The altitudinal distribution of bats of four Plecotus species in Croatia was analysed. P. austriacus and P. kolombatovici are restricted to lower altitudes, under 800 m a.s.l.. P. auritus inhabits continental forest habitats, preferably beech forests, with the exception of Pannonian lowland riparian forests of common oak which also lie under 100 m a.s.l.. As distinct from the other three Plecotus species, Croatian specimens of P. macrobullaris have an extremely wide altitudinal distribution, ranging from sea level to mountain tops, but with most localities under 800 m a.s.l.Analizirano je visinsko rasprostranjenje četiri vrste šišmiša roda Plecotus u Hrvatskoj. P. austriacus i P. kolombatovici su ograničeni na niže visine, do 800 m nad morem. P. auritus nastava kontinentalna šumska staništa, uglavnom bukove šume, ali u Panonskoj nizini i šume hrasta lužnjaka na nadmorskim visinama manjim od 100 m. Za razliku od ostalih vrsta roda Plecotus, nalazi vrste P. macrobullaris u Hrvatskoj imaju osobito široki raspon nadmorskih visina, od morske razine do vrhova planina, ali uz većinu nalazišta ispod 800 m nadmorske visine

I. Pavlinić

N. Tvrtković

English

Analizirano je visinsko rasprostranjenje četiri vrste šišmiša roda Plecotus u Hrvatskoj. P. austriacus i P. kolombatovici su ograničeni na niže visine, do 800 m nad morem. P. auritus nastava kontinentalna šumska staništa, uglavnom bukove šume, ali u Panonskoj nizini i šume hrasta lužnjaka na nadmorskim visinama manjim od 100 m. Za razliku od ostalih vrsta roda Plecotus, nalazi vrste P. macrobullaris u Hrvatskoj imaju osobito široki raspon nadmorskih visina, od morske razine do vrhova planina, ali uz većinu nalazišta ispod 800 m nadmorske visine.The altitudinal distribution of bats of four Plecotus species in Croatia was analysed. P. austriacus and P. kolombatovici are restricted to lower altitudes, under 800 m a.s.l.. P. auritus inhabits continental forest habitats, preferably beech forests, with the exception of Pannonian lowland riparian forests of common oak which also lie under 100 m a.s.l.. As distinct from the other three Plecotus species, Croatian specimens of P. macrobullaris have an extremely wide altitudinal distribution, ranging from sea level to mountain tops, but with most localities under 800 m a.s.l

Pavlinić, I.

Tvrtković, N.

