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Habitat selection and spatial use by the trawling bat Myotiscapaccinii (Bonaparte 1837)Author(s) David Almenar Joserra Aihartza Urtzi Goiti Egoitz Salsamendi and Inazio GarinSource Acta Chiropterologica 8(1)157-167 2006Published By Museum and Institute of Zoology Polish Academy of SciencesDOI httpdxdoiorg1031611733-5329(2006)8[157HSASUB]20CO2URL httpwwwbiooneorgdoifull1031611733-53292820062985B1573AHSASUB5D20CO3B2

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INTRODUCTION

Several bats species of the genus Myotisare referred to in the literature as lsquotrawlingMyotisrsquo or Myotis that forage over water(Siemers et al 2001 Fenton and Bogda-nowicz 2002) Formerly referred to as sub-genus Leuconoeuml (Findley 1972) this groupseems to be paraphylletic and its con-stituents share a series of morphologicalconvergences (Ruedi and Mayer 2001Fenton and Bogdanowicz 2002) such aslarge hind feet large wingspan and pointedwings (Findley 1972 Norberg and Ray-ner 1987) The species within this group

typically hunt low over water often gaffingpreys from the waterrsquos surface using theirlarge feet and uropatagium (Jones andRayner 1988 1991 Britton et al 1997Fenton and Bogdanowicz 2002) Typicalpreys are water-associated arthropods (Brit-ton et al 1997 Law and Urquhart 2000Flavin et al 2001) although some of thetrawling Myotis species also prey on fish(Robson 1984 Aihartza et al 2003 Ma etal 2003) Hunting on aquatic habitatscould be advantageous to bats in terms offoraging economy as insects usually swarmover water The reduced flight cost resultingfrom the ground effect (Aldridge 1988

Acta Chiropterologica 8(1) 157ndash167 2006PL ISSN 1508-1109 copy Museum and Institute of Zoology PAS

Habitat selection and spatial use by the trawling bat Myotis capaccinii (Bonaparte 1837)

DAVID ALMENAR1 2 JOSERRA AIHARTZA2 URTZI GOITI2 EGOITZ SALSAMENDI1 2and INAZIO GARIN2 3

1Estacioacuten Bioloacutegica de DoZana (CSIC) Avda Mariacutea Luisa sn Pabelloacuten del Peruacute 41013 Sevilla Spain2Zoologia eta Animali Zelulen Biologia Saila UPVEHU 644 PK 48080 Bilbao The Basque Country

3Corresponding author E-mail inaziogarinehues

Habitat selection and spatial use was studied in a population of Myotis capaccinii (Bonaparte 1837) in theeastern Iberian Peninsula during the spring of 2004 The radio-tracked bats used only aquatic habitats asforaging sites and most foraging activity concentrated on rivers Rivers were positively selected and showedthe highest preference rank Pools were also positively selected but only a single pool was used through thetracking period Foraging was not evenly distributed along rivers The features of the water surface furtherdetermined habitat selection Open waters with smooth surfaces were selected over cluttered surfaces or waterscompletely covered by vegetation This microhabitat preference is thought to be due to a greater efficiency inprey detection and capture over open calm waters Nonetheless the extremely high aggregation of foragingindividuals observed suggests that the distribution of prey might also affect the location of foraging sites alongrivers Thus conservation management of M capaccinii should ensure protection of low-flowing or stagnantwaters in rivers around the batsrsquo main caves

Key words Myotis capaccinii trawling bats radio-telemetry habitat selection spatial ecology Mediterranean

Rayner 1991) is also an advantage for a batflying low over water In addition trawlingbat species usually select calm and openwaters (Von Frenckell and Barclay 1987Boonman et al 1998 Rydell et al 1999Warren et al 2000) where target detectionmay be enhanced due to the smooth sur-faces (Boonman et al 1998 Rydell et al1999 Siemers et al 2001 2005) and re-duced sonic interferences (Von Frenckelland Barclay 1987 Mackey and Barclay1989) Nevertheless several species oftrawling bats are known to alternate theirforaging between aquatic and terrestrialhabitats (Jones and Rayner 1988 1991 Ar-nold et al 1998 Kalko et al 1998 Fentonand Bogdanowicz 2002)

Myotis capaccinii (Bonaparte 1837) is a Mediterranean troglophylous medium-sized species classed among the trawlingbats (Guilleacuten 1999 Spitzenberger and Hel-versen 2001) Traditionally it has beenlinked to inland waters and several obser-vations of hunting over water (usually calm)have been described (Ahleacuten 1990 Kalko1990 Spitzenberger and Helversen 2001)Although in Europe it is widely consideredan endangered species (Hutson et al 2001)there is little knowledge about its ecol-ogy similarly to other Mediterranean bats(Russo and Jones 2003) The large hind feetand pointed wings of M capaccinii are wellsuited to capture prey while trawlingthrough the water surface This species isknown to consume fish in the wild (Aihar-tza et al 2003)

Data published to date do not agree onthe relative use or relevance of terrestrialand aquatic environments in M capacciniiIn Corsica Courtois (1998) captured thespecies only on aquatic habitats althoughthe mist-nets used in trapping were not allo-cated in proportion to habitat availabilityCarmel and Safriel (1998) and Russo andJones (2003) performed multispecific re-search on the use of foraging habitat by

means of acoustic surveys in Israel andItaly respectively In both studies theaquatic habitats were the most frequentlyused by M capaccinii although some con-tacts were also made over terrestrial habitats(see also Cosson and Meacutedard 1999 Lanzaand Agnelli 2000 Spitzenberger and Hel-versen 2001) Given the limitations of themethods utilized these results are far fromconclusive no study has continuously sur-veyed individual bats or systematically re-corded the habitats involved in each kind ofbat activity

We expected an intensive use of aquaticenvironments by M capaccinii according toprevious data on its habitat use morpholo-gy and similarities to other trawling batsSpecifically we tested two predictions 1)terrestrial habitats if used will be not se-lected for 2) distribution and size of forag-ing sites will be linked to the availabilityand features of aquatic habitats We used radio-telemetry to determine 1) the degreeof use of terrestrial habitats 2) the use andselection of aquatic habitats at both themacrohabitat (landscape) and microhabitat(foraging site) level and 3) size and spatialdistribution of foraging sites To our knowl-edge no species-specific research on habi-tat selection and spatial use of M capacciniihas yet been published

MATERIALS AND METHODS

Study Area

The study area is located in the eastern IberianPeninsula in the province of ValPncia This region ischaracterised by a Mediterranean climate (Sanchis-Moll 1988) It is located in the low basin of theXuacutequer River where the river valley forms a largeflood plain The basin includes several tributarieswith varying degrees of steepness the most importantbeing the Albaida and Sellent rivers Flow of tributar-ies shows some degree of Mediterranean seasonalityThe altitude range is 20ndash600 m asl Riverbanks are steep in most of their courses Riverine vegetationis dominated by reeds (Arundo donax) and ripar-ian woodland is scarce Hills and slopes are mainly

158 D Almenar J Aihartza U Goiti E Salsamendi and I Garin

covered by Mediterranean scrubland and valleys andplains are dominated by orange groves As orangeagriculture demands irrigation water is highly man-aged through a big dam (called lsquoPresa de Tousrsquo)many lsquoassutsrsquo (the local name for a kind of small res-ervoir in the rivers) a vast network of irrigation ca-nals (05ndash8 m wide) and several ponds (about 3km2)Considering that it is a rural area the human popula-tion density is relatively high (142 inhabitantskm2 in2003 mdash source Instituto Nacional de Estadiacutestica)

Bat Population and Roosts

There are several caves in the area but only twoare important as bat roosts and inhabited by largegroups of M capaccinii during the nursery periodThe main roost is a limestone cave with more than500 individuals in June and about 150 individualsduring the study period (April 2004) The second a limestone chasm 84 km from the first one is alsoknown to be used from spring to autumn but thenumber of M capaccinii there is uncertain Bothroosts are shared with other cave-dwelling bats suchas Rhinolophus euryale R mehelyi Myotis blythii M myotis M emarginatus M nattereri and Minio-pterus schreibersii (D Almenar unpubl data)

Trapping Tagging and Data Collection

We radio-tagged 10 male and 10 female M ca-paccinii captured with a harp trap (Kunz and Kurta1988) on the 9th and 17th of April 2004 while enter-ing the main cave Forearm length body mass andsex were recorded for each individual After clippingthe fur between the batrsquos shoulder blades we attacheda 045 g PIP II radio-transmitter (Biotrack Ltd Dor-set UK) with Skinbond surgical adhesive (Smith andNephew Largo Florida USA) The transmitter repre-sented less than 5 of the batrsquos body mass (Aldridgeand Brigham 1988) except for four cases in whichthey were up to 57 Capture manipulation and tag-ging were performed with permission of the Valen-cian Government

The bats were tracked between the 10th and 30thof April from emergence to return They were trackedby car and on foot using up to six radio-receivers(five units were model 1000-XRS Wildlife MaterialsCarbondale USA and one was model FT-290RII AWagener Telemetrieanlagen Koumlln Germany) withthree-element Yaggi antennas Two to three bats weretracked each night simultaneously by teams of two tothree people Locations were obtained by homing-in(White and Garrot 1990) The batsrsquo time spent andactivity type in each location were recorded Activity

was classified as commuting when direct and quickdisplacements between roost and foraging sites or between two separate foraging locations occurr-ed Motionless usually weak signals coming from a shelter were classified as resting Periodic move-ments over a location were recorded as foraging Onlyforaging locations were analysed Foraging locationswere mapped in the field on orthoimages or 110000topographic maps and transferred afterward into a Geographical Information System (GIS mdash ArcView32 ESRI California USA)

Habitat Availability and Use

Photointerpretation of orthoimages (Institut Geo-grBfic ValenciB Valencia Spain) at 4 pixelsm resolu-tion with Arcview 32 GIS generated habitat mapswithin the Minimum Convex Polygon (MCP) en-compassing all radio-tracking locations and dailyroosts (White and Garrot 1990) First we distin-guished between terrestrial and aquatic habitatsSecondly three habitat categories were distinguishedwithin inland waters 1) rivers considering all the nat-ural watercourses with flowing water including smallreservoirs along them 2) canals considering onlythose not covered and more than 2 m wide 3) pondsconsidering only those with surface area greater than100 m2 Finally rivers were classified according tophysical features of water surface a) smooth definedas stagnant or slow flowing waters with smooth sur-face b) cluttered defined as water surfaces rippled orcluttered due to turbulence c) covered defined as wa-ter surfaces completely covered by vegetation (main-ly A donax) This mapping of habitats was checked inthe field River stretches were divided into homoge-neous sectors following the previous criteria This in-formation was transferred into GIS and the summedarea of each habitat category was estimated for thewhole MCP Availability of river water surfaces withdifferent physical features was estimated as thesummed area of each class in each individual foragingsite defined as the whole river sector between the twomost distant foraging locations

The bat location map was crossed with availablehabitats using Arcview 32 GIS to associate each fixwith the corresponding habitat category or river fea-ture class Foraging time spent by each bat on eachhabitat andor feature class was calculated

Habitat Selection

We used the methods proposed by Aebischer et al (1993) and Neu et al (1974) to analyse habitatselection The use of both is justified because they

Habitat of Myotis capaccinii 159

offer complementary information the former ranksthe relative preferences for habitatsfeatures and thelatter provides the significance of the selection ofeach category The method proposed by Aebischer et al (1993) is based on compositional analysis Itwas applied both to test the hypothesis of non-randomuse and to assess the rank of relative preference forhabitatsfeatures To assess the selection or rejectionof the habitat or river feature categories we used themethod proposed by Neu et al (1974) based on theestimation of Baileyrsquos simultaneous confidence inter-vals (Cherry 1996)

Compositional method was carried out usingCompos Analysis 51 (Smith Ecology Ltd Aberga-venny UK) Significance of Wilkrsquos and t-statisticswas obtained by randomisation with 1000 iterationsWhenever the use of a category was 0 it was re-placed by 0001 Because some categories were un-available for some bats we calculated the Wilkrsquoslambda as a weighted mean Neu et alrsquos method was carried out using Resource Selection for Win-dows (F Leban University of Idaho Moscow USA)For this test the time spent on each category wastransformed into the number of contacts consideringa minimum interval of 10 minutes to avoid autocorre-lation Confidence intervals for categories with lessthan five contacts were not taken into account (Man-ly et al 2002) For all tests significance was set at P lt 005 The confidence intervals were computed atthe 95 level

Spatial Use

In the case of linear elements such as rivers orcanals individual foraging sites were measured bylength and area of used stretches defined as the wholeriver or canal sector between the two most distant for-aging locations When used the length and area ofponds were added to these values Whenever the lastday roost was known we measured the minimum lin-ear distance between each foraging location and thatroost The individual foraging time in minutes wasused as an estimation of tracking effort to test the in-dependence of the measured variables from effortSquare-root transformation was applied to variablesto fit with test assumptions

RESULTS

We obtained usable tracking informa-tion for 10 females and seven males to-talling 7938 minutes of foraging time(Table 1) Tracking time was shorter thanexpected due to practical difficulties relatedto the physiography of the rivers and land-scape features The steep banks and the fre-quent changes in direction of the rivers aswell as the scarcity of vantage points overthem challenged continuous radio-tracking

160 D Almenar J Aihartza U Goiti E Salsamendi and I Garin

Body FAL Tracking TTF Foraging site DistanceCode Sex mass (g) (mm) nights (min) Lenght (m) Area (ha) Maximum 0 SD

611 X ndash ndash 4 380 4148 440 2504 781 1100628 X 79 415 4 272 2613 280 2265 1181 592640 Y ndash 409 4 406 1565 533 8840 7750 2097677 Y ndash 420 8 1217 3185 1137 9899 835 2341695 X 96 416 5 498 1165 387 600 331 194816 X 90 413 2 425 174 035 ndash ndash ndash832 X 85 424 4 1270 1623 534 957 299 212920 Y 92 409 7 939 3425 372 8488 7859 419930 Y 82 414 6 717 2785 1020 9669 2095 3601115 X 99 409 3 454 496 060 7692 7535 97143 Y 99 411 3 372 3922 411 9207 8504 652184 X 100 419 1 157 1592 540 8762 8460 196315 Y 100 418 2 115 539 050 1530 1400 121399 X 99 420 1 58 432 101 ndash ndash ndash539 Y 99 400 5 465 4184 452 9172 8613 518588 Y 99 416 1 151 941 290 8762 8653 154789 Y 98 414 1 42 100 040 8544 8544 ndash

