaje o rodu Murina. Zkratky: M – metacentrický; SM – submetacentrický; ST – subtelocentrický; A – akrocentrický; conv – konvenční barvení; G – G-pruhování; C – C-pruhování; AgNOR – barvení oblastí organisujících jadérko (NOR) stříbrem species 2n aurata1 cyclotis hilgendorfi2 hilgendorfi leucogaster leucogaster puta silvatica suilla ussuriensis 44 60 44 50? 44 44 56 44 58 44 50 44 50 44 56 44 58 44 56 FN M-SM ST 5 4 4 4 4 4* 4 3 4 4 4 – – 3 4 – – 4 4 3 A X Y method source 12 17 17 14 13 17 17 14 13 14 SM SM SM SM SM SM M M SM SM A – – A A A A A ST A conv conv conv G conv conv conv conv, AgNOR G, C, AgNOR G Ando et al. 1977 Rickart et al. 1999 Harada 1973 Harada et al. 1987 Ando et al. 1977 McBee et al. 1986 Lin et al. 2002 Ono & Obara 1994 this study Harada et al 1987 * chromosomes showing different morphology compared to the other species. 1 these Japanese specimens would probably be classified with M. ussuriensis today. 2 the subspecies leucogaster hilgendorfi has been elevated to species rank by Yoshiyuki (1989). 281 with a FN of 50 (Rautenbach et al. 1993). The Phoniscus atrox female presented here is the first Kerivoulinae species where banding methods have been applied. Comparing the three species studied, it becomes clear that the karyological variability is much greater in the subfamily Ke rivoulinae than in the Murininae and Myotinae. Chromosomal data from other Kerivoulinae species are therefore urgently needed for a karyological characterization of this subfamily. More chromosomal data are available from the subfamily Murininae. Conventionally stained karyotypes of 6 Murina species have been published (references see Table 1), all with a diploid chromosome number of 44. Concerning the number of subtelocentric chromosomal pairs, the data differ, in some cases even within the same species. To my opinion, this is more likely a matter of interpretation than of real differences, as the minute short arms are clearly visible only in high quality metaphases. On the other hand, chromosomal differences could probably point to hidden species. One example could be the M. leucogaster Milne-Edwards, 1872 specimen of McBee et al. (1986), showing a clearly different morphology of the metacentric pairs. An indication for cryptic species in the genus Murina was also revealed by cytochrome b sequence comparison of Murina cf. cyclotis Dobson, 1872 specimens from Laos (Stadelman et al. 2004). From the two other Murininae genera, Harpiola and Harpiocephalus, chromosome description has only be given for a single female specimen of Harpiocephalus mordax Thomas, 1923 with a 2n=40 (McBee et al. 1986). According to the 5 subtelocentric chromosomal pairs observed in this specimen, it could be suspected that there are also some pairs with euchromatic short arms as in Murina. From the genus Murina, G-banding patterns of two species, Murina ussuriensis and Murina hilgendorfi, have been published previously (Harada et al. 1987). The results are in good agreement with the data published here. Only the euchromatic short arm on chromosome 7, present in Murina and the genus Myotis, has not been mentioned by Harada et al. (1987). The distributional pattern of Nucleolus Organizing Regions has only been studied in M. sil vatica (Ono & Obara 1994). As in the genus Myotis and in M. suilla, multiple NORs on minute short arms of acrocentric chromosomes have been found in this species. As multiple NORs are also present in Phoniscus atrox, the distributional pattern of NORs is very