Activity and Foraging Habitats of Miniopterus

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Activity and foraging habitats of Miniopterus schreibersii (Chiroptera, Miniopteridae) in southern France : implications for its conservation Stéphane Vincent, Mélanie Némoz, Stephane Aulagnier

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Stéphane Vincent, Mélanie Némoz, Stephane Aulagnier. Activity and foraging habitats of Miniopterus schreibersii (Chiroptera, Miniopteridae) in southern France : implications for its conservation. Hys- trix, the Italian Journal of Mammalogy, Associazione Teriologica Italiana, 2011, 22 (1), pp.57-72. 10.4404/Hystrix-22.1-4524. hal-02651396

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APPENDIX Gazetteer ACTIVITY AND FORAGING HABITATS OF MINIOPTERUS SCHREIBERSII (CHIROPTERA, MINIOPTERIDAE) IN ~ Altitude Locality Governorate Coordinates [m a.s.l.] SOUTHERN FRANCE: IMPLICATIONS Aden Aden 12° 48‟ N, 45° 02‟ E 20 FOR ITS CONSERVATION Al Ahjur Al Mahwit 15° 31‟ N, 43° 52‟ E 2400 1 2 3 Al Anad Lahj 13° 17‟ N, 44° 45‟ E 385 STEPHANE VINCENT , MELANIE NEMOZ , STEPHANE AULAGNIER * Al Hadr, W of Lawdar Abyan 13° 53‟ N, 45° 48‟ E 1143 1 Al Makha Taizz 13° 19‟ N, 43° 15‟ E 5 Ligue pour la Protection des Oiseaux - Drôme, pôle Chiroptères, 10 rue Roch Grivel, 26400 Crest, France Al Nueimah, WNW of Buwayash Hadramaut 14° 36‟ N, 49° 06‟ E 285 2Société Française pour l’Etude et la Protection des Mammifères / Conservatoire Régional Am Rija‟, Wadi am Rija‟ Lahj 13° 02‟ N, 44° 34‟ E 300 des Espaces Naturels de Midi-Pyrénées, B.P. 43053, 31025 Toulouse cedex 03, France 3 Ancient Great Dam, W of Ma‟rib Ma‟rib 15° 24‟ N, 45° 16‟ E 1125 Société Française pour l’Etude et la Protection des Mammifères / Comportement et Ecolo- gie de la Faune Sauvage, Institut National de Recherche Agronomique, B.P. 52627, 31326 NNW of Ash Shuqayrah, Wadi Bani Khawlan Taizz 13° 20‟ N, 43° 43‟ E 470 Castanet Tolosan cedex, France; Ba Tays, Wadi Bana Abyan 13° 21‟ N, 45° 18‟ E 175 *Corresponding author, E-mail: [email protected] Damqawt, wadi NW of the village Al Mahra 16° 35‟ N, 52° 50‟ E 25 Received 4 October 2010; accepted 20 March 2011 Ghayl Ba Wazir, karstic hole Hadramaut 14° 48‟ N, 49° 23‟ E 120 Halhal, NE of Hajjah Hajjah 15 44‟ N, 43 37‟ E 1060 ABSTRACT - Comprehensive knowledge of roosting and foraging ecology is essential for W of Hammam Ali Dhamar 14° 41‟ N, 44° 07‟ E 1585 conserving bats. Therefore, the four-year LIFE Nature programme "Conservation of three cave-dwelling bats in Southern France" included an autoecological study of Miniopterus Hawf, gardens above the town Al Mahra 16° 39–40‟ N, 53° 03-05‟ E 230-735 schreibersii, a highly gregarious cave-dwelling species. In a colony of 3-5,000 adults, 21 Jebel Bura, W of Riqab Al Hudaydah 14° 52‟ N, 43° 25‟ E 320 females were radio-tracked during pregnancy and lactating periods, when some of them Kadamat Al „Abdali, Wadi Tuban Lahj 13° 08‟ N, 44° 51‟ E 200 switched roosts within a 30 km radius around the maternity colony. Every night, for about 6 desert NW of Lahj al Hutah Lahj 13° 10‟ N, 44° 49‟ E 260 hours, each bat flew far from the roost (4.1 to 29.2 km) to forage on several small feeding areas (1 to 9 over a few nights). Mean individual home-range estimation averaged 10837 ha Mashgab, S of Ash Shamsara Taizz 13° 21‟ N, 43° 57‟ E 1170 for pregnant females, 22318 ha for lactating females. Urban areas lighted by white street S of Najd An Nashamah Taizz 13° 22‟ N, 44° 02‟ E 1235 lamps were used extensively. Some bats also foraged selectively in deciduous or mixed Sana village, Sana‟a Sana‟a 15° 17‟ N, 44° 10‟ E 2490 woodlands and in orchards and parks. The importance of hedgerows was confirmed. Con- servation of M. schreibersii must be planned at a large scale, protecting a network of roosts desert 25 km WSW of Sayhut Al Mahra 15° 10‟ N, 51° 02‟ E 20 and promoting nature-friendly agricultural practices. Shuhayr Hadramaut 14° 41‟ N, 49° 24‟ E 40 Socotra, Faka Spring, Wadi Erher Hadramaut 12° 33‟ N, 54° 28‟ E 5 Key words: hunting activity, roosting behaviour, home range, urban areas, hedgerows, Socotra, Wadi Es Gego Hadramaut 12° 28‟ N, 54° 01‟ E 295 Schreibers’ bat, France

