In the Western Palaearctic, mountain environ-ments are diversity hotspots representing a signifi-cant proportion of the European biota (Grabherr etal., 2011). Their vertical climate zonation results inhigh habitat diversity (Körner, 2003; Väre et al.,2003; Nagy and Grabherr, 2009) that supports nu-merous chiropteran species (Ulrich et al., 2007),most concentrated at low and mid-elevations (Stutz,1989; Holzhaider and Zahn, 2001; McCain, 2007;Piksa et al., 2013), though some are also foundabove the timberline (Barataud, 2004). Bat prospec-tion efforts, however, are not balanced among differ-ent elevations, as bat diversity studies usually do notinclude alpine environments (but see Aellen, 1962;Garin et al., 2003; Barataud, 2004). Consequent-ly, bats’ use of alpine environments is liable to beunderestimated.
Plecotus macrobullaris, the mountain long-earedbat, is currently known in the Alpine Arc (Trizio etal., 2005; Arthur and Lemaire, 2009; Presetnik etal., 2009; Mattei-Roesli, 2010), Pyrenees (Garin etal., 2003), Dinaric Alps (Pavlinić and Tvrtković,2004; Tvrtković et al., 2005), some Mediterraneanislands (Benda et al., 2008), Anatolia (Karataş andSozen, 2006), Syria (Benda et al., 2006), the Cau -casus (Spitzenberger et al., 2006; Kiefer, 2008), andseveral mountainous regions in the Middle East(Spitzenberger et al., 2006; Kiefer, 2008). Never the -less, its distribution data is largely biased, as morethan half of the presence records come from theAlpine Arc and Dinaric Alps. Moreover, most of theeastern records are museum specimens, for whichonly proximate locations are provided (Spitzen -berger et al., 2006; Kiefer, 2008).
Although P. macrobullaris was initially thoughtto be an alpine species (Kiefer and Veith, 2002)
Review on the geographic and elevational distribution of the mountain
long-eared bat Plecotus macrobullaris, completed by utilising
a specific mist-netting technique
ANTTON ALBERDI1, 2, INAZIO GARIN1, OSTAIZKA AIZPURUA1, and JOXERRA AIHARTZA1
1Department of Zoology and Animal Cell Biology, Faculty of Science and Technology, University of The Basque CountryUPV/EHU, Sarriena z.g., Leioa E-48940, The Basque Country
The mountain long-eared bat, Plecotus macrobullaris, is a recently described species, and characterisation of its geographic andelevational distribution is still in progress. Captures in various environments led to a controversial ecological perception, with P. macrobullaris initially defined as an alpine species but subsequently found in the Mediterranean seashore and other lowland areas.Sampling efforts hitherto were uneven; this bat has been studied more thoroughly in Western Europe than in the eastern portion ofits range, and in lowlands more than highlands or alpine environments. For greater insight into its distribution pattern, we conducteda field survey in several mountain areas of its known range, using a novel mist-netting technique (described herein) that has provenvery useful for targeting and capturing low-flying open-space bats in alpine environments. We also gathered all available distributiondata on this species from published resources and by contacting researchers, obtaining records at 351 total localities (including 113 from other authors’ unpublished reports and 45 from our own fieldwork). We concluded that P. macrobullaris is present in themain Western Palearctic mountain ranges, extending from the Pyrenees to the Middle East, and has an elevational distribution fromsea level up to 2,800 m. The high number of these bats captured foraging above the timberline, in addition to the exclusivelymountainous distribution, indicate that the species is indeed alpine, showing a pattern similar to other highly mobile vertebratesrestricted to mountain areas and absent from flatlands. Nevertheless, its apparent elevational distribution may still be biased towardlower areas, due to the scarcity of surveys in high mountain habitats.
when described in 2002 in Austria (Kock, 2002;Spitzen berger et al., 2002, 2003), this was chal-lenged when the bat was found almost at sea level(Pavlinić and Tvrtković, 2004; Tvrtković et al., 2005;Presetnik et al., 2009), showing an elevational distribution even lower than that of Plecotus auritusin some cases (Mattei-Roesli, 2010).