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short communication / kratko priop}enjeALTITUDINAL DISTRIBUTIONOF FOUR PLECOTUS SPECIES(MAMMALIA, VESPERTILIONIDAE)OCCURRING IN CROATIAIGOR PAVLINI] & NIKOLA TVRTKOVI]Department of Zoology, Croatian Natural History Museum,Demetrova 1, HR-10000 Zagreb, CroatiaPavlini}, I. & Tvrtkovi}, N.: Altitudinal distribution of four Plecotus species (Mammalia,Vespertilionidae) occurring in Croatia. Nat. Croat., Vol. 13, No. 4, 395–401, 2004, Zagreb.The altitudinal distribution of bats of four Plecotus species in Croatia was analysed. P. austriacusand P. kolombatovici are restricted to lower altitudes, under 800 m a.s.l.. P. auritus inhabits continen-tal forest habitats, preferably beech forests, with the exception of Pannonian lowland riparian for-ests of common oak which also lie under 100 m a.s.l.. As distinct from the other three Plecotus spe-cies, Croatian specimens of P. macrobullaris have an extremely wide altitudinal distribution, rangingfrom sea level to mountain tops, but with most localities under 800 m a.s.l.Key words: Plecotus macrobullaris, P. auritus, P. austriacus, P. kolombatovici, altitudinal range, CroatiaPavlini}, N. & Tvrtkovi}, N.: Visinsko rasprostranjenje ~etiri vrste {i{mi{a roda Plecotus(Mammalia, Vespertilionidae) u Hrvatskoj. Nat. Croat., Vol. 13, No. 4, 395–401, 2004, Zagreb.Analizirano je visinsko rasprostranjenje ~etiri vrste {i{mi{a roda Plecotus u Hrvatskoj. P. au-striacus i P. kolombatovici su ograni~eni na ni`e visine, do 800 m nad morem. P. auritus nastavakontinentalna {umska stani{ta, uglavnom bukove {ume, ali u Panonskoj nizini i {ume hrasta lu-`njaka na nadmorskim visinama manjim od 100 m. Za razliku od ostalih vrsta roda Plecotus, nalazivrste P. macrobullaris u Hrvatskoj imaju osobito {iroki raspon nadmorskih visina, od morske razinedo vrhova planina, ali uz ve}inu nalazi{ta ispod 800 m nadmorske visine.Klju~ne rije~i: Plecotus macrobullaris, P. auritus, P. austriacus, P. kolombatovici, visinska rasprostra-njenost, HrvatskaINTRODUCTIONHANAK (1969) was the first to report differences in the altitudinal distribution ofPlecotus austriacus Fischer, 1829 and P. auritus Linnaeus, 1758 in central Europe.HORA^EK et al. (2004) summarized observations which characterise P. austriacus as aspecies that lives in the lowlands of Europe. The altitudinal range extends from theNAT. CROAT. VOL. 13 No 4 395¿401 ZAGREB December 31, 2004Croatian Natural History Museum, Demetrova 1, Zagreb, Croatiamountain to the submontane belt (600/800 m a.s.l.), with individual exceptions ofup to 1350 m a.s.l. (hibernacula – SPITZENBERGER, 2002). In Bulgaria, 80% of a totalof 75 sites of P. austriacus are situated below 1000 m a.s.l. (BENDA & IVANOVA, 2003).HORA^EK & \ULI] (2004) stated that P. auritus is a planar/colline to subalpinespecies of mesophile mixed broadleaved forests, with most findings ranging in ele-vation from 200 to 1920 m a.s.l., and winter findings reaching up to 2350 m in theAlps. Findings across northeast Europe are generally within the planar belt, mostlyin boreal coniferous forests. From 22 sites in Bulgaria, 86% lie in altitudes over 1000m a.s.l. (BENDA & IVANOVA, 2003). The Iberian subspecies, P. auritus begognae (PAZ,1994) has a similar distribution pattern in supra-Mediterranean forests lying higherthan 1000 m a.s.l.. The lowest nursery roost of P. auritus reported for Austria wasknown from 150 m a.s.l. (SPITZENBERGER, 2002).Like Plecotus austriacus, P. kolombatovici \uli}, 1980 is restricted to the planar /colline belt (\ULI], 1980; HANAK et al., 2001; SPITZENBERGER et al., 2002), but in theEuropean East Mediterranean. P. macrobullaris Kuzyakin, 1965, newly discovered tooccur in Croatia (KIEFER & VEITH, 2002; SPITZENBERGER et al., 2003) is distributed inthe Caucasus, Alps (SPITZENBERGER 2002; KIEFER & VEITH, 2002), Greek mountains(KIEFER & VEITH, 2002; SPITZENBERGER et al., 2003), Pyrenees (GARIN et al., 2003;KIEFER & HELVERSEN, 2004), and mountains of Crete, Syria and Iran (JUSTE et al.,2004). KIEFER & VEITH (2002), relying on a small sample of specimens from theWestern Alps, maintained that this species was restricted exclusively to themontane and alpine belt ranging from 800 to 1800 m a.s.l.. As exceptions, KIEFER &HELVERSEN (2004) noted findings from 480 m (Pesina near Verona, N-Italy; KIEFER &VEITH, 2002) and 688 m (Lienz, Austria; SPITZENBERGER et al., 2001). SPITZENBERGER(2002) reported an altitudinal range of this species in the Austrian Alps from 670 to1221 m a.s.l., and GARIN et al. (2003) found the summer population in the rockybare ground of Brecha de Rolando (Pyrenees) at an altitude of 2807 m.The aim of this paper is, to describe the altitudinal distribution of these fourPlecotus species in Croatia, using revised and new material from Croatia. Special at-tention is given to P. macrobullaris.MATERIAL AND METHODSThe total sample of altitudinal data of Plecotus in Croatia consists of findingsfrom 75 localities. Plecotus kolombatovici was found in 20 localities, P. macrobullaris in28, P. austriacus in 16 and P. auritus in 18 localities. Most of the data were collectedduring intensive field work in 2001 – 2004, investigating the distribution and iden-tification of P. macrobullaris. Altitudes were noted with GPS devices, and the resultswere proved on state geographical maps on a scale of 1:25.000. A list of localitiesand the method of species identification are presented in another paper (TVRTKOVI]et al., in press). Preserved specimens are housed in the mammal collections of theCroatian Natural History Museum in Zagreb; the Zoological Department of Facultyof Science, University of Zagreb; Natural History Museum Wien; Zoological Mu-seum Berlin; Zoological Museum Budapest; Senckenberg Museum, Frankfurt am396 Pavlini}, I. & Tvrtkovi}, N.: Altitudinal distribution of four Plecotus species occurring in CroatiaMain; Zoological Museum St. Petersburg, and the British Museum /Natural His-tory). The most intensive investigation was conducted in the region of Mt Velebit. De-scription of altitudinal belts and their connections with regional vegetation of thismountain are in TVRTKOVI] & KLETE^KI (1993).RESULTSOnly P. macrobullaris occured in all altitudinal zones in Croatia (Fig. 1), findingsranging from sea level to the top of mountains (max. 1800 m/asl; subalpine belt).The greater part of our findings (19 out of 28 localities) are from altitudes lowerthan 800 m a.s.l. (Fig. 1). According to our results, most P. austriacus and P. kolom-batovici were restricted to the planar or colline belt, and occurred just occasionallyin the submontane or montane belt. In the case of P. austriacus, the highest findingswere recorded in caves (Lokve in Gorski kotar area, 740 m a.s.l.; Bari}a pe}ina,Li~ko Petrovo Selo, 400 m a.s.l.). P. austriacus is restricted to areas with continentalinfluences (Istria, Kvarner Bay, part of Gorski Kotar and Lika, the Pannonian low-land). P. kolombatovici inhabits only the Mediterranean area near the Adriatic coast(from Istria to Dubrovnik) and is restricted completely to xerophilous habitats. Fif-Nat. Croat. Vol. 13(4), 2004 397Fig. 1. Distribution of four Plecotus species findings in different altitudinal belts in Croatia.teen records of the Dinaric subpopulation of P. auritus are distributed between 200to 1600 m a.s.l., but the Pannonian findings (3 sites) ranged from 90 to 900 m a.s.l.There is only one finding of this species from southern slopes (Fig. 2) near theAdriatic coast (@iva voda, N Velebit, 1240 m a.s.l.)DISCUSSIONIn case of Plecotus austriacus and P. kolombatovici our results corresponded withdata from \ULI] (1980) and HANAK et al. (2001). While populations of Plecotusauritus from the Dinaric Alps and Medvednica mountains had a similar altitudinaland ecological pattern to that described in HORA^EK & \ULI] (2004), populationsfrom the southwest part of Pannonian lowlands inhabited