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Dondini and Vergari Hystrix It. J. Mamm. (n.s.) 22(1) 2011: 99-104

1972 and 1997. Southwestern Naturalist Temple H.J., Terry A. 2007. The status and di- 44: 495-500. stribution of European . Luxem- PRESENT STATUS OF A RARE SPECIES, Schorcht W., Bontadina F., Schaub M. 2009. bourg: Office for Official Publications of LASIOPTERUS (SCHREBER, 1780), IN HUNGARY Variation of adult survival drives the European Communities, 48 pp. population dynamics in a migrating Vannini M. 1990. Lezioni di statistica. 1 forest bat. J. Anim. Ecol. 78: 1182-1190. Università di Firenze, 216 pp. PÉTER ESTÓK

1Bükk Mammalogical Society, 3300 Eger, Maklári út 77/A, Hungary 2Eszterházy Károly College, 3300 Eger, Eszterházy tér 1., Hungary Corresponding Author e-mail: [email protected]

Received 16 September 2010; accepted 29 May 2011

ABSTRACT - By reviewing all published and unpublished data, the present status of Nyctalus lasiopterus in Hungary was assessed. The species is known from 12 localities in the country, mostly situated in the North Hungarian Mountains. Only one stable population is known, which is located in the Mátra Mountains and has been present each summer since its discovery in 1995. Except for this population, the species is rarely found in other loca- tions. Radio-tracking resulted in the finding of 13 roosting trees, which were exclusively (Fagus sylvatica) in a montane forest. The largest roosting group consisted of more than 50 individuals. Sex ratio was highly female biased; adult males have rarely been found in the country. Lactating females and juveniles were captured regularly, which indicates the presence of a stable nursing population in the country. In Hungary, the main threat to N. lasiopterus is likely to be the logging of old beech forests. Further research is needed to draw a better picture of the Hungarian population of the species and strengthen the conservation of this magnificent bat.

Key words: greater noctule, distribution, sex ratio, reproduction, Hungary

RIASSUNTO - Status di una specie rara, Nyctalus lasiopterus (Schreber, 1780), in Un- gheria. Lo status attuale di Nyctalus lasiopterus in Ungheria è stato valutato analizzando tutti i dati, pubblicati e non, disponibili. La specie è segnalata in 12 località, perlopiù nelle montagne settentrionali. E’ nota una sola popolazione riproduttiva, nei Monti Mátra, che è stata segnalata ogni estate a partire dal 1995. Nelle altre località, la specie è stata rinvenuta solo sporadicamente. La radiotelemetria ha permesso di individuare 13 roost, tutti all’interno di faggi (Fagus sylvatica). Il gruppo di dimensioni maggiori consisteva di più di 50 esemplari. La sex ratio è decisamente in favore delle femmine, mentre esemplari maschi sono stati catturati solo raramente. Femmine allattanti e giovani dell’ anno sono stati cattu- rati con regolarità, suggerendo che la riproduzione della specie in Ungheria avviene rego- larmente. La minaccia principale alla sopravvivenza della specie è il taglio delle foreste di faggio mature. Ulteriori ricerche sono necessarie per ottenere un quadro più preciso della distribuzione della specie nel Paese e sostenerne la conservazione.

Parole chiave: nottola gigante, rapporto sessi, riproduzione, Ungheria

DOI: 10.4404/Hystrix-22.1-4506

INTRODUCTION region. Its distribution area extends from the Iberian Peninsula eastwards to Nyctalus lasiopterus (Schreber, 1780) the Urals, Iran and Kazakhstan, south- is a rare bat species of the Palaearctic wards to Morocco, Libya and possibly