TABLE 1 Sex body mass forearm length (FAL) tracking effort (tracking nights and TTF = tracking time of foraging activity) length and area of foraging sites and average and maximum travelling distances of 17 radio-tracked M capaccinii in the Xuacutequer River basin (Iberian Peninsula)

of most of the individuals Furthermore toensure that the observed locations were notbiased toward those sites most easily acces-sible once the signal of a target individualwas lost we searched for it thoroughly overthe whole study area This prevented biasedresults but further reduced the number of observed locations On average individualswere located during 36 nights (range 1ndash8)recording an average of 467 minutes of for-aging activity (range 42ndash1270)

Habitat Use and Selection

The MCP covered 745 km2 with an al-titude range of 25ndash417 m asl (see Fig 1)None of the bats was located foraging onterrestrial habitats All the foraging activityconcentrated on aquatic environments Thetracked bats spent almost all their foragingtime over rivers (943 unweighted accu-mulated foraging time) Three bats also foraged in a pond next to a river (55) A canal was also used (12) where onlyone bat foraged for several minutes per-forming some kind of lsquoforaging while com-mutingrsquo behaviour given that this bat alsofollowed the canal to reach the foraging

place in the river where it spent the rest ofthe night The use of aquatic habitats was non-random (compositional analysisweighted mean Wilkrsquos λ = 0036 Neu etalrsquos method G = 20154 P lt 0001 for bothcomparisons) Rivers and ponds were posi-tively selected Rivers were significantlypreferred over the rest of habitats whileponds were significantly preferred to canals(Tables 2 and 3)

Concerning river features the usedstretches were mainly stagnant or had slow-flowing waters eg small reservoirs orpools usually containing eutrophic andcloudy water Bats spent most of their timeover river stretches with smooth surfaces(847 unweighted accumulated foragingtime over rivers) although some activitywas recorded over rippled waters (147)or surfaces covered by vegetation (05)Use of each class was non-random (com-positional analysis weighted mean Wilkrsquos λ = 0268 Neu et alrsquos method G = 6830for both P lt 001) Smooth surfaces werepositively and significantly selected overthe rest of the features whereas clutter-ed and covered surfaces were negatively selected (Tables 2 and 3) Differences

Habitat of Myotis capaccinii 161

FIG 1 Map of the study area showing the aggregation of the foraging locations Rivers and canals are drawn ingrey (canals as dashed lines) Black dots represent foraging locations The asterisk represents the main cave

and the triangle the secondary cave The arrow shows the direction of the water flow

between cluttered and covered surfacescould not be tested by randomisation be-cause the degrees of freedom were less thanfive (Aebischer et al 1993) due to somelsquozerorsquo values of availability

Spatial Use

After emergence bats moved directly tothe foraging places Bats commuted overterrestrial habitats rivers or canals Al-though the availability of rivers exceeded26 km in length within the MCP all the for-aging activity concentrated in two collectiveforaging sectors mdash 44 km and 33 km longrespectively mdash situated 11 km apart fromeach other The use of spotlights revealed a high density of bats hunting in the forag-ing sites within these sectors whereas notrawling bat activity was observed in othersectors of the rivers Moreover all but oneof the tagged bats foraged exclusively inone of the two foraging sectors during thestudy period Within these sectors mostbats repeatedly changed the foraging siteswhere they flew upward and downwardfrom minutes to hours

This aggregation was not determined bylow availability of suitable foraging habi-tats Calm open waters amounted to 62 of the available river surfaces and these

two sectors comprised only 19 of thesesmooth surfaces

Between individual bats size of forag-ing sites varied from 017 to 418 km inlength (0 SD 194 145) and from 035 to 1137 ha in surface (393 317) Neithervariable was independent from tracking effort (Pearsonrsquos correlation coefficientr = 063 and r = 048 for both n = 17 and P lt 001)

On average the bats moved 37 km (SD = 38) each night to their foraging sitesThe farthest foraging location reached by an individual in a single night was 99 km fromits roost

DISCUSSION

Our results pertaining to habitat selec-tion show that M capaccinii is specialisedin feeding over water as previous studieshave indicated (Kalko 1990 Meacutedard andGuibert 1990 Carmel and Safriel 1998Courtois 1998 Cosson and Meacutedard 1999Spitzenberger and Helversen 2001 Russoand Jones 2003) The foraging activity inour study occurred over aquatic habitatswhereas use of terrestrial habitats waslinked to commuting Carmel and Safriel(1998) and Russo and Jones (2003) founda slightly higher use of terrestrial habitatthan we did although their study method

162 D Almenar J Aihartza U Goiti E Salsamendi and I Garin

Category 0 plusmn SE t-statistic df P-value Preference rankHabitats

River versus Pool 60 plusmn 100 603 16 0001 RivergtgtgtPoolgtgtgtCanalRiver versus Canal 90 plusmn 046 1951 16 0001Pool versus Canal 29 plusmn 120 262 16 001

River water surfaceSmooth versus Rippled 16 plusmn 068 231 14 001 SmoothgtgtgtRippledgtCoveredSmooth versus Covered 34 plusmn 159 211 4 ndashRippled versus Covered 17 plusmn 183 090 4 ndash

TABLE 2 Relative selection of habitat and physical features of rivers by 17 M capaccinii in spring in the XuacutequerRiver basin following compositional analysis Positive comparisons (indicating preference of numerator versusdenominator) are shown with the mean and SE values for log-ratio differences The t-statistic degrees offreedom and P-values estimated by means of randomisation are also presented Rankings of preferredcategories are provided in the last column Each category written before gt is preferred to those following thesymbol The symbol gtgtgt shows significant differences in preference

did not allow the identification of the kindof activity involved

Another selection pattern cannot be dis-carded where the population status or habi-tat availability are different The high popu-lation size of M capaccinii in the study areasuggests that the habitat quality is equallyhigh (sensu Garshelis 2000) In suboptimalhabitat conditions the species would possi-bly be forced to change its behaviour (Goi-ti et al 2003) The best-known species of

Habitat of Myotis capaccinii 163

Confidence intervals ProportionCategoryLower Upper available Selection

HabitatsCanal 0 0006 0111 ndashPool 0034 0071 0023 PositiveRiver 0923 0962 0866 Positive

River water surfaceSmooth 0807 0871 0737 PositiveRippled 0120 0182 0241 NegativeCovered 0001 0015 0022 Negative

TABLE 3 Selection of aquatic habitats and physical features of rivers by 17 M capaccinii following the Neu etalrsquos (1974) method The 95 Baileyrsquos confidence intervals (CI) for the proportion of use are shown and itscomparison to proportion available allows for assessing the significance of selection CI for canals is not reliablebecause the number of locations in this habitat was less than five

trawling bats are able to exploit niches other than water using different foragingstrategies (mainly aerial hawking) ie My-otis daubentonii (Jones and Rayner 1988Arnold et al 1998) M adversus (Jonesand Rayner 1991) and Noctilio albiventris(Kalko et al 1998)

Three putative factors might explain theobserved selection behaviour of M capac-cinii the water surfacesrsquo acoustical proper-ties the distribution of different kinds of

FIG 2 Use (O) versus availability (Q) of different aquatic habitats (left) and river features (right) resulting fromthe tracking of 17 M capaccinii Percentage use is measured as accumulated foraging time for all bats in each

class Percent availability is measured in an MCP encompassing all bat locations

water surfaces and prey availability Withinrivers tracked bats preferred open waterswith smooth surfaces as reported for othertrawling Myotis (Von Frenckell and Barclay1987 Boonman et al 1998 Rydell et al1999 Warren et al 2000) Siemers et al(2001) proved that smooth surfaces act aslsquoacoustic mirrorsrsquo improving target detec-tion for different Myotis including M ca-paccinii Calm waters present less high-frequency water noise interfering less inthe batsrsquo echolocating system (Von Frenc-kell and Barclay 1987 Mackey and Bar-clay 1989) In this sense target hiding bynoise and structural clutter presumablyleads to underexploitation of rippled watersurfaces in rivers Also the tracked individ-uals clumped together in one of the two riv-er sectors that harboured almost all of theforaging activity After leaving the roostthey moved consistently there without anyapparent search phase If physical featuresof water were the only factors leading to theselection of foraging sites we should expecta higher dispersion of individuals A morelikely explanation for this aggregation couldbe a patchy distribution of the availablefood among the acoustically suitable riverstretches Barclay et al (2000) found thatMyotis moluccarum foraged at 10 km fromthe roost and suggested that available foodin overlooked nearer water surfaces was in-sufficient to meet the batsrsquo needs The ex-treme aggregation observed in our studymay also be beneficial to bats throughgrouping and associated information trans-fer (Wilkinson 1995) presence of relativesandor minimization of predation risk Ifthis pattern of aggregated spatial use is re-lated to food availability we should expectvariation from aggregation to total disper-sion depending on the conditions in otherareas andor seasons Alternatively if theaggregation is determined by social factorswe should expect a similar pattern in otherareas andor seasons

Ponds were highly used by M capac-cinii in Israel although seemingly therewere no water-running rivers during thestudy period (Carmel and Safriel 1998)Despite the smooth and usually clutter-freewater surface of ponds we found that onlyone of the 40 available ponds in our studyarea was used and it was 100 m from one ofthe most used river stretches Low use ofponds located far from rivers has been re-ported in M daubentonii (Ciechanowski2002) Searching costs along the linearlycontinuous river stretches may be lowerthan over ponds because ponds are separat-ed by large unprofitable (terrestrial) areaswhilst rivers offer a landscape elementwhere both commuting and foraging arecompatible

Although water surfaces in canals werealso smooth these were the least preferredaquatic habitats likely because of the lackof substrate and the high water speed whichmight limit the development of insect com-munities Moreover speed at water surfaceis usually high in canals and this couldmake prey detection and capture by bats difficult Only one bat was detected forag-ing over a canal but the foraging lasted forseveral minutes and was performed whilethe inividual commuted along the canal tothe feeding place in a river Avoidance ofsurfaces completely covered by vegetationhas a more obvious explanation as it is vir-tually impossible for the bats to fly throughdense reed formations

Lakes have been also noted as exten-sively occupied by the species (Russo andJones 2003) There are no lakes in ourstudy area but we observed that the largereservoir of Tous located upstream in theXuacutequer River was not used It is probablethat lower water temperature delays theemergence of aquatic insects in the reser-voir

Myotis capaccinii is able to cover longdistances to reach its foraging sites as are

164 D Almenar J Aihartza U Goiti E Salsamendi and I Garin

other trawling species (Arnold et al 1998Barclay et al 2000) Recorded average and maximum distances are high comparedto other bats with similar wing shapes(Jones et al 1995) Aquatic habitats areusually arranged in a linear pattern or scat-tered along the landscape so M capacci-nii should be able to reach distant places es-pecially if available prey is not evenly dis-tributed throughout these habitats Thisability allows the species to exploit vast ar-eas around its main caves

This work presents the first publishedresults on habitat selection and spatial usefor M capaccinii providing useful informa-tion for the management of its foragingsites Aquatic habitats around 10 km of thebatsrsquo main caves must be protected espe-cially those consisting of open calm watersTransformation of low-flowing or stagnantwaters into fast-running waters could harmbat populations Canalization of riverscould reduce the availability of prey by re-ducing substrate quality and habitat diversi-ty and increasing water speed (Brookes1988) thereby negatively affecting M ca-paccinii The strong dependence of M ca-paccinii on aquatic habitats highlights theneed for involving this species in future en-vironmental impact assessment of anylarge-scale hydraulic transformations suchas big dams or water transfers betweenbasins

Eutrophication has a variable effectamong bats that linked to aquatic habitatsWhile Myotis macropus and M daubento-nii seem not to be negatively affected bythis process (Vaughan et al 1996 Law andUrquhart 2000) M dasycneme prefers toforage in waters with a low degree of eu-trophication (Van De Sijpe et al 2004)Meacutedard and Guibert (1990) associated M capaccinii with well-preserved Mediter-ranean streams and proposed that the deg-radation of this habitat (mainly by loss ofriverbank tree cover and eutrophication) is

involved in the rarefaction of the species inFrance Nevertheless Courtois (1998) didnot find any association of this species withgood-quality waters in Corsica Our resultsfit better with those in Corsica than with thescenario proposed by Meacutedard and Guibert(1990) given that the rivers in the studyarea are usually eutrophicated It is possiblethat eutrophication does not represent a sig-nificant hazard for M capaccinii

ACKNOWLEDEGMENTS

We wish to thank Christian Leidenberger for hisvaluable help in the fieldwork and also Toni AlcocerToni Castelloacute and Miguel Angel Monsalve We thankBrock Fenton and an anonymous referee for theircomments that helped to improve the manuscript andDavid Howe and Kate Johnson for kindly reviewingthe English This work was part of a LIFE project(LIFE00NATE7337) coordinated by Carlos IbaacuteZezEstacioacuten Bioloacutegica de DoZana (CSIC) and co-fundedby the Conselleria de Territori i Habitatge of the Ge-neralitat Valenciana and the European CommissionSupport was also provided to this project by theCPEMN ldquoLa Granja de El Salerrdquo and UPVEHU(through research grant 9UPV0076310-158492004) DA and ES were supported by the CSIC and UG by the Basque Government This study wasperformed with permission of the Valencian Gov-ernment

LITERATURE CITED

AEBISCHER N J P A ROBERTSON and R E KEN-WARD 1993 Compositional analysis of habitatuse from animal radio-tracking data Ecology 741313ndash1325

AHLEacuteN I 1990 Identification of bats in flight Swed-ish Society for Conservation of Nature Stock-holm 50 pp

AIHARTZA J R U GOITI D ALMENAR and I GARIN2003 Evidences of piscivory by Myotis capac-cinii (Bonaparte 1837) in Southern Iberian Pen-insula Acta Chiropterologica 5 193ndash198

ALDRIDGE H D J N 1988 Flight kinematics and en-ergetics in the little brown bat Myotis lucifugus(Chiroptera Vespertilionidae) with reference tothe influence of ground effect Journal of Zool-ogy (London) 216 507ndash518