planar riparian forests ofcommon oak situated at altitudes from 90 to 110 m a.s.l. (Turopolje forests, DrenovBok; both near Sava river). One P. auritus specimen was found wintering in theUviraljka swallow-hole; TVRTKOVI] et al., 2001) in the isolated Pannonian mountainPapuk at 855 m a.s.l.. This distribution is actually the same as in Austria (SPITZEN-BERGER, 2002).Records of P. macrobullaris from the planar and colline belts of the northern Adri-atic coast in Croatia as well those from the southern slopes of Alps in northern Italy(SPITZENBERGER et al., 2001; 2002; 2003) contradict the assumptions of KIEFER &VEITH (2002) and KIEFER & HELVERSEN (2004) about the altitudinal distribution of398 Pavlini}, I. & Tvrtkovi}, N.: Altitudinal distribution of four Plecotus species occurring in CroatiaFig. 2. Altitudinal distribution of different Plecotus species P. macrobullaris (PM),P. auritus (PR), P. kolombatovici (PK) and P. austriacus (PS), on southern and northernslopes of the Dinaric mountains near the Adriatic coast.this species only »at heights above 800 m«. All along the Adriatic coast, P. macro-bullaris occupies Mediterranean pubescent oak woods with hop-hornbeam and orien-tal hornbeam that cover the southern slopes of the coastal mountains from the sealevel to 800 m a.s.l.. From here, the altitudinal range of this species extends to thesubalpine belt consisting of beech or mountain pine scrub at elevations of 1550 to1800 m a.s.l..JUSTE et al. (2004) suggested that the small genetic differentiation within the P.macrobullaris lineage corresponds to a recent recolonization from a refuge east of themontane areas in western Europe that were not covered by glaciers during the IceAges. JUSTE et al. suggested the Middle East as a potential refuge, but they notedthat the finding of two differentiated »western« and »eastern« clades (SPITZENBERGERet al., 2003) would indicate the existence of more than one refuge. The recent distri-bution of P. macrobullaris suggests a glacial refuge along the northern and easternAdriatic coast.In Croatia, P. macrobullaris has a distribution pattern similar to that of some otherkarst rock-dwelling species, for example the voles Chionomys nivalis (KRY[TUFEK &KOVA^I], 1989) and the endemic Dinaromys bogdanovi (KRY[TUFEK, 1999). Unlike otherspecies, these rodents inhabit all altitudinal belts, but only in karstic limestone ar-eas with crevices and caves. Also most of the Austrian range of P. macrobullaris(SPITZENBERGER, 2002) lies in areas of Alpine karst or very close to it (BOBEK, 1961).Nursery roosts (SPITZENBERGER, 1993; 2002) are situated in churches mostly in the up-per Drau valley (Carinthia) within a few kilometres of the karst slopes. The samesituation with nursery roosts can be found in Croatia, particularly in Istria (Boljun), theLika region (Ogulin, Stajnica, Smiljan), @umberak Mt. (Lije{}e, So{ice, Ka{t, MrzloPolje) and Mt Medvednica (Podsused) (position of localities shown in TVRTKOVI] etal., in press).ACKNOWLEDGEMENTSThe field work was supported by Project 183007 of Croatian Ministry of Science,Education and Sport, and by three projects on the inventory of bats: PINMATRAproject, Project of the Plitvi~ka Jezera National Park and Project of @umberak Na-ture Park – Samoborsko gorje. We would also like to thank D. Holcer, B. Jal`i}, E.Klete~ki, Z. Tadi}, D. Vuku{i}, A. Vuku{i}, [. Tomaji}, BIUS, D. Radovi}, O. Vu-kadinovi}, D. Peli}, M. ]aleta, K. ^ivi} and V. Obersnel for collecting material. M.[a{i}, Z. [ajbl, M. Vukovi} and K. Lattinger helped during field work. We are grate-ful to F. Spitzenberger for providing data from mammal collections in the NaturalHistory Museums in Berlin, London, St. Petersburg and Vienna.Received November 20, 2004Nat. Croat. 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Visinsko rasprostranjenje četiri vrste šišmiša roda Plecotus (Mammalia, Vespertilionidae) u Hrvatskoj

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