5 km S of Suq ad Dabab Taizz 13° 30‟ N, 43° 57‟ E 1305 RIASSUNTO - Attività e habitat di caccia di Miniopterus schreibersii (Chiroptera, Mi- Wadi „Adim, palmeria N of Sah Hadramaut 15° 41‟ N, 48° 52‟ E 730 niopteridae) nel sud della Francia: implicazioni per la sua conservazione. Conoscere Wadi Al Lahm, W of Al Mahwit Al Mahwit 15° 26‟ N, 43° 29‟ E 850 dell’ecologia comportamentale dei pipistrelli è essenziale per la loro conservazione. Il pro- gramma LIFE-Natura “Conservazione dei pipistrelli troglofili del sud della Francia”, della Wadi Daw‟an, palmeria S of Al Khuraybah Hadramaut 15° 09‟ N, 48° 26‟ E 1005 durata di quattro anni, ha incluso lo studio auto ecologico di Miniopterus schreibersii, una Wadi Dhahr, NW of Sana‟a Sana‟a 15° 27‟ N, 44° 10‟ E 2245 specie altamente gregaria. In una colonia di 3-5000 adulti, 21 femmine sono state seguite Wadi Maytam, 12 km SE of Ibb Ibb 13° 52‟ N, 44° 18‟ E 1615 con la radiotelemetria durante la gestazione e l’allattamento,quando alcune di esse hanno Wadi Zabid, SE of Al Mawqir Al Hudaydah 14° 10‟ N, 43° 30‟ E 270 utilizzato diversi roost nel raggio di 30 km dal sito riproduttivo. Ogni notte, per circa 6 ore, ciascun individuo ha utilizzato da 1 a 9 piccole aree di caccia a una distanza di 4,1 – 29,2 Wadi Zabid, W of Qaryat al Hasib Al Hudaydah 14° 09‟ N, 43° 31‟ E 300 km dal roost. Le dimensioni medie del home range sono variate da 10837 ha per le femmi- Zabid, citadel Al Hudaydah 14° 12‟ N, 43° 19‟ E 125 ne in gestazione a 22318 ha per quelle in allattamento. Le aree urbane illuminate da lam-

57 Aulagnier et al. pioni con luce bianca sono state selezionate come aree di foraggiamento, secondariamente anche le aree boschive frammentate e i frutteti delimitati da siepi. I progetti di conservazione del pipistrello di Schreibers devono essere pianificati su larga scala, proteggendo una rete sufficiente di rifugi e favorendo pratiche agricole con minor impatto sulla diversità ambientale.

Parole chiave: attività di caccia, uso dei rifugi, area vitale, aree urbane, siepi, pipistrello di Schreibers, Francia

DOI: 10.4404/Hystrix-22.1-4524

INTRODUCTION commuting routes and flight speed have been studied by Constant and Schreibers' bat (Miniopterus schreiber- Cannonge, then Serra-Cobo et al. sii Kuhl, 1817), a western Palaearctic (2000) confirmed the role of rivers as species (Appleton et al. 2004; Tian et possible landmarks in the orientation al. 2004; Bilgin et al. 2006), is listed as flight of the species. Using chemilumi- "near threatened" in IUCN red lists, at nescent tags, Barataud (1992) reported the global (IUCN 2010), European the first data on hunting behaviour in a (Temple and Terry 2007) and Mediter- wooded valley of central France. ranean levels (Temple and Cuttelod Schreibers' bats foraged in the upper 2009). Due to the mass mortality re- part of the valley, at the edge of the corded in 2002 in its south-western forest, at the canopy level. range (Roué and Némoz 2002; In contrast, radio-telemetry revealed Barataud and Précigout 2003; De Lucas intense foraging activity in urban areas 2007; Boléat et al. 2008), it has been as well as in broad-leaved woodlands, listed as "vulnerable" in France (UICN as far as 30 km from the roost (Lugon Comité Français 2009), Spain (De Lu- et al. 2004). As sample size was small, cas 2007) and Portugal (Cabral et al. these findings needed to be confirmed 2005). For example, in France, from and tentatively related to the hunting of 211,100 hibernating bats in 1995 (Roué Schreibers' bats for preferred prey and Groupe Chiroptères SFEPM 1997), items in eastern France, namely Lepi- only 70,950 were registered in 2004 doptera (73 to 96 % of volume through (Groupe Chiroptères SFEPM 2010). the seasons; Lugon 1998). Therefore, this cave-dwelling species For this purpose and for developing deserves a high priority for conserva- management guidelines for the forag- tion measures, including legal and ing areas, the four-year LIFE Nature physical roost protection and also programme "Conservation of three friendly management for foraging ar- cave-dwelling bats in Southern France" eas. included an autoecological study of While Schreibers' bats gather in large Schreibers' bat. By associating the ra- colonies in underground roosts that are dio-tracking of some females with diet then quite easy to locate, both their analysis, we investigated the habitat habitat use and spatial ecology are very choice of M. schreibersii from one poorly known. As far back as 1957, maternity colony in southeastern