Nevertheless, most published records of thespecies (64% of all localities) correspond to mater-nity colonies in buildings at 400–1400 m abovemean sea level (a.s.l.). This bat has seldom beenrecorded when foraging at night, and in those fewcases was captured either at water points orcommuting routes (Tvrtković et al., 2005; Benda etal., 2006, 2008). Additionally, some captures at2400– 2800 m in the Pyrenees (Garin et al., 2003)and recent studies on its ecology (Arthur andLemaire, 2009; Alberdi et al., 2012) suggest that P. macrobullaris might be more linked to subalpineand alpine belts than previously thought. Thus, hith-erto published assessments of its geographic and ele va tional distribution may stem from incompleteand lowland-biased inventories.
As noted above, supraforestal habitats are sel-dom included in bat diversity studies (Stutz, 1989;Presetnik et al., 2009; Mattei-Roesli, 2010). In the literature’s few exceptional cases where nets have been set above the timberline, mountain passes were used as sampling places (e.g., Aellen,1962; Ber trand, 1992; Garin et al., 2003). How-ever, as passes are the shortest way to cross moun-tain chains, and many species — including forestalor even lowland species, e.g., Miniopterus schrei-bersii (Hutson et al., 2008) — fly through themwhen commuting between valleys (Aellen, 1962;Bertrand, 1992), that netting strategy may be mis-leading for distributional conclusions, because occasional dispersal is also likely to be recorded. Onthe other hand, placing nets in water points is usual-ly of meagre effectiveness in regions with highlyavailable drinking water. Bats (mainly the smallones) can utilise any of the thousands of watersources usually available in alpine environments,and capture rates are therefore low. Conversely, set-ting nets in open space — including supraforestalhabitats — is advised against by some authors due to alleged low effectiveness (Agnelli et al., 2006).However, it has been shown that the traditional perception of low capture rates in some foragingsites may be misleading, and that the perception of abundance of some species can change dramati-cally when using alternative methods (Carroll et al.,2002).
To elucidate the geographic and elevational dis-tribution of P. macrobullaris, we first present newdistribution data in several Western Palearcticmount ain ranges, achieved by a specific mist-nettingtechnique targeted to bats foraging in open supra -forestal habitats; second, we compile and reviewthis species’ published and unpublished records,paying particular attention to distribution, elevation,and sampling method.
MATERIALS AND METHODS
Ethics Statement
Capture and handling protocols met published guidelinesfor treatment of animals in research and teaching (Animal Be haviour Society, 2006) and were approved by the EthicsCom mittee at the University of The Basque Country (Refs.CEBA/238/2012/GARIN ATORRASAGASTI). All new cap-tures in the field were carried out under license provided by thecorresponding local agencies and governments.
Mist-netting in Open Supraforestal Habitats
To sample P. macrobullaris at higher elevations, we mist-netted bats during the summers (July–August) of 2009– 2012 inthe Pyrenees (Spain), Alps (Italy and Slovenia), PindosMountains (Greece), and Caucasus (Georgia). Overall, mist-nets were placed in 54 sites above the timberline, at 1450–2400m a.s.l. The timberline, defined as the line connecting the high-est limits of forest patches (Körner, 2003), which is usually artificially lowered to promote summer pasturing of livestock(Miehe and Miehe, 2000; Olsson et al., 2000; Dirnböck et al.,2003; Garcia-Gonzalez, 2008), ranges from 1500 m a.s.l. inmany areas in the Pyrenees and Maritime Alps (Italy) to 2500 ma.s.l. in the Caucasus. Considering that in mountainous areaslandscape usually changes drastically, with noticeable elevationand habitat differences, nets placed further than 1 km from eachother were considered to be different sampling sites.