ALDRIDGE H D J N and R M BRIGHAM 1988 Load carrying and manoeuvrability in an

Habitat of Myotis capaccinii 165

insectivorous bat a test of the 5 lsquorulersquo of radio-telemetry Journal of Mammalogy 69 379ndash382

ARNOLD A M BRAUN N BECKER and V STORCH1998 Beitrag zur Oumlkologie der Wasserfleder-maus (Myotis daubentoni) in Nordbaden Caroli-nea 56 103ndash110

BARCLAY R M R B J CHRUSZCZ and M RHODES2000 Foraging behaviour of the large-footed my-otis Myotis moluccarum (Chiroptera Vesperti-lionidae) in south-eastern Queensland AustralianJournal of Zoology 48 385ndash392

BOONMAN A M M BOONMAN F BRETSCHNEIDERand W A VAN DE GRIND 1998 Prey detection intrawling insectivorous bats duckweed affectshunting behaviour in Daubentonrsquos bat Myotisdaubentonii Behavioral Ecology and Sociobiol-ogy 44 99ndash107

BRITTON A R C G JONES J M V RAYNER A MBOONMAN and B VERBOOM 1997 Flight per-formance echolocation and foraging behaviourin pond bats Myotis dasycneme (ChiropteraVespertilionidae) Journal of Zoology (London)241 503ndash522

BROOKES A 1988 Channelized rivers perspectivesfor environmental management John Wiley andSons Chichester 342 pp

CARMEL Y and U SAFRIEL 1998 Habitat use bybats in a Mediterranean ecosystem in Israel mdashconservation implications Biological Conserva-tion 84 245ndash250

CHERRY S 1996 A comparison of confidence intervalmethods for habitat use-availability studies Jour-nal of Wildlife Management 60 653ndash658

CIECHANOWSKI M 2002 Community structure andactivity of bats (Chiroptera) over different waterbodies Mammalian Biology 67 276ndash285

COSSON E and P MEacuteDARD 1999 Murin de Capac-cinii Myotis capaccinii (Bonaparte 1837) Pp47ndash51 in Habitats et activiteacute de chasse des chi-roptPres menaceacutes en Europe synthPse de con-naissances actuelles en vue drsquoune gestion conser-vatrice (S Y ROUEacute and M BARATAUD eds) LeRhinolophe Vol Spec 136 pp

COURTOIS J 1998 Contribution B la connaissance dela reacutepartition et des caracteacuteristiques biologiquesdu murin de Capaccini (Myotis capaccinii) enCorse Arvicola 10 42ndash46

FENTON M B and W BOGDANOWICZ 2002 Rela-tionships between external morphology and for-aging behaviour bats in the genus Myotis Cana-dian Journal of Zoology 80 1004ndash1013

FINDLEY J S 1972 Phenetic relationships amongbats of the genus Myotis Systematic Zoology 2131ndash52

FLAVIN D A S S BIGGANE C B SHIEL P SMIDDY

and J S FAIRLEY 2001 Analysis of the diet ofDaubentonrsquos bat Myotis daubentonii in IrelandActa Theriologica 46 43ndash52

GARSHELIS D L 2000 Delusions in habitat evalua-tion measuring use selection and importancePp 111ndash164 in Research techniques in animalecology controversies and consequences (LBOITANI and T K FULLER eds) Columbia Uni-versity Press New York 442 pp

GOITI U J R AIHARTZA I GARIN and J ZABALA2003 Influence of habitat on the foraging behav-iour of the Mediterranean horseshoe bat Rhino-lophus euryale Acta Chiropterologica 5 75ndash84

GUILLEacuteN A 1999 Myotis capaccinii Pp 106ndash107 inThe atlas of European mammals (A J MITCHELL-JONES G AMORI W BOGDANOWICZ B KRY-ŠTUFEK P J H REIJNDERS F SPITZENBERGER M STUBBE J B M THISSEN V VOHRALIK andJ ZIMA eds) T amp AD Poyser Ltd London 484 pp

HUTSON A M S P MICKLEBURGH and P A RACEY(comp) 2001 Microchiropteran bats Global sta-tus survey and conservation action plan IUCNGland 259 pp

JONES G and J M V RAYNER 1988 Flight perfor-mance foraging tactics and echolocation in free-living Daubentonrsquos bats Myotis daubentoni (Chi-roptera Vespertilionidae) Journal of Zoology(London) 215 113ndash132

JONES G and J M V RAYNER 1991 Flight perfor-mance foraging tactics and echolocation in thetrawling insectivorous bat Myotis adversus (Chi-roptera Vespertilionidae) Journal of Zoology(London) 225 393ndash412

JONES G L DUVERGEacute and R D RANSOME 1995Conservation biology of an endangered speciesfield studies of greater horseshoe bat Pp309ndash324 in Ecology evolution and behaviour ofbats (P A RACEY and S M SWIFT eds) Sym-posia of the Zoological Society of London 67 1ndash421

KALKO E 1990 Field study on the echolocation andhunting behaviour of the long-fingered batMyotis capaccinii Bat Research News 3142ndash43

KALKO E K V H-U SCHNITZLER I KAIPF and AD GRINNELL 1998 Echolocation and foragingbehavior of the lesser bulldog bat Noctilio albi-ventris preadaptations for piscivory BehavioralEcology and Sociobiology 42 305ndash319

KUNZ T H and A KURTA 1988 Capture methodsand holding devices Pp 1ndash29 in Ecological andbehavioral methods for the study of bats (T HKUNZ ed) Smithsonian Institution Press Wash-ington DC 533 pp

166 D Almenar J Aihartza U Goiti E Salsamendi and I Garin

Habitat of Myotis capaccinii 167

LANZA B and P AGNELLI 2000 Long fingered batMyotis capaccinii (Bonaparte 1837) Pp 57ndash58in Italian mammals (M SPAGNESI S TOSO andA M DE MARINIS eds) Ministerio dellrsquoAmbien-te Servizio Conservazione della Natura and In-stituto Nazionale per la Fauna Selvatica A GhigiOzzano dellrsquoEmmilia 204 pp

LAW B and C A URQUHART 2000 Diet of the large-footed myotis Myotis macropus at a forest streamroost in northern New South Wales AustralianMammalogy 22 121ndash124

MA J G JONES S ZHANG J SHEN W METZNER L ZHANG and B LIANG 2003 Dietary analysisconfirms that Rickettrsquos big-footed bat (Myotisricketti) is a piscivore Journal of Zoology (Lon-don) 261 245ndash248

MACKEY R L and R M R BARCLAY 1989 The influence of physical clutter and noise on the ac-tivity of bats over water Canadian Journal of Zo-ology 67 1167ndash1170

MANLY B F J L L MCDONALD D L THOMAS T L MCDONALD and W P ERICKSON 2002 Re-source selection by animals Statistical designand analysis for field studies Kluwer AcademicPublishers Dordrecht 240 pp

MEacuteDARD P and E GUIBERT 1990 Disparition drsquounmilieu et rareacutefaction drsquoune espPce en France lemurin de Capaccini Myotis capaccinii (Bona-parte 1837) Mammalia 54 297ndash300

NEU C W C R BYERS and J M PEEK 1974 A technique for analysis of utilization-availab-ility data Journal of Wildlife Management 38541ndash545

NORBERG U M and J M V RAYNER 1987 Ecol-ogical morphology and flight in bats (MammaliaChiroptera) wing adaptations flight perfor-mance foraging strategy and echolocation Phil-osophical Transactions of the Royal Society ofLondon 316B 335ndash427

RAYNER J M V 1991 On the aerodynamics of ani-mal flight in ground effect Philosophical Trans-actions of the Royal Society of London 334B119ndash128

ROBSON S K 1984 Myotis adversus (ChiropteraVespertilionidae) Australiarsquos fish-eating batAustralian Mammalogy 7 51ndash52

RUEDI M and A MAYER 2001 Molecular systemat-ics of bats of the genus Myotis (Vespertilionidae)suggests deterministic ecomorphological conver-gences Molecular Phylogenetics and Evolution21 436ndash448

RUSSO D and G JONES 2003 Use of foraging habi-tats by bats in a Mediterranean area determined

by acoustic surveys conservation implicationsEcography 26 197ndash209

RYDELL J L A MILLER and M E JENSEN 1999Echolocation constraints of Daubentonrsquos bat foraging over water Functional Ecology 13247ndash255

SANCHIS-MOLL E J (ed) 1988 Guiacutea de la naturale-za de la Comunidad Valenciana Edicions Alfonsel MagnBnim Valencia 664 pp

SIEMERS B M P STILZ and H-U SCHNITZLER2001 The acoustic advantage of hunting at lowheights above water behavioural experiments onthe European lsquotrawlingrsquo bats Myotis capacciniiM dasycneme and M daubentonii Journal ofExperimental Biology 204 3843ndash3854

SIEMERS B M E BAUR and H-U SCHNITZLER2005 Acoustic mirror effect increases prey de-tection distance in trawling bats Naturwissen-schaften 92 272ndash276

SPITZENBERGER F and O VON HELVERSEN 2001Myotis capaccinii (Bonaparte 1837) mdash Langfuszlig-fledermaus Pp 281ndash302 in Handbuch der Saumlu-getiere Europas Band 4I Fledertiere I (J NIET-HAMMER and F KRAPP eds) Aula-Verlag Wies-baden 602 pp

VAN DE SIJPE M B VANDENDRIESSCHE P VOET J VANDENBERGHE J DUYCK E NAEYAERT M MANHAEVE and E MARTENS 2004 Summerdistribution of the pond bat Myotis dasycne-me (Chiroptera Vespertilionidae) in the west ofFlanders (Belgium) with regard to water qualityMammalia 68 377ndash386

VAUGHAN N G JONES and S HARRIS 1996 Effectsof sewage effluent on the activity of bats (Chi-roptera Vespertilionidae) foraging along riversBiological Conservation 78 337ndash343

VON FRENCKELL B and R M R BARCLAY 1987Bat activity over calm and turbulent water Cana-dian Journal of Zoology 65 219ndash222

WARREN R D D A WATERS J D ALTRINGHAM andD J BULLOCK 2000 The distribution of Dauben-tonrsquos bats (Myotis daubentonii) and pipistrellebats (Pipistrellus pipistrellus) (Vespertilionidae)in relation to small-scale variation in riverinehabitat Biological Conservation 92 85ndash91

WHITE G C and R A GARROT 1990 Analysis ofwildlife radio-tracking data Academic PressLondon 383 pp

WILKINSON G S 1995 Information transfer in batsPp 345ndash360 in Ecology evolution and behav-iour of bats (P A RACEY and S M SWIFT eds)Symposia of the Zoological Society of London67 1ndash421

Received 29 April 2005 accepted 13 January 2006

Rayner 1991) is also an advantage for a batflying low over water In addition trawlingbat species usually select calm and openwaters (Von Frenckell and Barclay 1987Boonman et al 1998 Rydell et al 1999Warren et al 2000) where target detectionmay be enhanced due to the smooth sur-faces (Boonman et al 1998 Rydell et al1999 Siemers et al 2001 2005) and re-duced sonic interferences (Von Frenckelland Barclay 1987 Mackey and Barclay1989) Nevertheless several species oftrawling bats are known to alternate theirforaging between aquatic and terrestrialhabitats (Jones and Rayner 1988 1991 Ar-nold et al 1998 Kalko et al 1998 Fentonand Bogdanowicz 2002)

Myotis capaccinii (Bonaparte 1837) is a Mediterranean troglophylous medium-sized species classed among the trawlingbats (Guilleacuten 1999 Spitzenberger and Hel-versen 2001) Traditionally it has beenlinked to inland waters and several obser-vations of hunting over water (usually calm)have been described (Ahleacuten 1990 Kalko1990 Spitzenberger and Helversen 2001)Although in Europe it is widely consideredan endangered species (Hutson et al 2001)there is little knowledge about its ecol-ogy similarly to other Mediterranean bats(Russo and Jones 2003) The large hind feetand pointed wings of M capaccinii are wellsuited to capture prey while trawlingthrough the water surface This species isknown to consume fish in the wild (Aihar-tza et al 2003)

Data published to date do not agree onthe relative use or relevance of terrestrialand aquatic environments in M capacciniiIn Corsica Courtois (1998) captured thespecies only on aquatic habitats althoughthe mist-nets used in trapping were not allo-cated in proportion to habitat availabilityCarmel and Safriel (1998) and Russo andJones (2003) performed multispecific re-search on the use of foraging habitat by

means of acoustic surveys in Israel andItaly respectively In both studies theaquatic habitats were the most frequentlyused by M capaccinii although some con-tacts were also made over terrestrial habitats(see also Cosson and Meacutedard 1999 Lanzaand Agnelli 2000 Spitzenberger and Hel-versen 2001) Given the limitations of themethods utilized these results are far fromconclusive no study has continuously sur-veyed individual bats or systematically re-corded the habitats involved in each kind ofbat activity

We expected an intensive use of aquaticenvironments by M capaccinii according toprevious data on its habitat use morpholo-gy and similarities to other trawling batsSpecifically we tested two predictions 1)terrestrial habitats if used will be not se-lected for 2) distribution and size of forag-ing sites will be linked to the availabilityand features of aquatic habitats We used radio-telemetry to determine 1) the degreeof use of terrestrial habitats 2) the use andselection of aquatic habitats at both themacrohabitat (landscape) and microhabitat(foraging site) level and 3) size and spatialdistribution of foraging sites To our knowl-edge no species-specific research on habi-tat selection and spatial use of M capacciniihas yet been published

MATERIALS AND METHODS

Study Area

The study area is located in the eastern IberianPeninsula in the province of ValPncia This region ischaracterised by a Mediterranean climate (Sanchis-Moll 1988) It is located in the low basin of theXuacutequer River where the river valley forms a largeflood plain The basin includes several tributarieswith varying degrees of steepness the most importantbeing the Albaida and Sellent rivers Flow of tributar-ies shows some degree of Mediterranean seasonalityThe altitude range is 20ndash600 m asl Riverbanks are steep in most of their courses Riverine vegetationis dominated by reeds (Arundo donax) and ripar-ian woodland is scarce Hills and slopes are mainly

158 D Almenar J Aihartza U Goiti E Salsamendi and I Garin

covered by Mediterranean scrubland and valleys andplains are dominated by orange groves As orangeagriculture demands irrigation water is highly man-aged through a big dam (called lsquoPresa de Tousrsquo)many lsquoassutsrsquo (the local name for a kind of small res-ervoir in the rivers) a vast network of irrigation ca-nals (05ndash8 m wide) and several ponds (about 3km2)Considering that it is a rural area the human popula-tion density is relatively high (142 inhabitantskm2 in2003 mdash source Instituto Nacional de Estadiacutestica)