58 Aulagnier et al. Foraging habitats of Miniopterus schreibersii pioni con luce bianca sono state selezionate come aree di foraggiamento, secondariamente France, at the edge of the Mediterra- tions dominate the landscape, including anche le aree boschive frammentate e i frutteti delimitati da siepi. I progetti di conservazio- nean region. We studied the pattern of mainly orchards and crops in the River ne del pipistrello di Schreibers devono essere pianificati su larga scala, proteggendo una nocturnal activity of bats and their use Rhône valley, and vineyards in the south- rete sufficiente di rifugi e favorendo pratiche agricole con minor impatto sulla diversità of space at both the individual and ern hills and the River Eygues valley. Hills ambientale. are covered with Mediterranean vegetation population levels, by identifying the of the holm-oak (Quercus ilex) and downy Parole chiave: attività di caccia, uso dei rifugi, area vitale, aree urbane, siepi, pipistrello di commuting routes, assessing home oak (Q. pubescens) series. Schreibers, Francia range areas, measuring distances from the roost to foraging sites and roost 1. Bat trapping, tagging and tracking DOI: 10.4404/Hystrix-22.1-4524 fidelity. Finally, we tried to identify the preferred foraging habitats after map- To reduce the impact on the colony and on INTRODUCTION commuting routes and flight speed ping their availability in the colony the individuals which rested during the have been studied by Constant and home range and analysing the diet of following diurnal period, bats were cap- Schreibers' bat (Miniopterus schreiber- Cannonge, then Serra-Cobo et al. the colony through the activity season. tured using a mistnet or a harp-trap as they sii Kuhl, 1817), a western Palaearctic (2000) confirmed the role of rivers as Radio-tracking was conducted during entered the roost. Each bat was identified species (Appleton et al. 2004; Tian et possible landmarks in the orientation and sexed, and both its forearm length and pregnancy and lactating periods, which body mass were measured. Adult females al. 2004; Bilgin et al. 2006), is listed as flight of the species. Using chemilumi- are the seasons of maximum energy "near threatened" in IUCN red lists, at nescent tags, Barataud (1992) reported were fitted with 0.64g, 0.80g or 0.86g (< demand for females (Racey and 6% body mass; Kenward, 1987) radio- the global (IUCN 2010), European the first data on hunting behaviour in a Speakman 1987, Kurta et al. 1990). transmitters (Holohil™, Biotrack™ and (Temple and Terry 2007) and Mediter- wooded valley of central France. Titley™), using surgical (Skinbond™) or ranean levels (Temple and Cuttelod Schreibers' bats foraged in the upper MATERIALS AND METHODS cosmetic adhesive (Duo Professional™), 2009). Due to the mass mortality re- part of the valley, at the edge of the and released in the tunnel after the adhesive corded in 2002 in its south-western forest, at the canopy level. From March to October a maternity colony got dry. We tagged 9 pregnant females in range (Roué and Némoz 2002; In contrast, radio-telemetry revealed May 2005 (3 the 14th, 1 the 19th, 3 the 22nd of Schreibers' bats (2500-3000 adults in th Barataud and Précigout 2003; De Lucas intense foraging activity in urban areas 2005, 4000-5000 in 2006) roosts in the and 2 the 25 ), and 12 lactating females in June 2006 (3 the 15th, 1 the 18th, 2 the 19th, 2007; Boléat et al. 2008), it has been as well as in broad-leaved woodlands, tunnel of Borie castle, located at Suze-land th listed as "vulnerable" in France (UICN as far as 30 km from the roost (Lugon Rousse (Drôme) in the south of the Rhône- 3 the 22 and 3 the 25 ). Transmitters detached after a few days. All procedures Comité Français 2009), Spain (De Lu- et al. 2004). As sample size was small, Alpes region (44°17’17”N, 4°50’17”E). They share the roost with about 600 greater were approved by the Ministry in charge of cas 2007) and Portugal (Cabral et al. these findings needed to be confirmed the Environment. 2005). For example, in France, from and tentatively related to the hunting of and lesser mouse-eared bats (Myotis myotis and M. bythii), and some Geoffroy's, Radio-tracking was carried out during two 211,100 hibernating bats in 1995 (Roué Schreibers' bats for preferred prey Bechstein's and Natterer's bats (M. emargi- weeks after the first captures by car and on and Groupe Chiroptères SFEPM 1997), items in eastern France, namely Lepi- natus, M. bechsteinii and M. nattereri, foot using 6 receivers (LA12Q AVM™; only 70,950 were registered in 2004 doptera (73 to 96 % of volume through respectively), as well as some lesser horse- Australis 26K Titley™) and either hand (Groupe Chiroptères SFEPM 2010). the seasons; Lugon 1998). shoe bats (Rhinolophus hipposideros). This held three-element Titley™ antennas or Therefore, this cave-dwelling species For this purpose and for developing tunnel, part of the Natura 2000 site FR four-element Televilt™ antennas. Each deserves a high priority for conserva- management guidelines for the forag- 8201676 "Sables du Tricastin", is burrowed night, from 21.00 to 7.00 (CEST - Central tion measures, including legal and ing areas, the four-year LIFE Nature in clayey limestone and sandstone, and European Summer Time), two to three physical roost protection and also programme "Conservation of three drains a swamp; water flows throughout the stationary teams located the bats every 5 min by triangulation from vantage points friendly management for foraging ar- cave-dwelling bats in Southern France" year. The surrounding area (named Tricastin) is (sometimes using a Null Peak AVM™ eas. included an autoecological study of an alluvial plain (rivers Rhône, Lez, Ou- antenna to improve the accuracy of the While Schreibers' bats gather in large Schreibers' bat. By associating the ra- vèze and Eygues) and low hills. The cli- direction of the signal), while the mobile colonies in underground roosts that are dio-tracking of some females with diet mate is Mediterranean, rainfall (750 mm operators determined bat locations mainly then quite easy to locate, both their analysis, we investigated the habitat per year) occurring irregularly through the with the "homing in" method (White and habitat use and spatial ecology are very choice of M. schreibersii from one year and rarely in summer, when the Mis- Garott 1990). All teams were co-ordinated poorly known. As far back as 1957, maternity colony in southeastern tral wind accentuates the drought. Cultiva- by walkie-talkie to ensure simultaneous

59 Aulagnier et al. bearings. In the field, bearings or locations 1981) of LOCATE II (NAMS, NSAC Can- were mapped on a 1/25.000© IGN map ada, 2000) and LOAS IV (Ecological implemented with a GPS grid. Only fixes Software Solutions, 2005) software. By this of active bats were used in habitat use method locations are calculated using, for analyses. Emergence and return times were each bearing, a fixed angular error, the also recorded. distance between the animal and the re- ceiver, and the angle between bearings. 2. Habitat categories This method provides a polygon when two bearings are available for one fix, and an In the area used by the colony, we identi- ellipse when three bearings have been re- fied 12 main habitat categories by field corded. surveys after both radio-tracking sessions: These locations and the homing in fixes 1. Deciduous or mixed woodlands frag- were stored in a Geographic Information mented by rivers, trails, roads or recently System (Arcview 3.2, ESRI California) and managed plots, which provide heterogene- superimposed to the SCAN25©IGN. Indi- ity and borders. vidual and colony home ranges were esti- 2. Closed and homogeneous deciduous or mated by the Minimum Convex Polygon mixed woodlands. (Mohr 1947) using the extension Animal 3. Heterogeneous pine woodlands includ- Movement (Hooge and Eichenlaub 2000) ing clearings and/or different storeys. of Arcview. Following Odum and Kuenzler 4. Dense pine woodlands. (1955), the minimum number of locations 5. Traditional orchards, parks or tree- for estimating the individual home range planted fallows. size was identified by calculating the mean 6. Pastures, meadows and scrublands de- of 100 bootstraps. Variation in home range limited by hedgerows or next to a wood- size between pregnant and lactating fe- land area. males was tested by a Mann-Whitney 7. Pastures, meadows and scrublands in an ranked test. open area. Foraging areas were identified by only the 8. Crops and vineyards delimited by hedge- homing in fixes. Habitat categories within a rows or next to a woodland area. 50 m radius circle were quantified (number of occurrences and percent area). Habitat 9. Crops and vineyards in an open area. selection relied on the comparison between 10. Urban areas with street lamps. used habitats (overall percent area of each 11. Urban areas without street lamps. habitat category within the circles) and 12. Lakes, swamps and rivers. available habitats (percent area of each Due to the very low occurrence of some habitat category within the colony home habitats (e.g. pine woods) or the difficulty range) using compositional analysis (Ae- in identifying them at the home range scale bischer et al. 1993). The unit of statistical (e.g. the presence of street lamps), to ana- independence was the individual tracked lyse habitat selection they were grouped bat. Following Almenar et al. (2006), to into five supra-categories: woodlands (1-4), assess the selection or rejection of each orchards and parks, open areas (6-9), urban habitat category, Bonferroni confidence areas (10-11), and water bodies. intervals were additionally calculated according to Neu et al. (1974). Analyses were 3. Data analysis computed with the animal movement extension (Hooge and Eichenlaub 2000) of Locations from triangulation and 95 % Arcview. error polygons were calculated using the The height of hedgerows was classified in Maximum Likelihood Estimator (Lenth four categories (0: no hedgerow, 1: hedge-