Most sampling sites were reached after 1–3 hours hiking,and all the sampling material was carried out in back-packs. Toreduce the likelihood of capturing mountain long-eared batsduring non-foraging activities, we placed mist-nets (2.5 m tall)in meadows and rocky areas, which are their putative foraginggrounds (Alberdi et al., 2012 — Fig. 1). To improve capturesuccess in such extensive habitats, we set nets at ground levelcombining units of different lengths that fitted to irregularitiesof terrain and were linked in lines in a zigzag formation (at ap-proximately 130° angles) for greater stability. The length of net-lines ranged 80–140 m depending on topography of samplingsites. Sticks were affixed to the ground from both sides of thenet-line using ropes and picks, so the mist-nets were less affect-ed by the changes of wind direction and intensity common inmountain environments. Nets were set during daylight, keptclosed until 15 minutes after dusk to avoid catching birds andbundling up beetles, then checked every 15 minutes, and weretypically kept open until 3 am, except when adverse meteoro-logical conditions occurred.
Species were identified in situ based on general mor-phology and biometric measurements (Dietz and Helver sen,
452 A. Alberdi, I. Garin, O. Aizpurua, and J. Aihartza
2004), and were subsequently genetically verified using mito-chondrial markers ND1 (ND1F2 and ND1R primers — Kawaiet al., 2002) and Cyt-b (Molcit-F and MVZ-16 primers — Smithand Patton, 1993; Ibáñez et al., 2006). We took note of the net-bag where bats were captured as indicative of their flightheight above the ground, and used Chi-squared test to assess theprobability of gathered values happening at random. We com-pared elevational distribution of sampling effort with capturesof P. macrobullaris by Kernel Density Esti mation (KDE). Chi-squared test and Kernel Density Estimation were performed using R 2.9.2 software available at CRAN (http://cran.r-project.org/).
Data Gathering and Review
We compiled all existing presence data on mountain long-eared bats by revisiting information published in scientific arti-cles, atlases, and books, and by contacting researchers, natureconservation organisations, and naturalists (Table 1).
RESULTS
Mist-netting in Open Supraforestal Habitats
After 54 sampling nights and 250 hours of net-ting effort, we captured 285 bats in 48 of the 54 sam-pled sites, at elevations between 1450 and 2400 ma.s.l. The captures of P. macrobullaris fitted wellthe trapping effort carried out throughout the eleva-tional range sampled, and no capture reduction was
observed at highest elevations (Fig. 2). Bats werecaught in subalpine and alpine meadows as well asbare rock areas with scarce vegetation. Captured in-dividuals belonged to ten species (Table 2), with themountain long-eared bat predominant: 197 P. ma-crobullaris constituted 70% of all captures, followedby Myotis nattereri sensu lato (12%; see Sa licini etal., 2011) and M. myotis (6%). We captured more fe-male P. macrobullaris (58%) than males (42%),whereas 65% of the other species were males.Plecotus macrobullaris and M. nattereri were cap-tured in the two lowest bags of the mist-nets, i.e. lessthan one metre above the ground on average andlower than expected by chance (χ2 = 45.8, P < 0.05and χ2 = 23.0, P < 0.05, d.f. = 4, respectively).
This thorough sampling in the Pyrenees increas -ed the known locations of P. macrobullaris from 5 to46 in three years; in a single summer, we were ableto capture 106 individuals in 17 of the 19 sampledsites. Additionally, we captured mountain long-eared bats at six different locations in the PindosMountains and Caucasus, in single-week surveys.
Distribution of Plecotus macrobullaris
Our mist-netting in alpine environments resultedin 41 new supraforestal localities of P. macrobul -laris. Additionally, we captured the species at four
Geographic and elevational distribution of Plecotus macrobullaris 453
FIG. 1. Net-setting and sampling sites: (A) Setting a net-line at Plan d’Aniz (Pyrenees), alpine meadow at 1750 m a.s.l., where 21individuals belonging to five species were captured; (B) Insolas (Pyrenees), alpine terrace at 2370 m a.s.l., where 14 P. macrobullariswere captured; (C) Viola (Alps), a former clogged lake at 2400 m a.s.l., where two M. nattereri were captured; and (D) La Ripera
(Pyrenees), valley bottom at 1550 m a.s.l., where 14 individuals belonging to four species were captured
A
B C D
other localities below the timberline, and we gath-ered 113 unpublished records by contacting otherauthors, for an overall compilation of 351 presencelocalities throughout its geographic distribution(Table 3).