Bat Population and Roosts

There are several caves in the area but only twoare important as bat roosts and inhabited by largegroups of M capaccinii during the nursery periodThe main roost is a limestone cave with more than500 individuals in June and about 150 individualsduring the study period (April 2004) The second a limestone chasm 84 km from the first one is alsoknown to be used from spring to autumn but thenumber of M capaccinii there is uncertain Bothroosts are shared with other cave-dwelling bats suchas Rhinolophus euryale R mehelyi Myotis blythii M myotis M emarginatus M nattereri and Minio-pterus schreibersii (D Almenar unpubl data)

Trapping Tagging and Data Collection

We radio-tagged 10 male and 10 female M ca-paccinii captured with a harp trap (Kunz and Kurta1988) on the 9th and 17th of April 2004 while enter-ing the main cave Forearm length body mass andsex were recorded for each individual After clippingthe fur between the batrsquos shoulder blades we attacheda 045 g PIP II radio-transmitter (Biotrack Ltd Dor-set UK) with Skinbond surgical adhesive (Smith andNephew Largo Florida USA) The transmitter repre-sented less than 5 of the batrsquos body mass (Aldridgeand Brigham 1988) except for four cases in whichthey were up to 57 Capture manipulation and tag-ging were performed with permission of the Valen-cian Government

The bats were tracked between the 10th and 30thof April from emergence to return They were trackedby car and on foot using up to six radio-receivers(five units were model 1000-XRS Wildlife MaterialsCarbondale USA and one was model FT-290RII AWagener Telemetrieanlagen Koumlln Germany) withthree-element Yaggi antennas Two to three bats weretracked each night simultaneously by teams of two tothree people Locations were obtained by homing-in(White and Garrot 1990) The batsrsquo time spent andactivity type in each location were recorded Activity

was classified as commuting when direct and quickdisplacements between roost and foraging sites or between two separate foraging locations occurr-ed Motionless usually weak signals coming from a shelter were classified as resting Periodic move-ments over a location were recorded as foraging Onlyforaging locations were analysed Foraging locationswere mapped in the field on orthoimages or 110000topographic maps and transferred afterward into a Geographical Information System (GIS mdash ArcView32 ESRI California USA)

Habitat Availability and Use

Photointerpretation of orthoimages (Institut Geo-grBfic ValenciB Valencia Spain) at 4 pixelsm resolu-tion with Arcview 32 GIS generated habitat mapswithin the Minimum Convex Polygon (MCP) en-compassing all radio-tracking locations and dailyroosts (White and Garrot 1990) First we distin-guished between terrestrial and aquatic habitatsSecondly three habitat categories were distinguishedwithin inland waters 1) rivers considering all the nat-ural watercourses with flowing water including smallreservoirs along them 2) canals considering onlythose not covered and more than 2 m wide 3) pondsconsidering only those with surface area greater than100 m2 Finally rivers were classified according tophysical features of water surface a) smooth definedas stagnant or slow flowing waters with smooth sur-face b) cluttered defined as water surfaces rippled orcluttered due to turbulence c) covered defined as wa-ter surfaces completely covered by vegetation (main-ly A donax) This mapping of habitats was checked inthe field River stretches were divided into homoge-neous sectors following the previous criteria This in-formation was transferred into GIS and the summedarea of each habitat category was estimated for thewhole MCP Availability of river water surfaces withdifferent physical features was estimated as thesummed area of each class in each individual foragingsite defined as the whole river sector between the twomost distant foraging locations

The bat location map was crossed with availablehabitats using Arcview 32 GIS to associate each fixwith the corresponding habitat category or river fea-ture class Foraging time spent by each bat on eachhabitat andor feature class was calculated

Habitat Selection

We used the methods proposed by Aebischer et al (1993) and Neu et al (1974) to analyse habitatselection The use of both is justified because they

Habitat of Myotis capaccinii 159

offer complementary information the former ranksthe relative preferences for habitatsfeatures and thelatter provides the significance of the selection ofeach category The method proposed by Aebischer et al (1993) is based on compositional analysis Itwas applied both to test the hypothesis of non-randomuse and to assess the rank of relative preference forhabitatsfeatures To assess the selection or rejectionof the habitat or river feature categories we used themethod proposed by Neu et al (1974) based on theestimation of Baileyrsquos simultaneous confidence inter-vals (Cherry 1996)

Compositional method was carried out usingCompos Analysis 51 (Smith Ecology Ltd Aberga-venny UK) Significance of Wilkrsquos and t-statisticswas obtained by randomisation with 1000 iterationsWhenever the use of a category was 0 it was re-placed by 0001 Because some categories were un-available for some bats we calculated the Wilkrsquoslambda as a weighted mean Neu et alrsquos method was carried out using Resource Selection for Win-dows (F Leban University of Idaho Moscow USA)For this test the time spent on each category wastransformed into the number of contacts consideringa minimum interval of 10 minutes to avoid autocorre-lation Confidence intervals for categories with lessthan five contacts were not taken into account (Man-ly et al 2002) For all tests significance was set at P lt 005 The confidence intervals were computed atthe 95 level

Spatial Use

In the case of linear elements such as rivers orcanals individual foraging sites were measured bylength and area of used stretches defined as the wholeriver or canal sector between the two most distant for-aging locations When used the length and area ofponds were added to these values Whenever the lastday roost was known we measured the minimum lin-ear distance between each foraging location and thatroost The individual foraging time in minutes wasused as an estimation of tracking effort to test the in-dependence of the measured variables from effortSquare-root transformation was applied to variablesto fit with test assumptions

RESULTS

We obtained usable tracking informa-tion for 10 females and seven males to-talling 7938 minutes of foraging time(Table 1) Tracking time was shorter thanexpected due to practical difficulties relatedto the physiography of the rivers and land-scape features The steep banks and the fre-quent changes in direction of the rivers aswell as the scarcity of vantage points overthem challenged continuous radio-tracking

160 D Almenar J Aihartza U Goiti E Salsamendi and I Garin

Body FAL Tracking TTF Foraging site DistanceCode Sex mass (g) (mm) nights (min) Lenght (m) Area (ha) Maximum 0 SD

611 X ndash ndash 4 380 4148 440 2504 781 1100628 X 79 415 4 272 2613 280 2265 1181 592640 Y ndash 409 4 406 1565 533 8840 7750 2097677 Y ndash 420 8 1217 3185 1137 9899 835 2341695 X 96 416 5 498 1165 387 600 331 194816 X 90 413 2 425 174 035 ndash ndash ndash832 X 85 424 4 1270 1623 534 957 299 212920 Y 92 409 7 939 3425 372 8488 7859 419930 Y 82 414 6 717 2785 1020 9669 2095 3601115 X 99 409 3 454 496 060 7692 7535 97143 Y 99 411 3 372 3922 411 9207 8504 652184 X 100 419 1 157 1592 540 8762 8460 196315 Y 100 418 2 115 539 050 1530 1400 121399 X 99 420 1 58 432 101 ndash ndash ndash539 Y 99 400 5 465 4184 452 9172 8613 518588 Y 99 416 1 151 941 290 8762 8653 154789 Y 98 414 1 42 100 040 8544 8544 ndash

TABLE 1 Sex body mass forearm length (FAL) tracking effort (tracking nights and TTF = tracking time of foraging activity) length and area of foraging sites and average and maximum travelling distances of 17 radio-tracked M capaccinii in the Xuacutequer River basin (Iberian Peninsula)

of most of the individuals Furthermore toensure that the observed locations were notbiased toward those sites most easily acces-sible once the signal of a target individualwas lost we searched for it thoroughly overthe whole study area This prevented biasedresults but further reduced the number of observed locations On average individualswere located during 36 nights (range 1ndash8)recording an average of 467 minutes of for-aging activity (range 42ndash1270)

Habitat Use and Selection

The MCP covered 745 km2 with an al-titude range of 25ndash417 m asl (see Fig 1)None of the bats was located foraging onterrestrial habitats All the foraging activityconcentrated on aquatic environments Thetracked bats spent almost all their foragingtime over rivers (943 unweighted accu-mulated foraging time) Three bats also foraged in a pond next to a river (55) A canal was also used (12) where onlyone bat foraged for several minutes per-forming some kind of lsquoforaging while com-mutingrsquo behaviour given that this bat alsofollowed the canal to reach the foraging

place in the river where it spent the rest ofthe night The use of aquatic habitats was non-random (compositional analysisweighted mean Wilkrsquos λ = 0036 Neu etalrsquos method G = 20154 P lt 0001 for bothcomparisons) Rivers and ponds were posi-tively selected Rivers were significantlypreferred over the rest of habitats whileponds were significantly preferred to canals(Tables 2 and 3)

Concerning river features the usedstretches were mainly stagnant or had slow-flowing waters eg small reservoirs orpools usually containing eutrophic andcloudy water Bats spent most of their timeover river stretches with smooth surfaces(847 unweighted accumulated foragingtime over rivers) although some activitywas recorded over rippled waters (147)or surfaces covered by vegetation (05)Use of each class was non-random (com-positional analysis weighted mean Wilkrsquos λ = 0268 Neu et alrsquos method G = 6830for both P lt 001) Smooth surfaces werepositively and significantly selected overthe rest of the features whereas clutter-ed and covered surfaces were negatively selected (Tables 2 and 3) Differences

Habitat of Myotis capaccinii 161

FIG 1 Map of the study area showing the aggregation of the foraging locations Rivers and canals are drawn ingrey (canals as dashed lines) Black dots represent foraging locations The asterisk represents the main cave

and the triangle the secondary cave The arrow shows the direction of the water flow

between cluttered and covered surfacescould not be tested by randomisation be-cause the degrees of freedom were less thanfive (Aebischer et al 1993) due to somelsquozerorsquo values of availability

Spatial Use

After emergence bats moved directly tothe foraging places Bats commuted overterrestrial habitats rivers or canals Al-though the availability of rivers exceeded26 km in length within the MCP all the for-aging activity concentrated in two collectiveforaging sectors mdash 44 km and 33 km longrespectively mdash situated 11 km apart fromeach other The use of spotlights revealed a high density of bats hunting in the forag-ing sites within these sectors whereas notrawling bat activity was observed in othersectors of the rivers Moreover all but oneof the tagged bats foraged exclusively inone of the two foraging sectors during thestudy period Within these sectors mostbats repeatedly changed the foraging siteswhere they flew upward and downwardfrom minutes to hours

This aggregation was not determined bylow availability of suitable foraging habi-tats Calm open waters amounted to 62 of the available river surfaces and these

two sectors comprised only 19 of thesesmooth surfaces

Between individual bats size of forag-ing sites varied from 017 to 418 km inlength (0 SD 194 145) and from 035 to 1137 ha in surface (393 317) Neithervariable was independent from tracking effort (Pearsonrsquos correlation coefficientr = 063 and r = 048 for both n = 17 and P lt 001)

On average the bats moved 37 km (SD = 38) each night to their foraging sitesThe farthest foraging location reached by an individual in a single night was 99 km fromits roost

DISCUSSION

Our results pertaining to habitat selec-tion show that M capaccinii is specialisedin feeding over water as previous studieshave indicated (Kalko 1990 Meacutedard andGuibert 1990 Carmel and Safriel 1998Courtois 1998 Cosson and Meacutedard 1999Spitzenberger and Helversen 2001 Russoand Jones 2003) The foraging activity inour study occurred over aquatic habitatswhereas use of terrestrial habitats waslinked to commuting Carmel and Safriel(1998) and Russo and Jones (2003) founda slightly higher use of terrestrial habitatthan we did although their study method

162 D Almenar J Aihartza U Goiti E Salsamendi and I Garin

Category 0 plusmn SE t-statistic df P-value Preference rankHabitats

River versus Pool 60 plusmn 100 603 16 0001 RivergtgtgtPoolgtgtgtCanalRiver versus Canal 90 plusmn 046 1951 16 0001Pool versus Canal 29 plusmn 120 262 16 001

River water surfaceSmooth versus Rippled 16 plusmn 068 231 14 001 SmoothgtgtgtRippledgtCoveredSmooth versus Covered 34 plusmn 159 211 4 ndashRippled versus Covered 17 plusmn 183 090 4 ndash

TABLE 2 Relative selection of habitat and physical features of rivers by 17 M capaccinii in spring in the XuacutequerRiver basin following compositional analysis Positive comparisons (indicating preference of numerator versusdenominator) are shown with the mean and SE values for log-ratio differences The t-statistic degrees offreedom and P-values estimated by means of randomisation are also presented Rankings of preferredcategories are provided in the last column Each category written before gt is preferred to those following thesymbol The symbol gtgtgt shows significant differences in preference

did not allow the identification of the kindof activity involved

Another selection pattern cannot be dis-carded where the population status or habi-tat availability are different The high popu-lation size of M capaccinii in the study areasuggests that the habitat quality is equallyhigh (sensu Garshelis 2000) In suboptimalhabitat conditions the species would possi-bly be forced to change its behaviour (Goi-ti et al 2003) The best-known species of

Habitat of Myotis capaccinii 163

Confidence intervals ProportionCategoryLower Upper available Selection

HabitatsCanal 0 0006 0111 ndashPool 0034 0071 0023 PositiveRiver 0923 0962 0866 Positive

River water surfaceSmooth 0807 0871 0737 PositiveRippled 0120 0182 0241 NegativeCovered 0001 0015 0022 Negative

TABLE 3 Selection of aquatic habitats and physical features of rivers by 17 M capaccinii following the Neu etalrsquos (1974) method The 95 Baileyrsquos confidence intervals (CI) for the proportion of use are shown and itscomparison to proportion available allows for assessing the significance of selection CI for canals is not reliablebecause the number of locations in this habitat was less than five

trawling bats are able to exploit niches other than water using different foragingstrategies (mainly aerial hawking) ie My-otis daubentonii (Jones and Rayner 1988Arnold et al 1998) M adversus (Jonesand Rayner 1991) and Noctilio albiventris(Kalko et al 1998)

Three putative factors might explain theobserved selection behaviour of M capac-cinii the water surfacesrsquo acoustical proper-ties the distribution of different kinds of