60 Aulagnier et al. Foraging habitats of Miniopterus schreibersii bearings. In the field, bearings or locations 1981) of LOCATE II (NAMS, NSAC Can- row < 2 m, 2: hedgerow between 2 and 10 2. Nocturnal activity were mapped on a 1/25.000© IGN map ada, 2000) and LOAS IV (Ecological m, 3: hedgerow > 10 m). The light colour implemented with a GPS grid. Only fixes Software Solutions, 2005) software. By this (white or orange) of street lamps at homing Pregnant females (14-28 May) emerged of active bats were used in habitat use method locations are calculated using, for in fixes was also recorded. An equal num- 59 ± 32 min after sunset and returned analyses. Emergence and return times were each bearing, a fixed angular error, the ber of locations, randomly sampled in the 140 ± 45 min before sunrise. Lactating distance between the animal and the re- also recorded. same proportion of habitat categories, were females (15-28 June) emerged 42 ± 17 ceiver, and the angle between bearings. described for subsequent comparisons. min after sunset and returned 113 ± 47 2. Habitat categories This method provides a polygon when two Significance of differences between the 2 min before sunrise. During all nights, bearings are available for one fix, and an two sets was checked by χ tests whenever In the area used by the colony, we identi- ellipse when three bearings have been re- applicable. Significance for all tests, com- no female returned temporarily to the fied 12 main habitat categories by field corded. puted by Minitab 12.2, was set at p < 0.05. roost. One lactating female even spent surveys after both radio-tracking sessions: These locations and the homing in fixes one day in a secondary roost, 19 km 1. Deciduous or mixed woodlands frag- were stored in a Geographic Information RESULTS distant from the colony, and came back System (Arcview 3.2, ESRI California) and mented by rivers, trails, roads or recently early on the following night. Total du- managed plots, which provide heterogene- superimposed to the SCAN25©IGN. Indi- 1. Radio-tracking data ration of activity was 333 ± 57 min for ity and borders. vidual and colony home ranges were esti- pregnant females, and 366 ± 38 min for mated by the Minimum Convex Polygon 2. Closed and homogeneous deciduous or lactating females (Fig. 1), the differ- mixed woodlands. (Mohr 1947) using the extension Animal The 21 females were tracked during Movement (Hooge and Eichenlaub 2000) ence being significant (Mann Whitney 3. Heterogeneous pine woodlands includ- only 2.4 ± 0.8 nights (maximum 4 W = 356, p = 0.029). ing clearings and/or different storeys. of Arcview. Following Odum and Kuenzler nights) due to the quick loss or break- (1955), the minimum number of locations 4. Dense pine woodlands. down of transmitters (Tab. 1). The 5. Traditional orchards, parks or tree- for estimating the individual home range 3. Home ranges and foraging areas size was identified by calculating the mean mean number of locations per female planted fallows. was 64.0 ± 61.3 (SD), the maximum 6. Pastures, meadows and scrublands de- of 100 bootstraps. Variation in home range Individual home ranges (Fig. 2) were limited by hedgerows or next to a wood- size between pregnant and lactating fe- number was 201, including 83 homing significantly larger for lactating fe- land area. males was tested by a Mann-Whitney in fixes. The minimum number of loca- males (22318 ± 7141 ha, n = 6) than for ranked test. 7. Pastures, meadows and scrublands in an tions for estimating individual home pregnant females (10837 ± 5399 ha, n open area. Foraging areas were identified by only the range size was obtained for 6 pregnant homing in fixes. Habitat categories within a = 6 Mann-Whitney W = 25 ; p = 8. Crops and vineyards delimited by hedge- and 6 lactating females. For this sam- 0.031). The smallest home range was rows or next to a woodland area. 50 m radius circle were quantified (number of occurrences and percent area). Habitat ple, the mean number of locations was 4465 ha for a pregnant female, and the 9. Crops and vineyards in an open area. selection relied on the comparison between 103.1 ± 50.3 (with no significant dif- 10. Urban areas with street lamps. largest was 29840 ha for a lactating used habitats (overall percent area of each ference between pregnant and lactating 11. Urban areas without street lamps. female (Fig. 3). habitat category within the circles) and 12. Lakes, swamps and rivers. females; Mann-Whitney W = 45.5, p = Commuting routes were very difficult available habitats (percent area of each Due to the very low occurrence of some 0.337). to identify due to the flight speed of the habitat category within the colony home Out of 17 bats tracked for more than bats and the large distance covered habitats (e.g. pine woods) or the difficulty range) using compositional analysis (Ae- in identifying them at the home range scale bischer et al. 1993). The unit of statistical one night, 6 pregnant and 1 lactating every night. The use of woodland bor- (e.g. the presence of street lamps), to ana- independence was the individual tracked females were located resting in diurnal ders, hedgerows; and riverine forests as lyse habitat selection they were grouped bat. Following Almenar et al. (2006), to roosts different from the Borie tunnel. landmarks suffered many exceptions, into five supra-categories: woodlands (1-4), assess the selection or rejection of each Two roosts were identified (under- including crossing a small mountain orchards and parks, open areas (6-9), urban habitat category, Bonferroni confidence ground transit roosts 15 and 19 km area. During windy nights, it was even areas (10-11), and water bodies. intervals were additionally calculated ac- distant from the colony, hosting mater- more difficult to locate the bats when cording to Neu et al. (1974). Analyses were nity colonies of greater and lesser commuting. Luckily they were very 3. Data analysis computed with the animal movement extension (Hooge and Eichenlaub 2000) of mouse-eared bats), and at least two faithful to their foraging areas, as re- Locations from triangulation and 95 % Arcview. other roosts were only located in a fa- vealed by the small number of distinct error polygons were calculated using the The height of hedgerows was classified in vourable area at a distance of 23 and 30 homing in fixes (Tab. 1), and it was Maximum Likelihood Estimator (Lenth four categories (0: no hedgerow, 1: hedge- km from the colony. then possible to find them after their

61 Aulagnier et al.

Table 1 - Tracking data and spatial use data of 21 female Schreibers' bats from the maternity colony of Suze-la-Rousse (France). * individuals reaching the minimum number of locations for estimating the home range size. Mean ± standard deviation.