The mountain long-eared bat is present in themain mountain ranges of Southern Europe, the NearEast, and the Middle East (Fig. 3): the Pyrenees (46locations), Alps (177), Dinaric Alps (80), PindosMountains (7), Taurus Mountains (9), KaçkarMountains (1), Upper and Lower Caucasus (11),Anti-Lebanon (4), Guneydogu Toroslar Mountains
(1), Alborz Mountains (2), and Zagros Mountains(5). It is also present in the Mediterranean islands ofCorsica (1) and Crete (7).
The total compiled data shows that the species’known localities span 0–2800 m a.s.l., though therecorded elevational distribution differs amongmountain ranges (Fig. 4). Approximately half (52%)of all known P. macrobullaris localities are roosts,mainly buildings (160 records) and the rest (23 re -cords) subterranean cavities such as caves, mines,and tunnels. Mist-netting provided 25% of therecords: 6% with nets set at water points, 14% in
454 A. Alberdi, I. Garin, O. Aizpurua, and J. Aihartza
TABLE 1. Information sources on the distribution of P. macrobullaris. For records appearing in more than one publication, only onewas counted
Type Observer Region # records
Unpublished C. Flaquer and X. Puig-Montserrat Pyrenees (Catalonia) 10Museu de Granollers-Ciències NaturalsJ. C. Albero Pyrenees (Spain) 1J. Girard-Claudon Alps and Prealps (France) 34LPO Coordination Rhône-AlpesGroupe Chiroptères Rhône-AlpesLigue pour la Protection des OiseauxDrôme et IsèreG. Kapfer Alps and Prealps (France) 24Groupe Chiroptères de ProvenceC. Schönbächler Alps (Switzerland) 1M. Spada Alps (Italy) 13Insubria UniversityParco Paneveggio-Pale di San Martino Alps (Italy) 2Database of Centre for Cartography of Slovenia 20Fauna and Flora (CKFF)K. Sachanowicz Albania 2P. Georgiakakis Crete (Greece) 7
Published Garin et al. (2003) Pyrenees (Spain) 2Dejean (2009) Pyrenees (France) 1Desmet (2008) Alps (France) 1Juste et al. (2004) Alps (France), Azerbaijan, Iran 3Ashrafi et al. (2010) Alps (Switzerland) 14Kiefer et al. (2002) Alps (Austria) 1Kiefer (2008) Alps (Austria, France), Dinaric Alps, Middle East 10Mattei-Roesli et al. (2010) Alps (Switzerland) 56Reiter and Zahn (2006) Alps (Austria, Switzerland) 1Spitzenberger et al. (2002) Alps (Austria) 2Spitzenberger et al. (2006) Alps (Austria, Italy), Dinaric Alps, Middle East 23Kiefer and von Helversen (2004) Corsica, Alps (France, Switzerland, Austria) 5Presetnik et al. (2009) Slovenia 26Ramovs et al. (2010) Slovenia 1Zagmajster et al. (2012) Southeastern Italy 2Tvrtković et al. (2005) Dinaric Alps (Croatia) 24Sachanowicz and Ciechanowski (2006) Dinaric Alps (Albania) 1Bekker and Boshamer (2007) Dinaric Alps (Macedonia) 1Benda et al. (2008) Crete (Greece) 4Karataş and Sozen (2006) Turkey 10Benda et al. (2004) Syria 3Yavruyan et al. (2007) Armenia 1
New data Present study Pyrenees (The Basque Country, Spain, Catalonia),Alps (Italy, Slovenia), Pindos Mts., Caucasus 45
Total 349
meadows, and 5% in unspecified sites. Captures in subalpine and alpine meadows during the pres-ent fieldwork represent almost half of the 85 com-piled outdoor localities. The capture method for an-other 83 records (23%) could not be determined(Table 3).