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Algeria (Ibáñez et al. 2004; Simmons thor - were collected to evaluate the present 2005). The species is considered to be status of N. lasiopterus in Hungary. Most rare all over its range, in many Euro- data were obtained between 1993 and pean countries only a few data on the 2010, by mist-netting over 400 nights in the species being available (Ibáñez et al. North Hungarian Mountains, mostly near drinking sites in deciduous forests (Estók 2004). N. lasiopterus is a typical tree- and Gombkötő 2007; Estók 2007b; Estók dwelling bat, roosting in trees all year unpublished). In 2009, the distribution of round. It has been found in the cavities the species at potential habitats of the of very different trees, including both North Hungarian Mountains was also in- deciduous and coniferous species, un- vestigated by the detection of echolocation der the dried leaves of palm trees calls (Estók and Görföl 2009); identifica- (Washingtonia filifera) (Ibáñez et al. tion of the calls was based on the character- 2004) and also in bat boxes (Dondini istics presented by Estók and Siemers and Vergari 2000). Only two subterra- (2009). Radio-tracking was performed at two localities, in the Mátra Mountains and nean occurrences of the species are in the Bükk Mountains (Estók et al. 2007; known: Beck and Gebhard (2000) re- Estók unpublished; Estók and Görföl ported a female specimen which was 2009). Data on sex-identified Hungarian found in a cellar of a castle in Switzer- specimens were used to assess the sex ratio land (possibly an accidental occur- of N. lasiopterus, by the Chi-squared test rence) and Tvrtković and Baltić (1996), for goodness of fit. Expected values were found the remains of 16 N. lasiopterus calculated on the base of 1:1 male-female in a cave in Croatia. In some parts of sex ratio. the Mediterranean area, it may prey upon small migrating birds in autumn RESULTS AND DISCUSSION and spring (Dondini and Vergari 2000; Ibáñez et al. 2001; Popa-Lisseanu et al. 1. Hungarian distribution 2007). The species has been studied actively The species is known from 12 localities in Hungary since the early 1990s. Be- in Hungary (Fig. 1). Most occurrences sides basic faunistics, radio-tracking are associated with the area of the surveys were also conducted to get in- North Hungarian Mountains. Most formation on its habitat requirements mist-netted individuals were captured and the sex ratio of the species was also in the Mátra Mountains, where the only evaluated (Estók 2007a, b; Estók and known stable population dwells. Regu- Gombkötő 2007; Estók et al. 2007; lar samplings at that most significant Estók and Görföl 2009). The aim of Hungarian locality in 2007 showed that this paper is to draw a picture of the the species possibly arrives in late present status of the species in Hun- spring/early summer and leaves the gary, based on the results of research territory in early autumn at the latest. carried out in the last two decades. 2. Habitat, roosting sites MATERIALS AND METHODS All but two specimens were recorded in All available Hungarian data on the species the North Hungarian Mountains, where - including unpublished results of the au- wide woodlands are present. The circum-

100 Estók P. Nyctalus lasiopterus in Hungary

Algeria (Ibáñez et al. 2004; Simmons thor - were collected to evaluate the present 2005). The species is considered to be status of N. lasiopterus in Hungary. Most rare all over its range, in many Euro- data were obtained between 1993 and pean countries only a few data on the 2010, by mist-netting over 400 nights in the species being available (Ibáñez et al. North Hungarian Mountains, mostly near drinking sites in deciduous forests (Estók 2004). N. lasiopterus is a typical tree- and Gombkötő 2007; Estók 2007b; Estók dwelling bat, roosting in trees all year unpublished). In 2009, the distribution of round. It has been found in the cavities the species at potential habitats of the of very different trees, including both North Hungarian Mountains was also in- deciduous and coniferous species, un- vestigated by the detection of echolocation der the dried leaves of palm trees calls (Estók and Görföl 2009); identifica- (Washingtonia filifera) (Ibáñez et al. tion of the calls was based on the character- 2004) and also in bat boxes (Dondini istics presented by Estók and Siemers and Vergari 2000). Only two subterra- (2009). Radio-tracking was performed at two localities, in the Mátra Mountains and nean occurrences of the species are in the Bükk Mountains (Estók et al. 2007; known: Beck and Gebhard (2000) re- Estók unpublished; Estók and Görföl ported a female specimen which was 2009). Data on sex-identified Hungarian found in a cellar of a castle in Switzer- specimens were used to assess the sex ratio Figure 1 - Hungarian occurrences of Nyctalus lasiopterus (triangles: data before 1975; cir- land (possibly an accidental occur- of N. lasiopterus, by the Chi-squared test cles: data after 1990; filled symbols: captured specimens or found remains; open symbol: rence) and Tvrtković and Baltić (1996), for goodness of fit. Expected values were acoustic data; grey circles correspond to the areas of Mátra Mountains (large circle) and found the remains of 16 N. lasiopterus calculated on the base of 1:1 male-female Bükk Mountains (small circle), their size being proportional to the number of observed in- in a cave in Croatia. In some parts of sex ratio. dividuals; squares: main cities; data from Estók 2007b and Estók and Görföl 2009). the Mediterranean area, it may prey upon small migrating birds in autumn RESULTS AND DISCUSSION stances of the two findings of the spe- terus was rediscovered in the Bükk and spring (Dondini and Vergari 2000; cies in lowlands suggest that they were Mountains, lactating females were also Ibáñez et al. 2001; Popa-Lisseanu et al. 1. Hungarian distribution migrating individuals. Our radio- mist-netted at six locations, bringing 2007). tracking data showed a frequent roost evidence of the occurrence of the spe- The species has been studied actively The species is known from 12 localities changing behaviour of N. lasiopterus cies’ reproduction in the country in Hungary since the early 1990s. Be- in Hungary (Fig. 1). Most occurrences (Estók et al. 2007; Estók and Görföl (Gombkötő et al. 1996). Currently, 137 sides basic faunistics, radio-tracking are associated with the area of the 2009), a behaviour typical of forest- records of sex-identified specimens are surveys were also conducted to get in- North Hungarian Mountains. Most dwelling bat species (Lewis 1995), known from Hungary. Two records, formation on its habitat requirements mist-netted individuals were captured which has also been reported for N. those of a migrating female captured in and the sex ratio of the species was also in the Mátra Mountains, where the only lasiopterus in maternity colonies in April, 1960 and its pup, which was evaluated (Estók 2007a, b; Estók and known stable population dwells. Regu- Spain (Popa-Lisseanu et al. 2008). Ra- born in captivity (Topál 1976) were Gombkötő 2007; Estók et al. 2007; lar samplings at that most significant dio-tracked used exclusively beech excluded from the analysis. Females (N Estók and Görföl 2009). The aim of Hungarian locality in 2007 showed that trees (Fagus sylvatica) as roosts (N = = 118) sharply dominate in both July the species possibly arrives in late 13), in a natural, old mountain beech (N = 78, χ2=49.3, df=1, p < 0.001) and this paper is to draw a picture of the 2 present status of the species in Hun- spring/early summer and leaves the forest (Aconito-Fagetum). The largest August (N = 43, χ =25.33, df=1, p < gary, based on the results of research territory in early autumn at the latest. roosting group consisted of more than 0.001) (Fig. 2). Probably some of the carried out in the last two decades. 50 individuals (Estók and Görföl few males (N = 17) captured in these 2. Habitat, roosting sites 2009). two months were subadult specimens. MATERIALS AND METHODS The stable reproduction of the species All but two specimens were recorded in 3. Reproduction and sex ratio in the country is supported by the regu- All available Hungarian data on the species the North Hungarian Mountains, where lar capture of lactating females in the - including unpublished results of the au- wide woodlands are present. The circum- Since the early 1990s, when N. lasiop- Mátra Mountains in the last two deca-