FIG 2 Use (O) versus availability (Q) of different aquatic habitats (left) and river features (right) resulting fromthe tracking of 17 M capaccinii Percentage use is measured as accumulated foraging time for all bats in each

class Percent availability is measured in an MCP encompassing all bat locations

water surfaces and prey availability Withinrivers tracked bats preferred open waterswith smooth surfaces as reported for othertrawling Myotis (Von Frenckell and Barclay1987 Boonman et al 1998 Rydell et al1999 Warren et al 2000) Siemers et al(2001) proved that smooth surfaces act aslsquoacoustic mirrorsrsquo improving target detec-tion for different Myotis including M ca-paccinii Calm waters present less high-frequency water noise interfering less inthe batsrsquo echolocating system (Von Frenc-kell and Barclay 1987 Mackey and Bar-clay 1989) In this sense target hiding bynoise and structural clutter presumablyleads to underexploitation of rippled watersurfaces in rivers Also the tracked individ-uals clumped together in one of the two riv-er sectors that harboured almost all of theforaging activity After leaving the roostthey moved consistently there without anyapparent search phase If physical featuresof water were the only factors leading to theselection of foraging sites we should expecta higher dispersion of individuals A morelikely explanation for this aggregation couldbe a patchy distribution of the availablefood among the acoustically suitable riverstretches Barclay et al (2000) found thatMyotis moluccarum foraged at 10 km fromthe roost and suggested that available foodin overlooked nearer water surfaces was in-sufficient to meet the batsrsquo needs The ex-treme aggregation observed in our studymay also be beneficial to bats throughgrouping and associated information trans-fer (Wilkinson 1995) presence of relativesandor minimization of predation risk Ifthis pattern of aggregated spatial use is re-lated to food availability we should expectvariation from aggregation to total disper-sion depending on the conditions in otherareas andor seasons Alternatively if theaggregation is determined by social factorswe should expect a similar pattern in otherareas andor seasons

Ponds were highly used by M capac-cinii in Israel although seemingly therewere no water-running rivers during thestudy period (Carmel and Safriel 1998)Despite the smooth and usually clutter-freewater surface of ponds we found that onlyone of the 40 available ponds in our studyarea was used and it was 100 m from one ofthe most used river stretches Low use ofponds located far from rivers has been re-ported in M daubentonii (Ciechanowski2002) Searching costs along the linearlycontinuous river stretches may be lowerthan over ponds because ponds are separat-ed by large unprofitable (terrestrial) areaswhilst rivers offer a landscape elementwhere both commuting and foraging arecompatible

Although water surfaces in canals werealso smooth these were the least preferredaquatic habitats likely because of the lackof substrate and the high water speed whichmight limit the development of insect com-munities Moreover speed at water surfaceis usually high in canals and this couldmake prey detection and capture by bats difficult Only one bat was detected forag-ing over a canal but the foraging lasted forseveral minutes and was performed whilethe inividual commuted along the canal tothe feeding place in a river Avoidance ofsurfaces completely covered by vegetationhas a more obvious explanation as it is vir-tually impossible for the bats to fly throughdense reed formations

Lakes have been also noted as exten-sively occupied by the species (Russo andJones 2003) There are no lakes in ourstudy area but we observed that the largereservoir of Tous located upstream in theXuacutequer River was not used It is probablethat lower water temperature delays theemergence of aquatic insects in the reser-voir

Myotis capaccinii is able to cover longdistances to reach its foraging sites as are

164 D Almenar J Aihartza U Goiti E Salsamendi and I Garin

other trawling species (Arnold et al 1998Barclay et al 2000) Recorded average and maximum distances are high comparedto other bats with similar wing shapes(Jones et al 1995) Aquatic habitats areusually arranged in a linear pattern or scat-tered along the landscape so M capacci-nii should be able to reach distant places es-pecially if available prey is not evenly dis-tributed throughout these habitats Thisability allows the species to exploit vast ar-eas around its main caves

This work presents the first publishedresults on habitat selection and spatial usefor M capaccinii providing useful informa-tion for the management of its foragingsites Aquatic habitats around 10 km of thebatsrsquo main caves must be protected espe-cially those consisting of open calm watersTransformation of low-flowing or stagnantwaters into fast-running waters could harmbat populations Canalization of riverscould reduce the availability of prey by re-ducing substrate quality and habitat diversi-ty and increasing water speed (Brookes1988) thereby negatively affecting M ca-paccinii The strong dependence of M ca-paccinii on aquatic habitats highlights theneed for involving this species in future en-vironmental impact assessment of anylarge-scale hydraulic transformations suchas big dams or water transfers betweenbasins

Eutrophication has a variable effectamong bats that linked to aquatic habitatsWhile Myotis macropus and M daubento-nii seem not to be negatively affected bythis process (Vaughan et al 1996 Law andUrquhart 2000) M dasycneme prefers toforage in waters with a low degree of eu-trophication (Van De Sijpe et al 2004)Meacutedard and Guibert (1990) associated M capaccinii with well-preserved Mediter-ranean streams and proposed that the deg-radation of this habitat (mainly by loss ofriverbank tree cover and eutrophication) is

involved in the rarefaction of the species inFrance Nevertheless Courtois (1998) didnot find any association of this species withgood-quality waters in Corsica Our resultsfit better with those in Corsica than with thescenario proposed by Meacutedard and Guibert(1990) given that the rivers in the studyarea are usually eutrophicated It is possiblethat eutrophication does not represent a sig-nificant hazard for M capaccinii

ACKNOWLEDEGMENTS

We wish to thank Christian Leidenberger for hisvaluable help in the fieldwork and also Toni AlcocerToni Castelloacute and Miguel Angel Monsalve We thankBrock Fenton and an anonymous referee for theircomments that helped to improve the manuscript andDavid Howe and Kate Johnson for kindly reviewingthe English This work was part of a LIFE project(LIFE00NATE7337) coordinated by Carlos IbaacuteZezEstacioacuten Bioloacutegica de DoZana (CSIC) and co-fundedby the Conselleria de Territori i Habitatge of the Ge-neralitat Valenciana and the European CommissionSupport was also provided to this project by theCPEMN ldquoLa Granja de El Salerrdquo and UPVEHU(through research grant 9UPV0076310-158492004) DA and ES were supported by the CSIC and UG by the Basque Government This study wasperformed with permission of the Valencian Gov-ernment

LITERATURE CITED

AEBISCHER N J P A ROBERTSON and R E KEN-WARD 1993 Compositional analysis of habitatuse from animal radio-tracking data Ecology 741313ndash1325

AHLEacuteN I 1990 Identification of bats in flight Swed-ish Society for Conservation of Nature Stock-holm 50 pp

AIHARTZA J R U GOITI D ALMENAR and I GARIN2003 Evidences of piscivory by Myotis capac-cinii (Bonaparte 1837) in Southern Iberian Pen-insula Acta Chiropterologica 5 193ndash198

ALDRIDGE H D J N 1988 Flight kinematics and en-ergetics in the little brown bat Myotis lucifugus(Chiroptera Vespertilionidae) with reference tothe influence of ground effect Journal of Zool-ogy (London) 216 507ndash518

ALDRIDGE H D J N and R M BRIGHAM 1988 Load carrying and manoeuvrability in an

Habitat of Myotis capaccinii 165

insectivorous bat a test of the 5 lsquorulersquo of radio-telemetry Journal of Mammalogy 69 379ndash382

ARNOLD A M BRAUN N BECKER and V STORCH1998 Beitrag zur Oumlkologie der Wasserfleder-maus (Myotis daubentoni) in Nordbaden Caroli-nea 56 103ndash110

BARCLAY R M R B J CHRUSZCZ and M RHODES2000 Foraging behaviour of the large-footed my-otis Myotis moluccarum (Chiroptera Vesperti-lionidae) in south-eastern Queensland AustralianJournal of Zoology 48 385ndash392

BOONMAN A M M BOONMAN F BRETSCHNEIDERand W A VAN DE GRIND 1998 Prey detection intrawling insectivorous bats duckweed affectshunting behaviour in Daubentonrsquos bat Myotisdaubentonii Behavioral Ecology and Sociobiol-ogy 44 99ndash107

BRITTON A R C G JONES J M V RAYNER A MBOONMAN and B VERBOOM 1997 Flight per-formance echolocation and foraging behaviourin pond bats Myotis dasycneme (ChiropteraVespertilionidae) Journal of Zoology (London)241 503ndash522

BROOKES A 1988 Channelized rivers perspectivesfor environmental management John Wiley andSons Chichester 342 pp

CARMEL Y and U SAFRIEL 1998 Habitat use bybats in a Mediterranean ecosystem in Israel mdashconservation implications Biological Conserva-tion 84 245ndash250

CHERRY S 1996 A comparison of confidence intervalmethods for habitat use-availability studies Jour-nal of Wildlife Management 60 653ndash658

CIECHANOWSKI M 2002 Community structure andactivity of bats (Chiroptera) over different waterbodies Mammalian Biology 67 276ndash285

COSSON E and P MEacuteDARD 1999 Murin de Capac-cinii Myotis capaccinii (Bonaparte 1837) Pp47ndash51 in Habitats et activiteacute de chasse des chi-roptPres menaceacutes en Europe synthPse de con-naissances actuelles en vue drsquoune gestion conser-vatrice (S Y ROUEacute and M BARATAUD eds) LeRhinolophe Vol Spec 136 pp

COURTOIS J 1998 Contribution B la connaissance dela reacutepartition et des caracteacuteristiques biologiquesdu murin de Capaccini (Myotis capaccinii) enCorse Arvicola 10 42ndash46

FENTON M B and W BOGDANOWICZ 2002 Rela-tionships between external morphology and for-aging behaviour bats in the genus Myotis Cana-dian Journal of Zoology 80 1004ndash1013

FINDLEY J S 1972 Phenetic relationships amongbats of the genus Myotis Systematic Zoology 2131ndash52

FLAVIN D A S S BIGGANE C B SHIEL P SMIDDY

and J S FAIRLEY 2001 Analysis of the diet ofDaubentonrsquos bat Myotis daubentonii in IrelandActa Theriologica 46 43ndash52

GARSHELIS D L 2000 Delusions in habitat evalua-tion measuring use selection and importancePp 111ndash164 in Research techniques in animalecology controversies and consequences (LBOITANI and T K FULLER eds) Columbia Uni-versity Press New York 442 pp

GOITI U J R AIHARTZA I GARIN and J ZABALA2003 Influence of habitat on the foraging behav-iour of the Mediterranean horseshoe bat Rhino-lophus euryale Acta Chiropterologica 5 75ndash84

GUILLEacuteN A 1999 Myotis capaccinii Pp 106ndash107 inThe atlas of European mammals (A J MITCHELL-JONES G AMORI W BOGDANOWICZ B KRY-ŠTUFEK P J H REIJNDERS F SPITZENBERGER M STUBBE J B M THISSEN V VOHRALIK andJ ZIMA eds) T amp AD Poyser Ltd London 484 pp

HUTSON A M S P MICKLEBURGH and P A RACEY(comp) 2001 Microchiropteran bats Global sta-tus survey and conservation action plan IUCNGland 259 pp

JONES G and J M V RAYNER 1988 Flight perfor-mance foraging tactics and echolocation in free-living Daubentonrsquos bats Myotis daubentoni (Chi-roptera Vespertilionidae) Journal of Zoology(London) 215 113ndash132

JONES G and J M V RAYNER 1991 Flight perfor-mance foraging tactics and echolocation in thetrawling insectivorous bat Myotis adversus (Chi-roptera Vespertilionidae) Journal of Zoology(London) 225 393ndash412

JONES G L DUVERGEacute and R D RANSOME 1995Conservation biology of an endangered speciesfield studies of greater horseshoe bat Pp309ndash324 in Ecology evolution and behaviour ofbats (P A RACEY and S M SWIFT eds) Sym-posia of the Zoological Society of London 67 1ndash421

KALKO E 1990 Field study on the echolocation andhunting behaviour of the long-fingered batMyotis capaccinii Bat Research News 3142ndash43

KALKO E K V H-U SCHNITZLER I KAIPF and AD GRINNELL 1998 Echolocation and foragingbehavior of the lesser bulldog bat Noctilio albi-ventris preadaptations for piscivory BehavioralEcology and Sociobiology 42 305ndash319

KUNZ T H and A KURTA 1988 Capture methodsand holding devices Pp 1ndash29 in Ecological andbehavioral methods for the study of bats (T HKUNZ ed) Smithsonian Institution Press Wash-ington DC 533 pp

166 D Almenar J Aihartza U Goiti E Salsamendi and I Garin

Habitat of Myotis capaccinii 167

LANZA B and P AGNELLI 2000 Long fingered batMyotis capaccinii (Bonaparte 1837) Pp 57ndash58in Italian mammals (M SPAGNESI S TOSO andA M DE MARINIS eds) Ministerio dellrsquoAmbien-te Servizio Conservazione della Natura and In-stituto Nazionale per la Fauna Selvatica A GhigiOzzano dellrsquoEmmilia 204 pp

LAW B and C A URQUHART 2000 Diet of the large-footed myotis Myotis macropus at a forest streamroost in northern New South Wales AustralianMammalogy 22 121ndash124

MA J G JONES S ZHANG J SHEN W METZNER L ZHANG and B LIANG 2003 Dietary analysisconfirms that Rickettrsquos big-footed bat (Myotisricketti) is a piscivore Journal of Zoology (Lon-don) 261 245ndash248

MACKEY R L and R M R BARCLAY 1989 The influence of physical clutter and noise on the ac-tivity of bats over water Canadian Journal of Zo-ology 67 1167ndash1170

MANLY B F J L L MCDONALD D L THOMAS T L MCDONALD and W P ERICKSON 2002 Re-source selection by animals Statistical designand analysis for field studies Kluwer AcademicPublishers Dordrecht 240 pp

MEacuteDARD P and E GUIBERT 1990 Disparition drsquounmilieu et rareacutefaction drsquoune espPce en France lemurin de Capaccini Myotis capaccinii (Bona-parte 1837) Mammalia 54 297ndash300

NEU C W C R BYERS and J M PEEK 1974 A technique for analysis of utilization-availab-ility data Journal of Wildlife Management 38541ndash545