PyreneesThe mountain long-eared bat has been recorded
throughout this entire mountain range except thewestern and eastern lower ends. Prior to this study,five locations were known in the Central Pyrenees(Spain, France, and Andorra — Garin et al., 2003;Kiefer and Helversen, 2004; Dejean, 2009; Flaqueret al., 2010). The uppermost height is a mount-ain pass at 2800 m a.s.l., near another mist-net-ting capture at 2400 m a.s.l. in Ordesa and MontePerdido National Park (Garin et al., 2003). Fe-male bats, including a lactating individual at 2800 m a.s.l., were captured at both locations. Weadded 42 new records (31 our own and 11 reportedby other authors), 40 of them obtained using mist-nets in subalpine and alpine meadows 1500–2450 m a.s.l.
AlpsPlecotus macrobullaris is distributed across this
entire mountain range, from the Provençal Prealps(France) to Slovenia and southern Austria. In theFrench Prealps and Alps, we gathered 42 and 19 re -cords, respectively (only three previously published— Juste et al., 2004; Kiefer and Helversen, 2004;Desmet, 2008). Some prealpine locations almostreach the Mediterranean seashore, and the highest is1940 m a.s.l. (J. Girard-Claudon, personal commu-nication; G. Kapfer, personal communication).
Known to be very common in the Central Alps, especially in the Swiss Valais (> 14 roostingsites) and Ticino (> 60 roosting sites — Ashrafi etal., 2010; Mattei-Roesli, 2010; Rutishauser et al.,2012), the species is also known in Liechten-stein with a single record (Kiefer and Helversen,2004) and in the Prealps near Italy’s Lake Maggio-re (Trizio et al., 2005). Almost all of those re-cords correspond to maternity colonies in villagebuildings at the valley bottom. We added the first recorded mountain long-eared bats in Aosta Val-ley, caught with mist-nets in alpine meadows at2050 m a.s.l.
Geographic and elevational distribution of Plecotus macrobullaris 455
FIG. 2. Kernel Density Estimation of the sampling effort (grey area) and captures of P. macrobullaris (black line) across thesupraforestal elevational gradient
TABLE 2. Summary of bats captured using the mist-netting technique described herein. An asterisk (*) indicates species captured inonly one place
Species Total individuals Males Females Elevation range
The species is known to be common in theEastern Alps as well: in Austria (21 records — Spi -tzen berger et al., 2002, 2006; Kiefer and Hel versen,2004; Reiter and Zahn, 2006; Kiefer 2008), Italy(unknown number — Chirichella et al., 2003), andnorthwestern Slovenia (10 records — Presetnik etal., 2009). M. Spada (personal communication) re-ported its presence at 13 locations in Italy, and wecaptured mountain long-eared bats at four moreEastern Alp locations, including two from mist-net-ting above the timberline in western Slovenia. Mostre cords in this area also correspond to maternityroosts below 1600 m a.s.l., though captures have occurred up to 2000 m in Italy (M. Spada, personalcommunication). One of our captures, in Triglav Na -tion al Park (Slovenia) at 1750 m a.s.l., is 800 mhigher than the highest hitherto published record ofP. ma crobullaris in Slovenia (Presetnik et al., 2009).
Dinaric Alps and Pindos MountainsPublished records locate the species in 59 points
stretching from southern Slovenia and southeasternItaly to Greece, where it inhabits mainly karsticmountainous regions but is lacking from the easternlowland plateau. It is known to range from sea levelup to 1800 m a.s.l. in the Dinaric Alps (Pavlinić andTvrtković, 2004; Tvrtković et al., 2005; Pre setnik etal., 2009). In Croatia, P. macrobullaris has been re corded at 22 localities; two locations are known in Trieste (Italy; Zagmajster et al., 2012), as also occurs in Bosnia-Herzegovina (Tvrtković et al.,2005); and single records are published for Monte -negro (Juste et al., 2004), Macedonia (Bekker andBosh amer, 2007), and Albania (Sachanowicz andCie chanowski, 2006).