similar in Myotinae, Kerivoulinae and Murininae. However, the ancestral location of the NORs for Vespertilionidae is not yet ascertained and therefore this feature is not suited for judging phylogenetic relationships amongst these subfamilies. SOUHRN Poprvé jsou uvedeny popisy chromosomů u trubkonosa vepřího (Murina suilla) (2n=44, FN=58) a vlnouška vroubkozubého (Phoniscus atrox) (2n=40). Karyotyp rodu Murina se liší od karyotypu rodu Myotis přítomností euchromatických krátkých ramen na chromosomech 12, 13 a 15. Chromosomová sada druhu Phoniscus atrox je typická extensivním přídavkem heterochromatických krátkých ramen a paracentrickou inversí na chromosomu 8. Redukce diploidního počtu chromosomů je způsobena Robertsonskými fusemi chromosomálních ramen 8+13 a 11+14. References Ando K., Tagawa T. & Uchida T. A., 1977: Considerations of karyotypic evolution within Vespertilioni dae. Experientia, 33: 877–879. Barratt E. M., Deaville R., Burland T. M. & Bruford M. W., 1997: DNA answers the call of pipistrelle bat species. Nature, 387: 138–139. 282 Bates P. J. J., Struebig M. J., Rossiter S. J., Kingston T., Sai S. L. O., Khin M. M., 2004: A new species of Kerivoula (Chiroptera: Vespertilionidae) from Myanmar (Burma). Acta Chiropterol., 6: 219–226. Bhattacharyya T. P., 2002: Taxonomic status of the genus Harpiola Thomas, 1915 (Mammalia: Chirop tera: Vespertilionidae), with a report of the occurrence of Harpiola grisea (Peters, 1872) in Mizoram, India. Proc. Zool. Soc. Calcutta, 55: 73–76. Bickham J. W., 1979: Banded karyotypes of 11 species of American bats (genus Myotis). Cytologia, 44: 789–797. Corbet G. B. & Hill J. E., 1992: The Mammals of the Indomalayan Region: a Systematic Review. Oxford University Press, 488 pp. Csorba G. & Bates P. J. J., 2005: Description of a new species of Murina from Cambodia (Chiroptera: Vespertilionidae: Murininae). Acta Chiropterol., 7: 1–7. Eick G. N., Jacobs D. S. & Matthee C. A., 2005: A nuclear DNA phylogenetic perspective on the evolution of echolocation and historical biogeography of extant bats (Chiroptera). Mol. Biol. Evol., 22: 1869–1886. Harada M., 1973: Chromosomes of nine chiropteran species in Japan (Chiroptera). La Kromosomo, 91: 2885–2895. Harada M., Ando K., Uchida T. A. & Takada S., 1987: A karyological study on two Japanese species of Murina (Mammalia: Chiroptera). J. Mammal. Soc. Japan, 12: 15–23. von Helversen O., Heller K.-G., Mayer F., Nemeth A., Volleth M. & Gombkötő P., 2001: Cryptic mammalian species: a new species of whiskered bat (Myotis alcathoe n. sp.) in Europe. Naturwissenschaften, 88: 217–223. Hill J. E., 1965: Asiatic bats of the genera Kerivoula and Phoniscus (Vespertilionidae). Mammalia, 29: 524–556. Hoofer S. R. & Van Den Bussche R. A., 2003: Molecular phylogenetics of the chiropteran family Vesper tilionidae. Acta Chiropterol., 5 (Supplement): 1–63. Kawai K., Nikaido M., Harada M., Matsumura S., Lin L.-K., Wu Y., Hasegawa M. & Okada N., 2002: Intra- and interfamily relationships of Vespertilionidae inferred from various markers including SINE insertion data. J. Mol. Evol., 55: 284–301. Kiefer A. & Veith M., 2001: A new species of long-eared bat from Europe (Chiroptera: Vespertilionidae). Myotis, 39: 5–16. Kiefer A., Mayer F., Kosuch J., von Helversen O. & Veith M., 2002: Conflicting molecular phylogenies of European long-eared bats (Plecotus) can be explained by cryptic diversity. Mol. Phylogenet. Evol., 25: 557–566. Koopman K. F., 1994: Chiroptera: Systematics. Pp.: 1–217. In: Niethammer J., Schliemann H. & Starck D. (eds.): Handbuch der Zoologie. Band VIII. Mammalia. Teilband 60. Walter de Gruyter, Berlin & New York, vii+224 pp. Kuo H. C., Fang Y. P., Csorba G. & Lee L. L., 2006: The definition of Harpiola (Vespertilionidae: Muri ninae) and the description of a new species from Taiwan. Acta Chiropterol., 8: 11–19. Lin L.