NORBERG U M and J M V RAYNER 1987 Ecol-ogical morphology and flight in bats (MammaliaChiroptera) wing adaptations flight perfor-mance foraging strategy and echolocation Phil-osophical Transactions of the Royal Society ofLondon 316B 335ndash427

RAYNER J M V 1991 On the aerodynamics of ani-mal flight in ground effect Philosophical Trans-actions of the Royal Society of London 334B119ndash128

ROBSON S K 1984 Myotis adversus (ChiropteraVespertilionidae) Australiarsquos fish-eating batAustralian Mammalogy 7 51ndash52

RUEDI M and A MAYER 2001 Molecular systemat-ics of bats of the genus Myotis (Vespertilionidae)suggests deterministic ecomorphological conver-gences Molecular Phylogenetics and Evolution21 436ndash448

RUSSO D and G JONES 2003 Use of foraging habi-tats by bats in a Mediterranean area determined

by acoustic surveys conservation implicationsEcography 26 197ndash209

RYDELL J L A MILLER and M E JENSEN 1999Echolocation constraints of Daubentonrsquos bat foraging over water Functional Ecology 13247ndash255

SANCHIS-MOLL E J (ed) 1988 Guiacutea de la naturale-za de la Comunidad Valenciana Edicions Alfonsel MagnBnim Valencia 664 pp

SIEMERS B M P STILZ and H-U SCHNITZLER2001 The acoustic advantage of hunting at lowheights above water behavioural experiments onthe European lsquotrawlingrsquo bats Myotis capacciniiM dasycneme and M daubentonii Journal ofExperimental Biology 204 3843ndash3854

SIEMERS B M E BAUR and H-U SCHNITZLER2005 Acoustic mirror effect increases prey de-tection distance in trawling bats Naturwissen-schaften 92 272ndash276

SPITZENBERGER F and O VON HELVERSEN 2001Myotis capaccinii (Bonaparte 1837) mdash Langfuszlig-fledermaus Pp 281ndash302 in Handbuch der Saumlu-getiere Europas Band 4I Fledertiere I (J NIET-HAMMER and F KRAPP eds) Aula-Verlag Wies-baden 602 pp

VAN DE SIJPE M B VANDENDRIESSCHE P VOET J VANDENBERGHE J DUYCK E NAEYAERT M MANHAEVE and E MARTENS 2004 Summerdistribution of the pond bat Myotis dasycne-me (Chiroptera Vespertilionidae) in the west ofFlanders (Belgium) with regard to water qualityMammalia 68 377ndash386

VAUGHAN N G JONES and S HARRIS 1996 Effectsof sewage effluent on the activity of bats (Chi-roptera Vespertilionidae) foraging along riversBiological Conservation 78 337ndash343

VON FRENCKELL B and R M R BARCLAY 1987Bat activity over calm and turbulent water Cana-dian Journal of Zoology 65 219ndash222

WARREN R D D A WATERS J D ALTRINGHAM andD J BULLOCK 2000 The distribution of Dauben-tonrsquos bats (Myotis daubentonii) and pipistrellebats (Pipistrellus pipistrellus) (Vespertilionidae)in relation to small-scale variation in riverinehabitat Biological Conservation 92 85ndash91

WHITE G C and R A GARROT 1990 Analysis ofwildlife radio-tracking data Academic PressLondon 383 pp

WILKINSON G S 1995 Information transfer in batsPp 345ndash360 in Ecology evolution and behav-iour of bats (P A RACEY and S M SWIFT eds)Symposia of the Zoological Society of London67 1ndash421

Received 29 April 2005 accepted 13 January 2006

covered by Mediterranean scrubland and valleys andplains are dominated by orange groves As orangeagriculture demands irrigation water is highly man-aged through a big dam (called lsquoPresa de Tousrsquo)many lsquoassutsrsquo (the local name for a kind of small res-ervoir in the rivers) a vast network of irrigation ca-nals (05ndash8 m wide) and several ponds (about 3km2)Considering that it is a rural area the human popula-tion density is relatively high (142 inhabitantskm2 in2003 mdash source Instituto Nacional de Estadiacutestica)

Bat Population and Roosts

There are several caves in the area but only twoare important as bat roosts and inhabited by largegroups of M capaccinii during the nursery periodThe main roost is a limestone cave with more than500 individuals in June and about 150 individualsduring the study period (April 2004) The second a limestone chasm 84 km from the first one is alsoknown to be used from spring to autumn but thenumber of M capaccinii there is uncertain Bothroosts are shared with other cave-dwelling bats suchas Rhinolophus euryale R mehelyi Myotis blythii M myotis M emarginatus M nattereri and Minio-pterus schreibersii (D Almenar unpubl data)

Trapping Tagging and Data Collection

We radio-tagged 10 male and 10 female M ca-paccinii captured with a harp trap (Kunz and Kurta1988) on the 9th and 17th of April 2004 while enter-ing the main cave Forearm length body mass andsex were recorded for each individual After clippingthe fur between the batrsquos shoulder blades we attacheda 045 g PIP II radio-transmitter (Biotrack Ltd Dor-set UK) with Skinbond surgical adhesive (Smith andNephew Largo Florida USA) The transmitter repre-sented less than 5 of the batrsquos body mass (Aldridgeand Brigham 1988) except for four cases in whichthey were up to 57 Capture manipulation and tag-ging were performed with permission of the Valen-cian Government

The bats were tracked between the 10th and 30thof April from emergence to return They were trackedby car and on foot using up to six radio-receivers(five units were model 1000-XRS Wildlife MaterialsCarbondale USA and one was model FT-290RII AWagener Telemetrieanlagen Koumlln Germany) withthree-element Yaggi antennas Two to three bats weretracked each night simultaneously by teams of two tothree people Locations were obtained by homing-in(White and Garrot 1990) The batsrsquo time spent andactivity type in each location were recorded Activity

was classified as commuting when direct and quickdisplacements between roost and foraging sites or between two separate foraging locations occurr-ed Motionless usually weak signals coming from a shelter were classified as resting Periodic move-ments over a location were recorded as foraging Onlyforaging locations were analysed Foraging locationswere mapped in the field on orthoimages or 110000topographic maps and transferred afterward into a Geographical Information System (GIS mdash ArcView32 ESRI California USA)

Habitat Availability and Use

Photointerpretation of orthoimages (Institut Geo-grBfic ValenciB Valencia Spain) at 4 pixelsm resolu-tion with Arcview 32 GIS generated habitat mapswithin the Minimum Convex Polygon (MCP) en-compassing all radio-tracking locations and dailyroosts (White and Garrot 1990) First we distin-guished between terrestrial and aquatic habitatsSecondly three habitat categories were distinguishedwithin inland waters 1) rivers considering all the nat-ural watercourses with flowing water including smallreservoirs along them 2) canals considering onlythose not covered and more than 2 m wide 3) pondsconsidering only those with surface area greater than100 m2 Finally rivers were classified according tophysical features of water surface a) smooth definedas stagnant or slow flowing waters with smooth sur-face b) cluttered defined as water surfaces rippled orcluttered due to turbulence c) covered defined as wa-ter surfaces completely covered by vegetation (main-ly A donax) This mapping of habitats was checked inthe field River stretches were divided into homoge-neous sectors following the previous criteria This in-formation was transferred into GIS and the summedarea of each habitat category was estimated for thewhole MCP Availability of river water surfaces withdifferent physical features was estimated as thesummed area of each class in each individual foragingsite defined as the whole river sector between the twomost distant foraging locations

The bat location map was crossed with availablehabitats using Arcview 32 GIS to associate each fixwith the corresponding habitat category or river fea-ture class Foraging time spent by each bat on eachhabitat andor feature class was calculated

Habitat Selection

We used the methods proposed by Aebischer et al (1993) and Neu et al (1974) to analyse habitatselection The use of both is justified because they

Habitat of Myotis capaccinii 159

offer complementary information the former ranksthe relative preferences for habitatsfeatures and thelatter provides the significance of the selection ofeach category The method proposed by Aebischer et al (1993) is based on compositional analysis Itwas applied both to test the hypothesis of non-randomuse and to assess the rank of relative preference forhabitatsfeatures To assess the selection or rejectionof the habitat or river feature categories we used themethod proposed by Neu et al (1974) based on theestimation of Baileyrsquos simultaneous confidence inter-vals (Cherry 1996)

Compositional method was carried out usingCompos Analysis 51 (Smith Ecology Ltd Aberga-venny UK) Significance of Wilkrsquos and t-statisticswas obtained by randomisation with 1000 iterationsWhenever the use of a category was 0 it was re-placed by 0001 Because some categories were un-available for some bats we calculated the Wilkrsquoslambda as a weighted mean Neu et alrsquos method was carried out using Resource Selection for Win-dows (F Leban University of Idaho Moscow USA)For this test the time spent on each category wastransformed into the number of contacts consideringa minimum interval of 10 minutes to avoid autocorre-lation Confidence intervals for categories with lessthan five contacts were not taken into account (Man-ly et al 2002) For all tests significance was set at P lt 005 The confidence intervals were computed atthe 95 level

Spatial Use

In the case of linear elements such as rivers orcanals individual foraging sites were measured bylength and area of used stretches defined as the wholeriver or canal sector between the two most distant for-aging locations When used the length and area ofponds were added to these values Whenever the lastday roost was known we measured the minimum lin-ear distance between each foraging location and thatroost The individual foraging time in minutes wasused as an estimation of tracking effort to test the in-dependence of the measured variables from effortSquare-root transformation was applied to variablesto fit with test assumptions

RESULTS

We obtained usable tracking informa-tion for 10 females and seven males to-talling 7938 minutes of foraging time(Table 1) Tracking time was shorter thanexpected due to practical difficulties relatedto the physiography of the rivers and land-scape features The steep banks and the fre-quent changes in direction of the rivers aswell as the scarcity of vantage points overthem challenged continuous radio-tracking

160 D Almenar J Aihartza U Goiti E Salsamendi and I Garin

Body FAL Tracking TTF Foraging site DistanceCode Sex mass (g) (mm) nights (min) Lenght (m) Area (ha) Maximum 0 SD

611 X ndash ndash 4 380 4148 440 2504 781 1100628 X 79 415 4 272 2613 280 2265 1181 592640 Y ndash 409 4 406 1565 533 8840 7750 2097677 Y ndash 420 8 1217 3185 1137 9899 835 2341695 X 96 416 5 498 1165 387 600 331 194816 X 90 413 2 425 174 035 ndash ndash ndash832 X 85 424 4 1270 1623 534 957 299 212920 Y 92 409 7 939 3425 372 8488 7859 419930 Y 82 414 6 717 2785 1020 9669 2095 3601115 X 99 409 3 454 496 060 7692 7535 97143 Y 99 411 3 372 3922 411 9207 8504 652184 X 100 419 1 157 1592 540 8762 8460 196315 Y 100 418 2 115 539 050 1530 1400 121399 X 99 420 1 58 432 101 ndash ndash ndash539 Y 99 400 5 465 4184 452 9172 8613 518588 Y 99 416 1 151 941 290 8762 8653 154789 Y 98 414 1 42 100 040 8544 8544 ndash

TABLE 1 Sex body mass forearm length (FAL) tracking effort (tracking nights and TTF = tracking time of foraging activity) length and area of foraging sites and average and maximum travelling distances of 17 radio-tracked M capaccinii in the Xuacutequer River basin (Iberian Peninsula)

of most of the individuals Furthermore toensure that the observed locations were notbiased toward those sites most easily acces-sible once the signal of a target individualwas lost we searched for it thoroughly overthe whole study area This prevented biasedresults but further reduced the number of observed locations On average individualswere located during 36 nights (range 1ndash8)recording an average of 467 minutes of for-aging activity (range 42ndash1270)

Habitat Use and Selection

The MCP covered 745 km2 with an al-titude range of 25ndash417 m asl (see Fig 1)None of the bats was located foraging onterrestrial habitats All the foraging activityconcentrated on aquatic environments Thetracked bats spent almost all their foragingtime over rivers (943 unweighted accu-mulated foraging time) Three bats also foraged in a pond next to a river (55) A canal was also used (12) where onlyone bat foraged for several minutes per-forming some kind of lsquoforaging while com-mutingrsquo behaviour given that this bat alsofollowed the canal to reach the foraging

place in the river where it spent the rest ofthe night The use of aquatic habitats was non-random (compositional analysisweighted mean Wilkrsquos λ = 0036 Neu etalrsquos method G = 20154 P lt 0001 for bothcomparisons) Rivers and ponds were posi-tively selected Rivers were significantlypreferred over the rest of habitats whileponds were significantly preferred to canals(Tables 2 and 3)

Concerning river features the usedstretches were mainly stagnant or had slow-flowing waters eg small reservoirs orpools usually containing eutrophic andcloudy water Bats spent most of their timeover river stretches with smooth surfaces(847 unweighted accumulated foragingtime over rivers) although some activitywas recorded over rippled waters (147)or surfaces covered by vegetation (05)Use of each class was non-random (com-positional analysis weighted mean Wilkrsquos λ = 0268 Neu et alrsquos method G = 6830for both P lt 001) Smooth surfaces werepositively and significantly selected overthe rest of the features whereas clutter-ed and covered surfaces were negatively selected (Tables 2 and 3) Differences

Habitat of Myotis capaccinii 161

FIG 1 Map of the study area showing the aggregation of the foraging locations Rivers and canals are drawn ingrey (canals as dashed lines) Black dots represent foraging locations The asterisk represents the main cave

and the triangle the secondary cave The arrow shows the direction of the water flow

between cluttered and covered surfacescould not be tested by randomisation be-cause the degrees of freedom were less thanfive (Aebischer et al 1993) due to somelsquozerorsquo values of availability

Spatial Use

After emergence bats moved directly tothe foraging places Bats commuted overterrestrial habitats rivers or canals Al-though the availability of rivers exceeded26 km in length within the MCP all the for-aging activity concentrated in two collectiveforaging sectors mdash 44 km and 33 km longrespectively mdash situated 11 km apart fromeach other The use of spotlights revealed a high density of bats hunting in the forag-ing sites within these sectors whereas notrawling bat activity was observed in othersectors of the rivers Moreover all but oneof the tagged bats foraged exclusively inone of the two foraging sectors during thestudy period Within these sectors mostbats repeatedly changed the foraging siteswhere they flew upward and downwardfrom minutes to hours