We gathered information about 20 additional unpublished localities in Slovenia (CKFF, 2011), inthe northern edge of the Dinaric range and its vicin-ity, corresponding mainly to maternity colonies inchurches below 1000 m a.s.l. Two more unpublishedlocations were gathered from Albania (K. Sacha -nowicz, personal communication) and four morefrom the Pindos Mountains of Greece (P. Ge or - giakakis, personal communication). We also mist-netted mountain long-eared bats at three other loca-tions in Greece, all in supraforestal environments1600–1950 m a.s.l.
Anatolia and Anti-Lebanon MountainsAll Levant records were previously published: in
the Anatolian peninsula, the species has been foundin the Köse Dağları mountain range in the north (onerecord — Karataş and Sozen, 2006) and near the
456 A. Alberdi, I. Garin, O. Aizpurua, and J. AihartzaT
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Taurus Mountains in the southwest (nine records —Juste et al., 2004; Karataş and Sozen, 2006; Spi -tzenberger et al., 2006). On the east face of the Anti-Lebanon mountain range, it has been recorded infour places (Benda et al., 2004; Shehab et al., 2007).The estimated elevation range of P. macrobullarisin Anatolia is quite imprecise, as many capture locations are not exactly known; nonetheless, all of those records are above 1000 m a.s.l., with at leastone reaching 1900 m (Karataş and Sozen, 2006). Its
elevational distribution is similar in the Anti-Le b -anon, with captures between 1000 and 1400 m.
Caucasus, Zagros, and Alborz MountainsIn the Upper Caucasus, only a few records of the
species (including the holotype) in the northern Cau -casian countries (Spitzenberger et al., 2006; Kiefer,2008) precede this study, whereas in the LesserCaucasus, P. macrobullaris has been reported inGeorgia, Armenia, and Azerbaijan (Kiefer, 2008).
Geographic and elevational distribution of Plecotus macrobullaris 457
FIG. 3. Presence of P. macrobullaris on an elevation map of the Western Palearctic
FIG. 4. Elevational distribution (m a.s.l.) of P. macrobullaris in several mountain chains. Boxes limit the 5% and 95% percentiles;vertical lines show minimum and maximum values and horizontal lines indicate the mean value. Note that elevation data of severallocations, mainly in Anatolia, Caucasus, Zagros and Alborz Mountains refer to the municipalities rather than to the precise capture
sites, and consequently, the lower limits of elevation ranges may be biased downwards
Mountain ranges
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There is a single record in southeastern Turkey’sGuneydogu Toroslar Mountains (Kurdistan) (Kiefer,2008). Four records are known in the central andnorthern Zagros Mountains (Spitzenberger et al.,2006; Kiefer, 2008) and two more on the border be-tween Iran and Azerbaijan, at the northern edge ofthe Alborz mountain range, next to the Caspian Sea(Spitzenberger et al., 2006; Kiefer, 2008). All cap-tures in the eastern regions of the Zagros and AlborzMountains occurred between 1200 and 2500 m a.s.l.(Spitzenberger et al., 2006; Kiefer, 2008). We mist-netted the species at three localities in the GeorgianCaucasus (Khevsureti and Tusheti regions) above2000 m, which are its highest known records in thatmountain range.
Corsica and CretePlecotus macrobullaris has been found in two
Med iterranean islands: a single hitherto publishedrecord from Corsica (Kiefer and Helversen, 2004),and in Crete, four reported localities (Benda et al.,2008) plus three unpublished findings (P. Georgia -kakis, personal communication). All of these re cordscorrespond to captures in cave entrances or build ingprospections. The species is known to occur at up to1550 m a.s.l. in Crete (Benda et al., 2008).
DISCUSSION
Mist-netting Above the Timberline
Our four-year experience sampling for bats in thevarious European mountain chains has shown thatthe mist-netting method we used is valuable forcatching hard-to-trap bats in supraforestal environ-ments. Eight of the ten species captured were caughtseveral times and in different mountain ranges(Table 1), suggesting that they forage frequently inalpine habitats. Myotis nattereri (sensu lato) and P. macrobullaris were the only species captured inthe four surveyed mountain chains, and were thetwo most abundant bat species we found in supra -forestal habitats. Previous works in such environ-ments yielded similar results, with undeterminedPle co tus spp. or P. auritus being the most frequent-ly encountered bats at high elevations (Aellen, 1962;Bertrand, 1992; Barataud, 2004). This suggests that the undetermined Plecotus species recorded using ultrasound detectors in such studies (Ba-rataud, 2004) most likely belong to P. macrobul -laris. Sim ilarly, bats previously identified as P. au -ritus at non-forested locations at high elevations(Aellen, 1962; Bertrand, 1992) may actually have
been P. ma cro bullaris, as the latter species was notyet described when those studies were performedbut is now known to be common in those surveyedareas.