Code No of No of No of No of dis- Maximum Maximum Convex nights locations homing-in tinct hom- distance (km) Polygon (ha) tracked ing-in 5-01 * 3 68 25 3 19.3 10550 5-02 2 11 0 0 - 4420 5-03 1 24 12 1 11.2 - 5-04 * 3 184 78 1 12.7 6009 5-05 * 3 91 30 3 18.5 4465 5-06 * 4 130 43 5 22.4 18000 5-07 * 2 54 22 9 15.9 9751 5-08 * 3 201 83 3 14.7 16250 5-09 2 29 12 1 16.6 2864 Pregnant 2.6 ± 0.9 88.0 ± 69.7 33.9 ± 29.1 2.9 ± 2.8 16.4 ± 3.6 9038.6 ± 5665.1 6-01 * 4 63 20 2 29.2 29840 6-02 2 2 0 0 - 358 6-03 3 15 0 0 - 3344 6-04 * 3 110 24 3 21.5 25330 6-05 * 3 78 0 0 - 24410 6-06 1 10 9 2 6.4 97 6-07 * 3 137 29 4 26.2 22230 6-08 2 1 0 0 - - 6-09 * 3 68 5 4 21.2 23340 6-10 * 1 53 5 2 20.8 8758 6-11 1 5 0 0 - 5987 6-12 2 10 0 0 - 6700 Lactating 2.3 ± 1.0 46.0 ± 46.1 7.7 ± 10.6 1.4 ± 1.6 20.9 ± 7.8 13672.1 ± 11310.9 outward journey which was usually Mean distance between two foraging longer than the return journey. A flight areas was 2896 ± 2834 m. The distance speed of 40-50 km/h was then re- between the maternity roost and forag- corded. ing areas (16.4 ± 5.7 km, N = 43, Fig. Foraging areas were identified for 8 4) was significantly larger for lactating pregnant and 6 lactating females. The females than for pregnant females number of foraging areas (2.0 ± 2.3) (Mann Whitney W = 550, p = 0.008). ranged from 1 (during 3 nights for a The maximum was 29.2 km and the pregnant female) to 9 (during two minimum was 4.1 km for two lactating nights for another pregnant female). females.

62 Aulagnier et al. Foraging habitats of Miniopterus schreibersii

Table 1 - Tracking data and spatial use data of 21 female Schreibers' bats from the mater- 500 nity colony of Suze-la-Rousse (France). * individuals reaching the minimum number of locations for estimating the home range size. Mean ± standard deviation.

400 Code No of No of No of No of dis- Maximum Maximum Convex nights locations homing-in tinct hom- distance (km) Polygon (ha) tracked ing-in 300

5-01 * 3 68 25 3 19.3 10550 (min) activity nocturnal of 5-02 2 11 0 0 - 4420 5-03 1 24 12 1 11.2 - 200 Duration 5-04 * 3 184 78 1 12.7 6009 5-05 * 3 91 30 3 18.5 4465 Pregnant (9) Lactating (12)

5-06 * 4 130 43 5 22.4 18000 Figure 1 - Nocturnal activity (total duration) of pregnant and lactating Schreibers' bat fe- 5-07 * 2 54 22 9 15.9 9751 males from the maternity colony of Suze-la-Rousse (France). 5-08 * 3 201 83 3 14.7 16250 5-09 2 29 12 1 16.6 2864 N Pregnant 2.6 ± 0.9 88.0 ± 69.7 33.9 ± 29.1 2.9 ± 2.8 16.4 ± 3.6 9038.6 ± 5665.1 6-01 * 4 63 20 2 29.2 29840 6-02 2 2 0 0 - 358 6-03 3 15 0 0 - 3344 6-04 * 3 110 24 3 21.5 25330 6-05 * 3 78 0 0 - 24410 6-06 1 10 9 2 6.4 97 6-07 * 3 137 29 4 26.2 22230

6-08 2 1 0 0 - - Maternity roost 6-09 * 3 68 5 4 21.2 23340 Roosts used during the study Other known roosts 6-10 * 1 53 5 2 20.8 8758 Pregnant female home range 6-11 1 5 0 0 - 5987 Lactating female home range 10 km 6-12 2 10 0 0 - 6700 Colony home range Lactating 2.3 ± 1.0 46.0 ± 46.1 7.7 ± 10.6 1.4 ± 1.6 20.9 ± 7.8 13672.1 ± 11310.9 Figure 2 - Roosts and home ranges (individual and colony) of pregnant and lactating Schreibers' bat females from the maternity colony of Suze-la-Rousse (France). outward journey which was usually Mean distance between two foraging The colony home range, evaluated by However some bats flew out of this longer than the return journey. A flight areas was 2896 ± 2834 m. The distance all radio-tracking data, was smaller for range and probably reached a distance speed of 40-50 km/h was then re- between the maternity roost and forag- pregnant females (89359 ha) than for of 32-35 km from the roost (according corded. ing areas (16.4 ± 5.7 km, N = 43, Fig. lactating females (162997 ha). The to imprecise locations). Foraging areas were identified for 8 4) was significantly larger for lactating maximum home range over the two pregnant and 6 lactating females. The females than for pregnant females periods was 168438 ha. The longest number of foraging areas (2.0 ± 2.3) (Mann Whitney W = 550, p = 0.008). 4. Habitat selection ranged from 1 (during 3 nights for a The maximum was 29.2 km and the distance precisely recorded from the pregnant female) to 9 (during two minimum was 4.1 km for two lactating roost was 29.7 km for a lactating fe- Urban areas were by far the most used nights for another pregnant female). females. male (24.7 km for a pregnant female). habitat category (54.0 %), followed

63 Aulagnier et al.

35000

30000

25000

20000

15000

10000

5000 Individual home(ha) rangesIndividual 0 Pregnant (6) Lactating (6)

Figure 3 - Home range areas of pregnant and lactating Schreibers' bat females from the maternity colony of Suze-la-Rousse (France).

and foraging (km) area and 5

0 Distance betwwen maternity roost maternity betwwenDistance Pregnant (27) Lactating (16)

Figure 4 - Flight distances between the roost and foraging areas of pregnant and lactating Schreibers' bat females from the maternity colony of Suze-la-Rousse (France).