This aggregation was not determined bylow availability of suitable foraging habi-tats Calm open waters amounted to 62 of the available river surfaces and these

two sectors comprised only 19 of thesesmooth surfaces

Between individual bats size of forag-ing sites varied from 017 to 418 km inlength (0 SD 194 145) and from 035 to 1137 ha in surface (393 317) Neithervariable was independent from tracking effort (Pearsonrsquos correlation coefficientr = 063 and r = 048 for both n = 17 and P lt 001)

On average the bats moved 37 km (SD = 38) each night to their foraging sitesThe farthest foraging location reached by an individual in a single night was 99 km fromits roost

DISCUSSION

Our results pertaining to habitat selec-tion show that M capaccinii is specialisedin feeding over water as previous studieshave indicated (Kalko 1990 Meacutedard andGuibert 1990 Carmel and Safriel 1998Courtois 1998 Cosson and Meacutedard 1999Spitzenberger and Helversen 2001 Russoand Jones 2003) The foraging activity inour study occurred over aquatic habitatswhereas use of terrestrial habitats waslinked to commuting Carmel and Safriel(1998) and Russo and Jones (2003) founda slightly higher use of terrestrial habitatthan we did although their study method

162 D Almenar J Aihartza U Goiti E Salsamendi and I Garin

Category 0 plusmn SE t-statistic df P-value Preference rankHabitats

River versus Pool 60 plusmn 100 603 16 0001 RivergtgtgtPoolgtgtgtCanalRiver versus Canal 90 plusmn 046 1951 16 0001Pool versus Canal 29 plusmn 120 262 16 001

River water surfaceSmooth versus Rippled 16 plusmn 068 231 14 001 SmoothgtgtgtRippledgtCoveredSmooth versus Covered 34 plusmn 159 211 4 ndashRippled versus Covered 17 plusmn 183 090 4 ndash

TABLE 2 Relative selection of habitat and physical features of rivers by 17 M capaccinii in spring in the XuacutequerRiver basin following compositional analysis Positive comparisons (indicating preference of numerator versusdenominator) are shown with the mean and SE values for log-ratio differences The t-statistic degrees offreedom and P-values estimated by means of randomisation are also presented Rankings of preferredcategories are provided in the last column Each category written before gt is preferred to those following thesymbol The symbol gtgtgt shows significant differences in preference

did not allow the identification of the kindof activity involved

Another selection pattern cannot be dis-carded where the population status or habi-tat availability are different The high popu-lation size of M capaccinii in the study areasuggests that the habitat quality is equallyhigh (sensu Garshelis 2000) In suboptimalhabitat conditions the species would possi-bly be forced to change its behaviour (Goi-ti et al 2003) The best-known species of

Habitat of Myotis capaccinii 163

Confidence intervals ProportionCategoryLower Upper available Selection

HabitatsCanal 0 0006 0111 ndashPool 0034 0071 0023 PositiveRiver 0923 0962 0866 Positive

River water surfaceSmooth 0807 0871 0737 PositiveRippled 0120 0182 0241 NegativeCovered 0001 0015 0022 Negative

TABLE 3 Selection of aquatic habitats and physical features of rivers by 17 M capaccinii following the Neu etalrsquos (1974) method The 95 Baileyrsquos confidence intervals (CI) for the proportion of use are shown and itscomparison to proportion available allows for assessing the significance of selection CI for canals is not reliablebecause the number of locations in this habitat was less than five

trawling bats are able to exploit niches other than water using different foragingstrategies (mainly aerial hawking) ie My-otis daubentonii (Jones and Rayner 1988Arnold et al 1998) M adversus (Jonesand Rayner 1991) and Noctilio albiventris(Kalko et al 1998)

Three putative factors might explain theobserved selection behaviour of M capac-cinii the water surfacesrsquo acoustical proper-ties the distribution of different kinds of

FIG 2 Use (O) versus availability (Q) of different aquatic habitats (left) and river features (right) resulting fromthe tracking of 17 M capaccinii Percentage use is measured as accumulated foraging time for all bats in each

class Percent availability is measured in an MCP encompassing all bat locations

water surfaces and prey availability Withinrivers tracked bats preferred open waterswith smooth surfaces as reported for othertrawling Myotis (Von Frenckell and Barclay1987 Boonman et al 1998 Rydell et al1999 Warren et al 2000) Siemers et al(2001) proved that smooth surfaces act aslsquoacoustic mirrorsrsquo improving target detec-tion for different Myotis including M ca-paccinii Calm waters present less high-frequency water noise interfering less inthe batsrsquo echolocating system (Von Frenc-kell and Barclay 1987 Mackey and Bar-clay 1989) In this sense target hiding bynoise and structural clutter presumablyleads to underexploitation of rippled watersurfaces in rivers Also the tracked individ-uals clumped together in one of the two riv-er sectors that harboured almost all of theforaging activity After leaving the roostthey moved consistently there without anyapparent search phase If physical featuresof water were the only factors leading to theselection of foraging sites we should expecta higher dispersion of individuals A morelikely explanation for this aggregation couldbe a patchy distribution of the availablefood among the acoustically suitable riverstretches Barclay et al (2000) found thatMyotis moluccarum foraged at 10 km fromthe roost and suggested that available foodin overlooked nearer water surfaces was in-sufficient to meet the batsrsquo needs The ex-treme aggregation observed in our studymay also be beneficial to bats throughgrouping and associated information trans-fer (Wilkinson 1995) presence of relativesandor minimization of predation risk Ifthis pattern of aggregated spatial use is re-lated to food availability we should expectvariation from aggregation to total disper-sion depending on the conditions in otherareas andor seasons Alternatively if theaggregation is determined by social factorswe should expect a similar pattern in otherareas andor seasons

Ponds were highly used by M capac-cinii in Israel although seemingly therewere no water-running rivers during thestudy period (Carmel and Safriel 1998)Despite the smooth and usually clutter-freewater surface of ponds we found that onlyone of the 40 available ponds in our studyarea was used and it was 100 m from one ofthe most used river stretches Low use ofponds located far from rivers has been re-ported in M daubentonii (Ciechanowski2002) Searching costs along the linearlycontinuous river stretches may be lowerthan over ponds because ponds are separat-ed by large unprofitable (terrestrial) areaswhilst rivers offer a landscape elementwhere both commuting and foraging arecompatible

Although water surfaces in canals werealso smooth these were the least preferredaquatic habitats likely because of the lackof substrate and the high water speed whichmight limit the development of insect com-munities Moreover speed at water surfaceis usually high in canals and this couldmake prey detection and capture by bats difficult Only one bat was detected forag-ing over a canal but the foraging lasted forseveral minutes and was performed whilethe inividual commuted along the canal tothe feeding place in a river Avoidance ofsurfaces completely covered by vegetationhas a more obvious explanation as it is vir-tually impossible for the bats to fly throughdense reed formations

Lakes have been also noted as exten-sively occupied by the species (Russo andJones 2003) There are no lakes in ourstudy area but we observed that the largereservoir of Tous located upstream in theXuacutequer River was not used It is probablethat lower water temperature delays theemergence of aquatic insects in the reser-voir

Myotis capaccinii is able to cover longdistances to reach its foraging sites as are

164 D Almenar J Aihartza U Goiti E Salsamendi and I Garin

other trawling species (Arnold et al 1998Barclay et al 2000) Recorded average and maximum distances are high comparedto other bats with similar wing shapes(Jones et al 1995) Aquatic habitats areusually arranged in a linear pattern or scat-tered along the landscape so M capacci-nii should be able to reach distant places es-pecially if available prey is not evenly dis-tributed throughout these habitats Thisability allows the species to exploit vast ar-eas around its main caves

This work presents the first publishedresults on habitat selection and spatial usefor M capaccinii providing useful informa-tion for the management of its foragingsites Aquatic habitats around 10 km of thebatsrsquo main caves must be protected espe-cially those consisting of open calm watersTransformation of low-flowing or stagnantwaters into fast-running waters could harmbat populations Canalization of riverscould reduce the availability of prey by re-ducing substrate quality and habitat diversi-ty and increasing water speed (Brookes1988) thereby negatively affecting M ca-paccinii The strong dependence of M ca-paccinii on aquatic habitats highlights theneed for involving this species in future en-vironmental impact assessment of anylarge-scale hydraulic transformations suchas big dams or water transfers betweenbasins

Eutrophication has a variable effectamong bats that linked to aquatic habitatsWhile Myotis macropus and M daubento-nii seem not to be negatively affected bythis process (Vaughan et al 1996 Law andUrquhart 2000) M dasycneme prefers toforage in waters with a low degree of eu-trophication (Van De Sijpe et al 2004)Meacutedard and Guibert (1990) associated M capaccinii with well-preserved Mediter-ranean streams and proposed that the deg-radation of this habitat (mainly by loss ofriverbank tree cover and eutrophication) is

involved in the rarefaction of the species inFrance Nevertheless Courtois (1998) didnot find any association of this species withgood-quality waters in Corsica Our resultsfit better with those in Corsica than with thescenario proposed by Meacutedard and Guibert(1990) given that the rivers in the studyarea are usually eutrophicated It is possiblethat eutrophication does not represent a sig-nificant hazard for M capaccinii

ACKNOWLEDEGMENTS

We wish to thank Christian Leidenberger for hisvaluable help in the fieldwork and also Toni AlcocerToni Castelloacute and Miguel Angel Monsalve We thankBrock Fenton and an anonymous referee for theircomments that helped to improve the manuscript andDavid Howe and Kate Johnson for kindly reviewingthe English This work was part of a LIFE project(LIFE00NATE7337) coordinated by Carlos IbaacuteZezEstacioacuten Bioloacutegica de DoZana (CSIC) and co-fundedby the Conselleria de Territori i Habitatge of the Ge-neralitat Valenciana and the European CommissionSupport was also provided to this project by theCPEMN ldquoLa Granja de El Salerrdquo and UPVEHU(through research grant 9UPV0076310-158492004) DA and ES were supported by the CSIC and UG by the Basque Government This study wasperformed with permission of the Valencian Gov-ernment

LITERATURE CITED

AEBISCHER N J P A ROBERTSON and R E KEN-WARD 1993 Compositional analysis of habitatuse from animal radio-tracking data Ecology 741313ndash1325

AHLEacuteN I 1990 Identification of bats in flight Swed-ish Society for Conservation of Nature Stock-holm 50 pp

AIHARTZA J R U GOITI D ALMENAR and I GARIN2003 Evidences of piscivory by Myotis capac-cinii (Bonaparte 1837) in Southern Iberian Pen-insula Acta Chiropterologica 5 193ndash198

ALDRIDGE H D J N 1988 Flight kinematics and en-ergetics in the little brown bat Myotis lucifugus(Chiroptera Vespertilionidae) with reference tothe influence of ground effect Journal of Zool-ogy (London) 216 507ndash518

ALDRIDGE H D J N and R M BRIGHAM 1988 Load carrying and manoeuvrability in an

Habitat of Myotis capaccinii 165

insectivorous bat a test of the 5 lsquorulersquo of radio-telemetry Journal of Mammalogy 69 379ndash382

ARNOLD A M BRAUN N BECKER and V STORCH1998 Beitrag zur Oumlkologie der Wasserfleder-maus (Myotis daubentoni) in Nordbaden Caroli-nea 56 103ndash110

BARCLAY R M R B J CHRUSZCZ and M RHODES2000 Foraging behaviour of the large-footed my-otis Myotis moluccarum (Chiroptera Vesperti-lionidae) in south-eastern Queensland AustralianJournal of Zoology 48 385ndash392

BOONMAN A M M BOONMAN F BRETSCHNEIDERand W A VAN DE GRIND 1998 Prey detection intrawling insectivorous bats duckweed affectshunting behaviour in Daubentonrsquos bat Myotisdaubentonii Behavioral Ecology and Sociobiol-ogy 44 99ndash107

BRITTON A R C G JONES J M V RAYNER A MBOONMAN and B VERBOOM 1997 Flight per-formance echolocation and foraging behaviourin pond bats Myotis dasycneme (ChiropteraVespertilionidae) Journal of Zoology (London)241 503ndash522

BROOKES A 1988 Channelized rivers perspectivesfor environmental management John Wiley andSons Chichester 342 pp

CARMEL Y and U SAFRIEL 1998 Habitat use bybats in a Mediterranean ecosystem in Israel mdashconservation implications Biological Conserva-tion 84 245ndash250

CHERRY S 1996 A comparison of confidence intervalmethods for habitat use-availability studies Jour-nal of Wildlife Management 60 653ndash658

CIECHANOWSKI M 2002 Community structure andactivity of bats (Chiroptera) over different waterbodies Mammalian Biology 67 276ndash285

COSSON E and P MEacuteDARD 1999 Murin de Capac-cinii Myotis capaccinii (Bonaparte 1837) Pp47ndash51 in Habitats et activiteacute de chasse des chi-roptPres menaceacutes en Europe synthPse de con-naissances actuelles en vue drsquoune gestion conser-vatrice (S Y ROUEacute and M BARATAUD eds) LeRhinolophe Vol Spec 136 pp

COURTOIS J 1998 Contribution B la connaissance dela reacutepartition et des caracteacuteristiques biologiquesdu murin de Capaccini (Myotis capaccinii) enCorse Arvicola 10 42ndash46

FENTON M B and W BOGDANOWICZ 2002 Rela-tionships between external morphology and for-aging behaviour bats in the genus Myotis Cana-dian Journal of Zoology 80 1004ndash1013

FINDLEY J S 1972 Phenetic relationships amongbats of the genus Myotis Systematic Zoology 2131ndash52

FLAVIN D A S S BIGGANE C B SHIEL P SMIDDY

and J S FAIRLEY 2001 Analysis of the diet ofDaubentonrsquos bat Myotis daubentonii in IrelandActa Theriologica 46 43ndash52

GARSHELIS D L 2000 Delusions in habitat evalua-tion measuring use selection and importancePp 111ndash164 in Research techniques in animalecology controversies and consequences (LBOITANI and T K FULLER eds) Columbia Uni-versity Press New York 442 pp

GOITI U J R AIHARTZA I GARIN and J ZABALA2003 Influence of habitat on the foraging behav-iour of the Mediterranean horseshoe bat Rhino-lophus euryale Acta Chiropterologica 5 75ndash84