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Figure 2 - Sex ratio of Nyctalus lasiopterus captured in Hungary between 1933 and 2010 (Data from: Topál 1959; Gombkötő et al. 1996; Czájlik and Harmos 1997; Matis et al. 2003; Estók unpublished data). des. In other parts of the North Hungar- of the oldest European records of the ian Mountains, the netting of reproduc- species originates from canton Uri, tive females is sporadic, insufficient to Switzerland, based on the collection of prove the species’ continuous repro- two adult females and one non-volant duction at those localities. juvenile (Fatio 1869). Data on repro- duction also exist from the eastern part CONCLUSIONS of its range (Abelencev and Popov 1956). Recent records of lactating fe- Nyctalus lasiopterus is a rare bat spe- males from Cyprus indicate that the cies in Hungary, its known range being species is also reproducing in the east- limited to the North Hungarian Moun- ern part of the Mediterranean (Benda et tains. As all recent Hungarian locations al. 2007). In Slovakia, nursery roosts of the species are either inside or next to possibly exist, since post-lactating fe- beech forests the species probably se- males and juveniles have been captured lects old beech trees as roosts, but fur- several times (Uhrin et al. 2006). On ther radio-tracking surveys are needed the other hand, strong male dominance to confirm this hypothesis. Currently, and the absence of females in the nurs- the species’stable reproduction is ing period have been reported for known for only one locality, however Greece (Helversen and Weid 1990), further nursing colonies are probably and northern Italy (Ravenna, Emilia- present in other parts of the North Hun- Romagna region) (Lanza 1959), which garian Mountains. Besides the well possibly represent mating and overwin- known Spanish population, available tering sites of the species. records suggest that nursing localities On the whole, despite the apparent exist also in other parts of Europe. One wide range of this species in Europe,