Our best capture performance occurred in flatmeadows at hanging valley bottoms, natural alpineterraces, or clogged lakes (Fig. 1). Conversely, mist-netting attempts were not so successful in either extremely humid zones such as moors or in overlydry meadows. We achieved better results in short-grass meadows — both moderately grazed and non-grazed — than in grasslands with tall vegetation,presumably because the capture of moths in shortgrass may be less hampered by clutter. As bats werecaptured in supraforestal environments only fromJuly to Sep tember, we cannot discard the possibilitythat seasonal changes in their foraging elevationrange or habitat preferences are the result of changesin meteorology and/or prey availability in meadows.
Most methods for capturing bats in their huntinggrounds have been devised for cluttered environ-ments (Kunz and Kurta, 1988; Sedlock, 2001; Car -roll et al., 2002). The mist-netting method detailedherein, however, has shown great success in openenvironments, particularly for catching low-flyingbats in their foraging grounds, in high-mountain ar-eas where water is not a limited resource. None the -less, as with every other bat-capture method used,this technique may also be prone to bias. Low-flyingbats such as P. macrobullaris, M. nattereri, and M. my-otis have the highest probability of being caught,where as species such as Hypsugo savii or Ta dari dateniotis usually fly several metres above the reach of nets set at ground level; hence, the abundance of the two latter species in alpine environments willprobably be underestimated by this method.
Geographic Distribution
The Alps, Northern Dinaric Alps, and Pyreneeshave been most surveyed and hold 85% of the cur-rent total localities of P. macrobullaris. Therefore,its actual distribution area may still be underrepre-sented, and additional sampling efforts in othermountain ranges are still needed for a complete assessment of its presence. Nevertheless, the dataherein allow inferring a quite precise picture of boththe geographic and elevational distribution of the mountain long-eared bat, which appears restrict-ed to mountain chains or their close surround-ings and absent from non-mountainous regions. This broad pattern can be also seen regionally, e.g.,in Switzerland (Ashrafi, 2010; Rutishauser et al.,
458 A. Alberdi, I. Garin, O. Aizpurua, and J. Aihartza
2012), Slo venia (Presetnik et al., 2009), and Croatia(Tvrtko vić et al., 2005), where P. macrobullaris ispresent in mountainous areas but is scarce or lackingin lowlands.
This alpine distribution pattern, unparalleledamong bats, is only shared by a handful of speciesfrom other vertebrate taxa. Among mammals, thedistribution of the snow vole (Chionomys nivalis)re sembles that of P. macrobullaris (Kryštufek andAmori, 2008). Moreover, birds such as the alpine ac-centor (Prunella collaris), alpine chough (Pyrrho co -rax graculus), and white-winged snowfinch (Monti -fringilla nivalis) show the highest similarity to thisbat’s pattern (Aulagnier et al., 2008; Sven sson et al., 2009), as their distribution is closely related tomountain ranges and they all breed in alpine envi-ronments. Furthermore, these alpine birds are alsoharboured in Corsica and Crete (Svensson et al.,2009) — the two Mediterranean islands where P. ma crobullaris has been recorded.