Table 2 - Selection of habitats classified according to the main landscape categories by 14 female Schreibers' bats from the maternity colony of Suze-la-Rousse (France): compositional analysis (signs indicate whether the habitat placed in the corresponding row was more (+) or less (-) preferred than that in the corresponding column; + or -: non significant trends, +++ or ---: significant differences, p < 0.05).

Orchards and Open Urban Water Woodlands Rank parks areas areas bodies Woodlands + + --- + 3 Orchards and parks - + --- +++ 2 Open areas - - --- + 1 Urban areas +++ +++ +++ +++ 4 Water bodies ------0

64 Aulagnier et al. Foraging habitats of Miniopterus schreibersii

35000 Table 3 - Selection of habitats classified according to the main landscape categories by 14 female Schreibers' bats from the maternity colony of Suze-la-Rousse: Bonferroni confi- 30000 dence intervals.

25000 Bonferroni confidence Available 20000 Used intervals p-value proportion proportion Selection 15000 Lower Upper Woodlands 0.155 0.286 0.1394 0.1906 Negative p < 0.0001 10000 Orchards and parks 0.091 0.023 0.0718 0.1105 Positive p < 0.0001 5000 Open areas 0.198 0.626 0.1769 0.2331 Negative p < 0.0001 Individual home(ha) rangesIndividual 0 Urban areas 0.540 0.032 0.4898 0.5592 Positive p < 0.0001 Pregnant (6) Lactating (6) Water bodies 0.015 0.017 0.0077 0.0253 None

Figure 3 - Home range areas of pregnant and lactating Schreibers' bat females from the maternity colony of Suze-la-Rousse (France). 0% Fragmented deciduous woodlands 7% 41% Closed deciduous woodlands 17% 12% 29% Heterogeneous pine woodlands 35 0% 0% Dense pine woodlands 0% 30 Orchards and parks 9% 25 Pastures with hedgerows Pastures without hedgerows 20 3% 1% 2% 3% Cultures with hedgerows 15 0% Cultures without hedgerows 10% 9% 10 36% Lighted urban areas 4% 4% 6% and foraging (km) area and 5 5% 2% Dark urban areas Water bodies 0 Pregnant Lactating Distance betwwen maternity roost maternity betwwenDistance Pregnant (27) Lactating (16)

Figure 4 - Flight distances between the roost and foraging areas of pregnant and lactating Figure 5 - Habitat use of pregnant and lactating Schreibers' bat females from the maternity Schreibers' bat females from the maternity colony of Suze-la-Rousse (France). colony of Suze-la-Rousse (France): percentages of homing in areas in each habitat category. Table 2 - Selection of habitats classified according to the main landscape categories by 14 female Schreibers' bats from the maternity colony of Suze-la-Rousse (France): composi- by open areas (19.8 %), woodlands (15.5 woodlands, were negatively selected, tional analysis (signs indicate whether the habitat placed in the corresponding row was %), orchards and parks (9.1 %), and while the use of water bodies was op- more (+) or less (-) preferred than that in the corresponding column; + or -: non significant water bodies (1.5 %). According to portunistic. trends, +++ or ---: significant differences, p < 0.05). compositional analysis, habitats were At a finer scale, foraging areas were not used according to their availability mainly urban areas (with and without (Λ = 0.1318, χ² = 28.37, df = 4, p < street lamps; number of occurrences = Orchards and Open Urban Water Woodlands Rank 0.0001). Schreibers' bat used urban 31; percent area = 46) and (heterogene- parks areas areas bodies areas >>> woodlands >>> orchards and ous) deciduous or mixed woodlands (n Woodlands + + --- + 3 parks > open areas > water bodies = 19; %area = 22), followed by crops Orchards and parks - + --- +++ 2 (Tab. 2). Bonferroni confidence inter- and vineyards (n = 11; %area = 8), Open areas - - --- + 1 vals (χ² = 6.64, df = 1, p = 0.0012) con- pastures, meadows and scrublands (n = firmed the selection for urban areas and 8; %area = 4), delimited by hedgerows Urban areas +++ +++ +++ +++ 4 outlined a preference for orchards and or next to woodland, orchards and Water bodies ------0 parks (Tab. 3). Open areas, as well as parks (n = 7; %area = 9) and water bo-

65 Aulagnier et al. dies (n = 6; %area = 4). By contrast, no Homing in fixes included significantly fix was recorded in dense pine woods, more and higher hedgerows than those and very few were recorded in closed randomly sampled in the colony home and homogeneous deciduous or mixed range (97 % vs. 70 %, and 94 % vs. 60 woodlands (n =1; %area = 1). Most % respectively, χ² = 22.43; df = 3; p < foraging areas included more than two 0.0001; Fig. 6). habitat categories (81%), with one Finally, in urban areas, Schreibers' bats dominant habitat category (75%), ex- foraged preferably in lighted areas (χ² = cept for those in (heterogeneous) de- 17.19; df = 1; p < 0.0001), and exclu- ciduous or mixed woodlands and urban sively in areas lit by white street lamps areas with street lamps. (Fig. 7), whereas orange lamps prevail in the region (63%). Urban areas without street lamps and water bodies were used only by preg- DISCUSSION nant females; (heterogeneous) deciduous or mixed woodlands were more Whether the duration of nocturnal ac- used by lactating females, while crops tivity had been already recorded by and vineyards delimited by hedgerows direct observation at the entrance of or next to woodland were more used roosts, and foraging areas described by by pregnant females (Fig. 5). the detection of echolocation calls, tele-

Hedgerow height

No hedgerow < 2 m 2 – 10 m > 10 m

Homing in Random

Figure 6 - Habitat use of Schreibers' bat females from the maternity colony of Suze-la- Rousse (France): comparison of hedgerow height between random sites and homing in areas.

0% 27% orange lamps

white lamps 49% 51% 57% without street lamps 16%

Homing in Random

Figure 7 - Habitat use of Schreibers' bat females from the maternity colony of Suze-la- Rousse (France): comparison of lighting between random and homing in urban areas.