102 Estók P. Nyctalus lasiopterus in Hungary

nursing areas seem to be very localised. REFERENCES There is a significant gap in our current knowledge on the distribution and Abelencev V.I., Popov B.M. 1956. Order status of the species, especially in the Bats. Chiroptera. In: Abelencev V.I., eastern part of the species’ range. Pidoplicko I.G., Popov B.M. (Eds.). Nursery colonies are extremely sensi- Fauna of Ukraine. Volume 1: Mam- mals. Number 1.Vidavnictvo, Kijev, tive seasonal aggregations, therefore pp. 229–446. identifying them is fundamental for the Beck A., Gebhard J. 2000. Riesenabend- effective conservation of N. lasiop- segler (Nyctalus lasiopterus) im terus. The stable nursing colonies of Schloss Hallwyl, Kanton Aargau, the species in Hungary have high con- gefunden. Pro Chiroptera 1: 1–3. servation importance, which has to be Benda P., Hanák V., Horáček I., Hulva P., recognised by local authorities. The Lučan R., Ruedi M. 2007. Bats recent disappearance of N. lasiopterus (Mammalia: Chiroptera) of the Eastern from Bükk Mountains, its second most Mediterranean. Part 5. Bat fauna of significant Hungarian site, where it was Cyprus: review of records with con- firmation of six species new for the is- Figure 2 - Sex ratio of Nyctalus lasiopterus captured in Hungary between 1933 and 2010 regularly mist-netted till 1997, points land and description of a new subspe- (Data from: Topál 1959; Gombkötő et al. 1996; Czájlik and Harmos 1997; Matis et al. out the vulnerability of the species. In cies. Acta Soc. Zool. Bohem. 71: 71– 2003; Estók unpublished data). Hungary, the main threat to N. lasiop- 130. terus is likely to be forestry, i.e. the Czájlik P., Harmos K. 1997. Data on the des. In other parts of the North Hungar- of the oldest European records of the logging of old beech forests. Unfortu- occurrence of bat species in the Kékes ian Mountains, the netting of reproduc- species originates from canton Uri, nately the area of old natural and semi- Észak forest reserve. Fol. Hist. Nat. tive females is sporadic, insufficient to Switzerland, based on the collection of natural beech forests is very limited in Mus. Matr. 22: 345–348 [In Hungarian prove the species’ continuous repro- two adult females and one non-volant the country. Further research at both with English summary]. duction at those localities. juvenile (Fatio 1869). Data on repro- known localities and other potential Dondini G., Vergari S. 2000. Carnivory in duction also exist from the eastern part roosting areas is needed to strengthen the (Nyctalus CONCLUSIONS of its range (Abelencev and Popov lasiopterus) in Italy. J. Zool. (London) the conservation of this magnificent 251: 233–236. 1956). Recent records of lactating fe- bat. Nyctalus lasiopterus is a rare bat spe- males from Cyprus indicate that the Estók P. 2007a. Seasonal changes in the cies in Hungary, its known range being species is also reproducing in the east- sex ratio of Nyctalus-species in North- ACKNOWLEDGEMENTS limited to the North Hungarian Moun- ern part of the Mediterranean (Benda et East Hungary. Acta Zoologica Aca- tains. As all recent Hungarian locations al. 2007). In Slovakia, nursery roosts demiae Scientiarum Hungaricae 53(1): of the species are either inside or next to possibly exist, since post-lactating fe- This paper is dedicated to Professor Emeri- 89–95. tus Benedetto Lanza, who presented valu- Estók P. 2007b. The year of Greater beech forests the species probably se- males and juveniles have been captured able data on this magnificent bat species, Noctule (Nyctalus lasiopterus lects old beech trees as roosts, but fur- several times (Uhrin et al. 2006). On Nyctalus lasiopterus. [Schreber, 1780]) 2007 - review of ther radio-tracking surveys are needed the other hand, strong male dominance I would like to thank Tamás Görföl for his Hungarian data and new results. The to confirm this hypothesis. Currently, and the absence of females in the nurs- help in the field. The research was sup- Proceedings of the Vth Conference on the species’stable reproduction is ing period have been reported for ported by the Hungarian Ministry of Envi- the Bat Conservation in Hungary (Pécs known for only one locality, however Greece (Helversen and Weid 1990), ronment and Water (K-36-08-00042A, K- 2005, december 3-4.) and VIth further nursing colonies are probably and northern Italy (Ravenna, Emilia- 36-09-00326W, K-36-10-00009B), the Conference on the Bat Conservation in present in other parts of the North Hun- Romagna region) (Lanza 1959), which EUROBATS Projects Initiative and the Hungary (Mártély 2007, október 12- garian Mountains. Besides the well possibly represent mating and overwin- Max Planck Society. I would like to thank 14.). CSEMETE Természet- és known Spanish population, available tering sites of the species. two anonymous referees for their useful Környezetvédelmi Egyesület, Szeged, records suggest that nursing localities On the whole, despite the apparent comments on an earlier version of this pp. 80–84 [in Hungarian with English exist also in other parts of Europe. One wide range of this species in Europe, manuscript. summary].

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