-K., Motokawa M. & Harada M., 2002: Karyology of ten vespertilionid bats (Chiroptera: Vesper tilionidae) from Taiwan. Zool. Stud., 41: 347–354. Maeda K. & Matsumura S., 1998: Two new species of vespertilionid bats, Myotis and Murina (Vespertilioni dae: Chiroptera) from Yanbaru, Okinawa Island, Okinawa Prefecture, Japan. Zool. Sci., 15: 301–307. Mayer F. & von Helversen O., 2001: Sympatric distribution of two cryptic bat species across Europe. Biol. J. Linn. Soc., 74: 365–374. McBee K., Bickham J. W., Yenbutra S., Nabhitabhata J. & Schlitter D. A., 1986: Standard karyology of nine species of vespertilionid bats (Chiroptera: Vespertilionidae) from Thailand. Ann. Carnegie Mus., 55: 95–116. Miller G. S., 1907: The families and genera of bats. Bull. U. S. Natl. Mus., 57: 1–282. Nowak R. M., 1994: Walker’s Bats of the World. The Johns Hopkins University Press, Baltimore, Mary land, USA, 287 pp. 283 Ono T. & Obara Y., 1994: Karyotypes and Ag-NOR variations in Japanese vespertilionid bats (Mammalia: Chiroptera). Zool. Sci., 11: 473–484. Pevzner P. & Tesler G., 2003: Human and mouse genomic sequences reveal extensive breakpoint reuse in mammalian evolution. Proc. Natl. Acad. Sci., 100: 7672–7677. Rautenbach I. L., Bronner G. N. & Schlitter D. A., 1993: Karyotypic data and attendant systematic implications for the bats of southern Africa. Koedoe, 36: 87–104. Rickart E. A., Mercier J. A. & Heaney L. R., 1999: Cytogeography of Philippine bats (Mammalia: Chi roptera). Proc. Biol. Soc. Washington, 112: 453–469. Stadelmann B., Jacobs D., Schoeman C. & Ruedi M., 2004: Phylogeny of African Myotis bats (Chiroptera, Vespertilionidae) inferred from cytochrome b sequences. Acta Chiropterol., 6: 177–192. Teeling E. C., Springer M. S., Madsen O., Bates P., O’Brien S. J. & Murphy W. J., 2005: A molecular phylogeny for bats illuminates biogeography and the fossil record. Science, 307(5709): 580–584. Van Den Bussche R. A. & Hoofer S. R., 2001: Evaluating monophyly of Nataloidea (Chiroptera) with mitochondrial DNA sequences. J. Mammal., 82: 320–327. Vanitharani J., Rajendran A., Bates P. J. J., Harrison D. L. & Pearch M. J., 2003: A taxonomic reasse ssment of Kerivoula lenis Thomas, 1916 (Chiroptera, Vespertilionidae) including a first record from peninsular India. Acta Chiropterol., 5: 49–60. Volleth M., 1987: Differences in the location of nucleolus organizer regions in European vespertilionid bats. Cytogenet. Cell Genet., 44: 186–197. Volleth M. & Heller K. G., 1994: Phylogenetic relationships of vespertilionid genera (Mammalia: Chi roptera) as revealed by karyological analysis. Ztschr. Zool. System. Evolutionsforsch., 32: 11–34. Volleth M. & Tidemann C. R., 1989: Chromosome studies in three genera of Australian vespertilionid bats and their systematic implications. Ztschr. Säugetierk., 54: 215–222. Volleth M., Bronner G., Göpfert M. C., Heller K.-G., von Helversen & Yong H. S., 2001: Karyotype comparison and phylogenetic relationships of Pipistrellus-like bats (Vespertilionidae; Chiroptera; Mam malia). Chromosome Res., 9: 25–46. Yoshiyuki M., 1989: A Systematic Study of the Japanese Chiroptera. National Science Museum, Tokyo, 242 pp. 284