66 Aulagnier et al. Foraging habitats of Miniopterus schreibersii dies (n = 6; %area = 4). By contrast, no Homing in fixes included significantly metry revealed innovative results on Schreibers' bat in eastern France (Guil- fix was recorded in dense pine woods, more and higher hedgerows than those the behaviour of Schreibers’ bats. laume and Roué, 2006) revealed a and very few were recorded in closed randomly sampled in the colony home However, radio-tracking of Schreibers' maximum distance of only 10 km. In and homogeneous deciduous or mixed range (97 % vs. 70 %, and 94 % vs. 60 bats proved to be tricky. First, radio- contrast, with 11 tracked bats Lugon et woodlands (n =1; %area = 1). Most % respectively, χ² = 22.43; df = 3; p < transmitters were quickly lost or broke al. (2004) recorded a maximum dis- foraging areas included more than two 0.0001; Fig. 6). down, the maximum number of track- tance between roost and foraging areas habitat categories (81%), with one Finally, in urban areas, Schreibers' bats ing nights (N = 4) being much shorter of 29 km in a nearby area. This huge dominant habitat category (75%), ex- foraged preferably in lighted areas (χ² = than those reported for two other cave- dispersal around the roost, recorded in cept for those in (heterogeneous) de- 17.19; df = 1; p < 0.0001), and exclu- dwelling species, such as the Mediter- two very different French regions, may ciduous or mixed woodlands and urban sively in areas lit by white street lamps ranean horseshoe bat Rhinolophus eu- reduce intra-specific competition areas with street lamps. (Fig. 7), whereas orange lamps prevail ryale (N = 9; Goiti et al. 2003), and among the large number of bats in the in the region (63%). Urban areas without street lamps and long-fingered bat Myotis capaccinii (N colony. As suggested by Goiti et al. water bodies were used only by preg- = 15; Biscardi et al. 2007). Their loss (2006) for the Mediterranean horseshoe DISCUSSION nant females; (heterogeneous) decidu- was probably due to the highly gregari- bat, this hypothesis is supported by the ous behaviour of the species, including longer distances and larger home ous or mixed woodlands were more Whether the duration of nocturnal ac- used by lactating females, while crops intense rubbings and mutual grooming, ranges recorded for lactating females, tivity had been already recorded by and possible interactions with the whose energetic demand is maximum and vineyards delimited by hedgerows direct observation at the entrance of or next to woodland were more used mouse-eared bats sharing the roost, as (Racey and Speakman 1987) and flight roosts, and foraging areas described by most recovered radiotransmitters were ability cleared of the embryo's body by pregnant females (Fig. 5). the detection of echolocation calls, tele- nibbled. Second, some females mass (Norberg and Rayner 1987). switched roost, even during the mater- Mean time of emergence (ca. 50 min Hedgerow height nity period, although they are known after sunset) was in agreement with for their attachment to the maternity previous records for this time of the roost (Rodrigues et al. 2010). Third, the year (Topál in Boye 2004), however No hedgerow high speed of the flight of this species, Schreibers' bats started to emerge < 2 m already recorded by Constant and Can- sooner than reported by Constant and 2 – 10 m nonge (1957) was confirmed. Schreib- Cannonge (1957). Contrary to most > 10 m ers' bats were reported to fly usually species (Racey 1982; Henry et al. 2002; between 5 and 10 m above ground, up Russo et al. 2002), maternal care does Homing in Random to the tree top level, but, mainly during not occur during the night, and hardly Figure 6 - Habitat use of Schreibers' bat females from the maternity colony of Suze-la- windy nights, they may shelter close to any resting period was recorded. A Rousse (France): comparison of hedgerow height between random sites and homing in woodland borders and hedgerows, similar result was obtained in eastern areas. where they are more difficult to locate France (Lugon et al. 2004) for 10 fe- (Weid and Von Helversen 1987; Lugon males. The total duration of activity 0% 1999; Lugon and Roué 2002). Fourth, was even longer during the lactating 27% the distances between diurnal roosts, as period, when the night is shorter. So orange lamps well as those between the maternity females probably balance the need for white lamps 49% roost and foraging areas reached more feeding far away from the roost and 51% 57% than 30 km, while the maximum re- reproduction constraints, reaching the without street lamps 16% corded distance was 15 km for the maximum during lactation (Racey and long-fingered bat (Biscardi et al. 2007) Speakman 1987). At the end of July, Homing in Random and 9 km for the Mediterranean horse- when the young-of-the-year are already

Figure 7 - Habitat use of Schreibers' bat females from the maternity colony of Suze-la- shoe bat (Goiti et al. 2006). A prelimi- flying, Barataud (1992) recorded some Rousse (France): comparison of lighting between random and homing in urban areas. nary four-night study conducted with returns to the roost after two hours,

67 Aulagnier et al. without distinguishing between adults The importance of woodland bor- and young. ders and hedgerows (Constant and Defining home ranges for such a long- Cannonge 1957; Barataud 1992; distance moving species is quite con- Lugon and Roué 2002) was con- fusing as Schreibers' bats usually for- firmed by our results. In open areas age on very small areas (e.g. 50 m of a street or hedgerow). Variation in the (crops, vineyards, pastures, mead- size of Minimum Convex Polygon ows and scrublands), hedgerows or mainly depended on both the distance woodland borders over 2 m high between foraging areas and the roost, were significantly chosen by bats and the number of foraging areas. De- for foraging. The vertical structure fining home range sizes using the Ker- of vegetation was also a determinant nel method (Worton 1989) would not in woodlands, where Schreibers' provide a good estimate of the spatial bats favoured fragmented deciduous ecology of a species which uses such a plots. In eastern France, 74.6 % of small part of its range as foraging ar- locations in forests were recorded eas. So we analysed habitat use by Schreibers' bats by the field description along roads and trails (Lugon et al. of homing in fixes. 2004). Deciduous woodlands were Confirming the results of Lugon et al. negatively selected in our study, a re- (2004), urban areas were the main for- sult mainly due to pregnant females aging habitats, and lactating females which possibly did not find substantial preferred lii areas, as numerous bat food supply there in May 2005. In species do in Europe and North Amer- June 2006, lactating females bene- ica (Rydell and Racey 1995). Further- fited from an outbreak of Asian more, Schreibers' bats only foraged in gypsy moth (Lymantria dispar) in areas lit by white street lamps. After holm-oak woodlands, and were Rydell (1992), it is now well known probably attracted by this abundant that the bluish-white light of mercury- resource. Contrary to pregnant fe- vapour street lamps, which emit ultra- males in May 2005, they neglected violet radiation, attracts insects, including moths, optimising energy gains by riverine forests, a habitat used in bats. In contrast, low-pressure sodium southern Italy (Russo and Jones lamps, which emit monochromatic 2003) and possibly offering an ear- orange light, do not attract insects lier availability of prey. (Racey 1998). For northern bats, Rydell (1992) reported a gross energy intake CONCLUSIONS more than twice higher for bats foraging around street lamps (0.5 kJ / min) The foraging behaviour of Schreibers' than in woodland (0.2 kJ / min). Arlet- bat proved to be different from most of taz et al. (1998) suggested that this insectivorous bat species. Although the foraging strategy could influence re- common pipistrelle (Pipistrellus pipi- production success, and then be se- strellus), Kuhl's pipistrelle (P. kuhlii), lected for. as well as the northern bat and the