Elevational Distribution
Captures of mountain long-eared bats at 41 local-ities above the timberline — 19 between 2000 and2800 m a.s.l. — confirm that the species uses alpineenvironments regularly, as suggested by the highprevalence of alpine moths in its diet (Alberdi et al.,2012). Except in the Pyrenees, where no nets wereerected above the previously known uppermost lim-it (2800 m a.s.l. — Garin et al., 2003), our mist-netsampling in supraforestal environments has in-creased the top recorded elevations of the species inthe surveyed mountain ranges. Thus, the known ele-vation range of mountain long-eared bats has nowrisen 400 m in the Western Alps (to 2050 m a.s.l.)and 800 m in the Julian Alps (to 1750 m). Similarly,the highest localities in the Pindos Mount ains andthe Caucasus have now been reset at 1960 m and2100 m a.s.l., respectively. It is noteworthy thatwhile the uppermost elevation of P. macrobullarisincreased significantly from sampling in alpine en-vironments, that of the other long-eared bats did not.This suggests that the perception of P. auritus inhab-iting higher environments than P. macrobullaris incertain places (Mattei-Roesli 2010) may be an arte-fact of sampling effort bias toward lower areas.Moreover, since the highest altitude belts are un-represented in our sample, we cannot discard the elevational range of P. macrobullaris to be evenbroader than illustrated here. The fact that the cap-ture rate did not show any noticeable reductionwhen sampling at higher elevation (Fig. 2) supports
this idea and is consistent with previous capture re -cords (up to 2807 m a.s.l. — Garin et al., 2003).
On the other hand, the reasons for lower capturesuccess in supraforestal habitats in the Alps than inthe Pyrenees may extend beyond a lesser samplingeffort. We cannot discard the possibility that bats’lesser use of supraforestal habitats in the Alps thanin other mountain chains may result from theirharsher climatic conditions; with temperatures ap-proximately 1ºC colder at 1500 m and 0.5ºC colderat 2000 m than those of the closest ranges (data fromHijmans et al., 2005; WorldClim database, http://www.worldclim.org/), the Alps are the coldestmountain range in the Southwestern Palearctic.Moreover, the elevational range of for ests is broad-er in the Alps than in other mountain chains, leadingto a larger separation between lowland and highlandopen areas by woodlands.
Conclusions
Surveys conducted in various European moun-tain ranges have shown that several bat speciesmake use of alpine environments, with the mountainlong-eared bat being an outstanding example.Supra forestal captures, and our thorough review ofpublished and unpublished locality records, depict P. macrobullaris as abundant in alpine environ-ments, strictly linked to mountain regions, with thesame pattern as other alpine vertebrates. The reasonswhy P. macrobullaris is tied to mountainous areas,however, are unknown.
Evaluation of the samplings in alpine environ-ments shows that the reported elevational distribu-tion of bats is generally biased toward lower alti-tudes, so further survey efforts at higher elevationsare desirable to obtain a complete distributional andecological view of this group of mammals. Themist-netting methodology detailed in this work hasproven particularly useful for capturing low-flyingbats foraging in open supraforestal environments.
ACKNOWLEDGEMENTS
We greatly appreciate the collaboration of all the institu-tions that kindly provided information about P. macrobullaris:Museu de Granollers-Ciències Naturals, Ligue pour la Pro tec -tion des Oiseaux de Rhône-Alpes, Groupe Chiroptères Rhône-Alpes, Ligue pour la Protection des Oiseaux Drôme et Isère,Groupe Chiroptères de Provence, Università degli Studidell’Insubria, Parco Paneveggio-Pale di San Martino, and theCentre for Cartography of Fauna and Flora (CKFF). We are especially thankful to C. Flaquer, X. Puig-Montserrat, J. C. Al bero, J. Girard-Claudon, G. Kapfer, C. Schönbächler, M.Spada, P. Pre setnik, K. Sachanowicz, and P. Georgiakakis,
Geographic and elevational distribution of Plecotus macrobullaris 459
whose information made this work possible, and to our Geor -gian friends, Archil Kikodze and Vakho Kirikashvili, who sokindly guided us in the Caucasus Mts. and introduced us to theirpeople. We are very grateful as well to the agencies that licensedour captures in the surveyed geographic areas. The Basque Gov -ern ment (research projects IT385-07 and IT301-10, and pre -doctoral grants BFI-2010-190 and BFI-2009-252) and the Uni -versity of The Basque Country UPV/EHU supported this work.
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