68 Aulagnier et al. Foraging habitats of Miniopterus schreibersii without distinguishing between adults The importance of woodland bor- common serotine (Eptesicus serotinus), scher, Yoann Peyrard, Thomas Deana, and young. ders and hedgerows (Constant and were reported to preferably forage in Alexandre Bringard, Frédéric Manalt, Max Lucet, Aurélie Foucher, Jean-Loup Defining home ranges for such a long- Cannonge 1957; Barataud 1992; urban areas lighted by white street distance moving species is quite con- Firmery, Laura Defreine, Claire Planchon, Lugon and Roué 2002) was con- lamps (Haffner and Stutz 1985-1986; Vincent Foray, and 33 other radio-tracking fusing as Schreibers' bats usually for- firmed by our results. In open areas Rydell 1992; Blake et al. 1994; Catto et participants. It was part of the LIFE-Nature age on very small areas (e.g. 50 m of a al. 1995), none of these species forage (crops, vineyards, pastures, mead- programme "Conservation of three cave- street or hedgerow). Variation in the 30 km away from their daily roost. dwelling bats in Southern France" size of Minimum Convex Polygon ows and scrublands), hedgerows or Another unlikely feature, according to (LIFE04NAT/FR/000080), also supported mainly depended on both the distance woodland borders over 2 m high Henry et al. (2002), was the continuous by the DIREN Rhône-Alpes, the Conseil between foraging areas and the roost, were significantly chosen by bats overnight foraging activity, even during Régional Rhône-Alpes, the Drôme depart- and the number of foraging areas. De- for foraging. The vertical structure ment, the Nature and Découvertes and the lactating period, and the roost switches. MAVA foundations. fining home range sizes using the Ker- of vegetation was also a determinant Long-distance foraging flights imply nel method (Worton 1989) would not in woodlands, where Schreibers' that conservation measures for REFERENCES provide a good estimate of the spatial bats favoured fragmented deciduous Schreibers' bat must be planned at a ecology of a species which uses such a plots. In eastern France, 74.6 % of large scale. Efforts should focus on the Aebischer N.J., Roberston P.A., Kenward small part of its range as foraging ar- locations in forests were recorded protection of a network of roosts, keep- R.E. 1993. Compositional analysis of eas. So we analysed habitat use by habitat use from animal radio-tracking along roads and trails (Lugon et al. ing in mind that this species is reluctant Schreibers' bats by the field description to enter grilled sites (Moeschler 1995), data. Ecology 74(5): 1313-1325. 2004). Deciduous woodlands were of homing in fixes. and also the improvement of hedge- Alcalde J.T. 2003. Impacto de los parques negatively selected in our study, a re- eólicos sobre las poblaciones de mur- Confirming the results of Lugon et al. rows, so as to assist commuting and sult mainly due to pregnant females ciélagos. Barbastella 2: 3-6. (2004), urban areas were the main for- foraging areas and connectivity be- which possibly did not find substantial Almenar D., Aihartza J., Goiti U., Salsa- aging habitats, and lactating females tween favourable habitats and the pro- mendi E., Garin I. 2006. Habitat selec- preferred lii areas, as numerous bat food supply there in May 2005. In motion of nature-friendly agricultural tion and spatial use by the trawling bat species do in Europe and North Amer- June 2006, lactating females bene- practices instead of massive pesticide Myotis capaccinii (Bonaparte, 1837). ica (Rydell and Racey 1995). Further- fited from an outbreak of Asian spread. The tendency to replace mer- Acta Chiropterol. 8(1): 157-167. Appleton B.R., McKenzie J.A., Christidis more, Schreibers' bats only foraged in gypsy moth (Lymantria dispar) in cury vapour lamps with sodium lamps, areas lit by white street lamps. After L. 2004. Molecular systematics and holm-oak woodlands, and were which use less energy but do not attract biogeography of the bent-wing bat Rydell (1992), it is now well known probably attracted by this abundant insects, is unfavourable to Schreibers’ complex Miniopterus schreibersii that the bluish-white light of mercury- resource. Contrary to pregnant fe- bat. On the other hand, the former at- (Kuhl, 1817) (Chiroptera: Vespertili- vapour street lamps, which emit ultra- males in May 2005, they neglected tract moths, that then become unavail- onidae). Mol. Phyl. Evol. 31(2): 431- violet radiation, attracts insects, includ- 439. riverine forests, a habitat used in able to gleaning bat species (Racey ing moths, optimising energy gains by 1998), and possibly concentrate bats Arlettaz R., Berthoud G., Desfayes M. southern Italy (Russo and Jones 1998. Tendance démographiques op- bats. In contrast, low-pressure sodium into relatively small risky areas. Due to 2003) and possibly offering an ear- posées chez deux espèces sympa- lamps, which emit monochromatic its high speed flight, Schreibers' bat triques de chauves-souris, Rhinolophus orange light, do not attract insects lier availability of prey. might suffer from the impact of wind hipposideros et Pipistrellus pipistrel- (Racey 1998). For northern bats, Rydell farms (Alcalde 2003). Mitigation and / lus : un possible lien de cause à effet. (1992) reported a gross energy intake CONCLUSIONS or avoidance measures should be a Rhinolophe 13: 35-41. more than twice higher for bats forag- priority for stakeholders. Barataud M. 1992. L'activité crépusculaire et nocturne de 18 espèces de Chirop- ing around street lamps (0.5 kJ / min) The foraging behaviour of Schreibers' than in woodland (0.2 kJ / min). Arlet- bat proved to be different from most of tères, révélée par marquage lumines- ACKNOWLEDGEMENTS cent et suivi acoustique. Rhinolophe, taz et al. (1998) suggested that this insectivorous bat species. Although the 9: 33-57. foraging strategy could influence re- common pipistrelle (Pipistrellus pipi- This study was made possible thanks to the Barataud M., Précigout L. 2003. Drama- production success, and then be se- strellus), Kuhl's pipistrelle (P. kuhlii), kind permission of the tunnel's owners and tique déclin des minioptères sur les lected for. as well as the northern bat and the all the bat workers, including Robin Let- sites d'hibernation. Plecotus 13: 9.

69 Aulagnier et al.

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