GUILLEacuteN A 1999 Myotis capaccinii Pp 106ndash107 inThe atlas of European mammals (A J MITCHELL-JONES G AMORI W BOGDANOWICZ B KRY-ŠTUFEK P J H REIJNDERS F SPITZENBERGER M STUBBE J B M THISSEN V VOHRALIK andJ ZIMA eds) T amp AD Poyser Ltd London 484 pp

HUTSON A M S P MICKLEBURGH and P A RACEY(comp) 2001 Microchiropteran bats Global sta-tus survey and conservation action plan IUCNGland 259 pp

JONES G and J M V RAYNER 1988 Flight perfor-mance foraging tactics and echolocation in free-living Daubentonrsquos bats Myotis daubentoni (Chi-roptera Vespertilionidae) Journal of Zoology(London) 215 113ndash132

JONES G and J M V RAYNER 1991 Flight perfor-mance foraging tactics and echolocation in thetrawling insectivorous bat Myotis adversus (Chi-roptera Vespertilionidae) Journal of Zoology(London) 225 393ndash412

JONES G L DUVERGEacute and R D RANSOME 1995Conservation biology of an endangered speciesfield studies of greater horseshoe bat Pp309ndash324 in Ecology evolution and behaviour ofbats (P A RACEY and S M SWIFT eds) Sym-posia of the Zoological Society of London 67 1ndash421

KALKO E 1990 Field study on the echolocation andhunting behaviour of the long-fingered batMyotis capaccinii Bat Research News 3142ndash43

KALKO E K V H-U SCHNITZLER I KAIPF and AD GRINNELL 1998 Echolocation and foragingbehavior of the lesser bulldog bat Noctilio albi-ventris preadaptations for piscivory BehavioralEcology and Sociobiology 42 305ndash319

KUNZ T H and A KURTA 1988 Capture methodsand holding devices Pp 1ndash29 in Ecological andbehavioral methods for the study of bats (T HKUNZ ed) Smithsonian Institution Press Wash-ington DC 533 pp

166 D Almenar J Aihartza U Goiti E Salsamendi and I Garin

Habitat of Myotis capaccinii 167

LANZA B and P AGNELLI 2000 Long fingered batMyotis capaccinii (Bonaparte 1837) Pp 57ndash58in Italian mammals (M SPAGNESI S TOSO andA M DE MARINIS eds) Ministerio dellrsquoAmbien-te Servizio Conservazione della Natura and In-stituto Nazionale per la Fauna Selvatica A GhigiOzzano dellrsquoEmmilia 204 pp

LAW B and C A URQUHART 2000 Diet of the large-footed myotis Myotis macropus at a forest streamroost in northern New South Wales AustralianMammalogy 22 121ndash124

MA J G JONES S ZHANG J SHEN W METZNER L ZHANG and B LIANG 2003 Dietary analysisconfirms that Rickettrsquos big-footed bat (Myotisricketti) is a piscivore Journal of Zoology (Lon-don) 261 245ndash248

MACKEY R L and R M R BARCLAY 1989 The influence of physical clutter and noise on the ac-tivity of bats over water Canadian Journal of Zo-ology 67 1167ndash1170

MANLY B F J L L MCDONALD D L THOMAS T L MCDONALD and W P ERICKSON 2002 Re-source selection by animals Statistical designand analysis for field studies Kluwer AcademicPublishers Dordrecht 240 pp

MEacuteDARD P and E GUIBERT 1990 Disparition drsquounmilieu et rareacutefaction drsquoune espPce en France lemurin de Capaccini Myotis capaccinii (Bona-parte 1837) Mammalia 54 297ndash300

NEU C W C R BYERS and J M PEEK 1974 A technique for analysis of utilization-availab-ility data Journal of Wildlife Management 38541ndash545

NORBERG U M and J M V RAYNER 1987 Ecol-ogical morphology and flight in bats (MammaliaChiroptera) wing adaptations flight perfor-mance foraging strategy and echolocation Phil-osophical Transactions of the Royal Society ofLondon 316B 335ndash427

RAYNER J M V 1991 On the aerodynamics of ani-mal flight in ground effect Philosophical Trans-actions of the Royal Society of London 334B119ndash128

ROBSON S K 1984 Myotis adversus (ChiropteraVespertilionidae) Australiarsquos fish-eating batAustralian Mammalogy 7 51ndash52

RUEDI M and A MAYER 2001 Molecular systemat-ics of bats of the genus Myotis (Vespertilionidae)suggests deterministic ecomorphological conver-gences Molecular Phylogenetics and Evolution21 436ndash448

RUSSO D and G JONES 2003 Use of foraging habi-tats by bats in a Mediterranean area determined

by acoustic surveys conservation implicationsEcography 26 197ndash209

RYDELL J L A MILLER and M E JENSEN 1999Echolocation constraints of Daubentonrsquos bat foraging over water Functional Ecology 13247ndash255

SANCHIS-MOLL E J (ed) 1988 Guiacutea de la naturale-za de la Comunidad Valenciana Edicions Alfonsel MagnBnim Valencia 664 pp

SIEMERS B M P STILZ and H-U SCHNITZLER2001 The acoustic advantage of hunting at lowheights above water behavioural experiments onthe European lsquotrawlingrsquo bats Myotis capacciniiM dasycneme and M daubentonii Journal ofExperimental Biology 204 3843ndash3854

SIEMERS B M E BAUR and H-U SCHNITZLER2005 Acoustic mirror effect increases prey de-tection distance in trawling bats Naturwissen-schaften 92 272ndash276

SPITZENBERGER F and O VON HELVERSEN 2001Myotis capaccinii (Bonaparte 1837) mdash Langfuszlig-fledermaus Pp 281ndash302 in Handbuch der Saumlu-getiere Europas Band 4I Fledertiere I (J NIET-HAMMER and F KRAPP eds) Aula-Verlag Wies-baden 602 pp

VAN DE SIJPE M B VANDENDRIESSCHE P VOET J VANDENBERGHE J DUYCK E NAEYAERT M MANHAEVE and E MARTENS 2004 Summerdistribution of the pond bat Myotis dasycne-me (Chiroptera Vespertilionidae) in the west ofFlanders (Belgium) with regard to water qualityMammalia 68 377ndash386

VAUGHAN N G JONES and S HARRIS 1996 Effectsof sewage effluent on the activity of bats (Chi-roptera Vespertilionidae) foraging along riversBiological Conservation 78 337ndash343

VON FRENCKELL B and R M R BARCLAY 1987Bat activity over calm and turbulent water Cana-dian Journal of Zoology 65 219ndash222

WARREN R D D A WATERS J D ALTRINGHAM andD J BULLOCK 2000 The distribution of Dauben-tonrsquos bats (Myotis daubentonii) and pipistrellebats (Pipistrellus pipistrellus) (Vespertilionidae)in relation to small-scale variation in riverinehabitat Biological Conservation 92 85ndash91

WHITE G C and R A GARROT 1990 Analysis ofwildlife radio-tracking data Academic PressLondon 383 pp

WILKINSON G S 1995 Information transfer in batsPp 345ndash360 in Ecology evolution and behav-iour of bats (P A RACEY and S M SWIFT eds)Symposia of the Zoological Society of London67 1ndash421

Received 29 April 2005 accepted 13 January 2006

offer complementary information the former ranksthe relative preferences for habitatsfeatures and thelatter provides the significance of the selection ofeach category The method proposed by Aebischer et al (1993) is based on compositional analysis Itwas applied both to test the hypothesis of non-randomuse and to assess the rank of relative preference forhabitatsfeatures To assess the selection or rejectionof the habitat or river feature categories we used themethod proposed by Neu et al (1974) based on theestimation of Baileyrsquos simultaneous confidence inter-vals (Cherry 1996)

Compositional method was carried out usingCompos Analysis 51 (Smith Ecology Ltd Aberga-venny UK) Significance of Wilkrsquos and t-statisticswas obtained by randomisation with 1000 iterationsWhenever the use of a category was 0 it was re-placed by 0001 Because some categories were un-available for some bats we calculated the Wilkrsquoslambda as a weighted mean Neu et alrsquos method was carried out using Resource Selection for Win-dows (F Leban University of Idaho Moscow USA)For this test the time spent on each category wastransformed into the number of contacts consideringa minimum interval of 10 minutes to avoid autocorre-lation Confidence intervals for categories with lessthan five contacts were not taken into account (Man-ly et al 2002) For all tests significance was set at P lt 005 The confidence intervals were computed atthe 95 level

Spatial Use

In the case of linear elements such as rivers orcanals individual foraging sites were measured bylength and area of used stretches defined as the wholeriver or canal sector between the two most distant for-aging locations When used the length and area ofponds were added to these values Whenever the lastday roost was known we measured the minimum lin-ear distance between each foraging location and thatroost The individual foraging time in minutes wasused as an estimation of tracking effort to test the in-dependence of the measured variables from effortSquare-root transformation was applied to variablesto fit with test assumptions

RESULTS

We obtained usable tracking informa-tion for 10 females and seven males to-talling 7938 minutes of foraging time(Table 1) Tracking time was shorter thanexpected due to practical difficulties relatedto the physiography of the rivers and land-scape features The steep banks and the fre-quent changes in direction of the rivers aswell as the scarcity of vantage points overthem challenged continuous radio-tracking

160 D Almenar J Aihartza U Goiti E Salsamendi and I Garin

Body FAL Tracking TTF Foraging site DistanceCode Sex mass (g) (mm) nights (min) Lenght (m) Area (ha) Maximum 0 SD

611 X ndash ndash 4 380 4148 440 2504 781 1100628 X 79 415 4 272 2613 280 2265 1181 592640 Y ndash 409 4 406 1565 533 8840 7750 2097677 Y ndash 420 8 1217 3185 1137 9899 835 2341695 X 96 416 5 498 1165 387 600 331 194816 X 90 413 2 425 174 035 ndash ndash ndash832 X 85 424 4 1270 1623 534 957 299 212920 Y 92 409 7 939 3425 372 8488 7859 419930 Y 82 414 6 717 2785 1020 9669 2095 3601115 X 99 409 3 454 496 060 7692 7535 97143 Y 99 411 3 372 3922 411 9207 8504 652184 X 100 419 1 157 1592 540 8762 8460 196315 Y 100 418 2 115 539 050 1530 1400 121399 X 99 420 1 58 432 101 ndash ndash ndash539 Y 99 400 5 465 4184 452 9172 8613 518588 Y 99 416 1 151 941 290 8762 8653 154789 Y 98 414 1 42 100 040 8544 8544 ndash

TABLE 1 Sex body mass forearm length (FAL) tracking effort (tracking nights and TTF = tracking time of foraging activity) length and area of foraging sites and average and maximum travelling distances of 17 radio-tracked M capaccinii in the Xuacutequer River basin (Iberian Peninsula)

of most of the individuals Furthermore toensure that the observed locations were notbiased toward those sites most easily acces-sible once the signal of a target individualwas lost we searched for it thoroughly overthe whole study area This prevented biasedresults but further reduced the number of observed locations On average individualswere located during 36 nights (range 1ndash8)recording an average of 467 minutes of for-aging activity (range 42ndash1270)

Habitat Use and Selection

The MCP covered 745 km2 with an al-titude range of 25ndash417 m asl (see Fig 1)None of the bats was located foraging onterrestrial habitats All the foraging activityconcentrated on aquatic environments Thetracked bats spent almost all their foragingtime over rivers (943 unweighted accu-mulated foraging time) Three bats also foraged in a pond next to a river (55) A canal was also used (12) where onlyone bat foraged for several minutes per-forming some kind of lsquoforaging while com-mutingrsquo behaviour given that this bat alsofollowed the canal to reach the foraging

place in the river where it spent the rest ofthe night The use of aquatic habitats was non-random (compositional analysisweighted mean Wilkrsquos λ = 0036 Neu etalrsquos method G = 20154 P lt 0001 for bothcomparisons) Rivers and ponds were posi-tively selected Rivers were significantlypreferred over the rest of habitats whileponds were significantly preferred to canals(Tables 2 and 3)

Concerning river features the usedstretches were mainly stagnant or had slow-flowing waters eg small reservoirs orpools usually containing eutrophic andcloudy water Bats spent most of their timeover river stretches with smooth surfaces(847 unweighted accumulated foragingtime over rivers) although some activitywas recorded over rippled waters (147)or surfaces covered by vegetation (05)Use of each class was non-random (com-positional analysis weighted mean Wilkrsquos λ = 0268 Neu et alrsquos method G = 6830for both P lt 001) Smooth surfaces werepositively and significantly selected overthe rest of the features whereas clutter-ed and covered surfaces were negatively selected (Tables 2 and 3) Differences

Habitat of Myotis capaccinii 161

FIG 1 Map of the study area showing the aggregation of the foraging locations Rivers and canals are drawn ingrey (canals as dashed lines) Black dots represent foraging locations The asterisk represents the main cave

and the triangle the secondary cave The arrow shows the direction of the water flow

between cluttered and covered surfacescould not be tested by randomisation be-cause the degrees of freedom were less thanfive (Aebischer et al 1993) due to somelsquozerorsquo values of availability

Spatial Use

After emergence bats moved directly tothe foraging places Bats commuted overterrestrial habitats rivers or canals Al-though the availability of rivers exceeded26 km in length within the MCP all the for-aging activity concentrated in two collectiveforaging sectors mdash 44 km and 33 km longrespectively mdash situated 11 km apart fromeach other The use of spotlights revealed a high density of bats hunting in the forag-ing sites within these sectors whereas notrawling bat activity was observed in othersectors of the rivers Moreover all but oneof the tagged bats foraged exclusively inone of the two foraging sectors during thestudy period Within these sectors mostbats repeatedly changed the foraging siteswhere they flew upward and downwardfrom minutes to hours

This aggregation was not determined bylow availability of suitable foraging habi-tats Calm open waters amounted to 62 of the available river surfaces and these

two sectors comprised only 19 of thesesmooth surfaces

Between individual bats size of forag-ing sites varied from 017 to 418 km inlength (0 SD 194 145) and from 035 to 1137 ha in surface (393 317) Neithervariable was independent from tracking effort (Pearsonrsquos correlation coefficientr = 063 and r = 048 for both n = 17 and P lt 001)

On average the bats moved 37 km (SD = 38) each night to their foraging sitesThe farthest foraging location reached by an individual in a single night was 99 km fromits roost

DISCUSSION

Our results pertaining to habitat selec-tion show that M capaccinii is specialisedin feeding over water as previous studieshave indicated (Kalko 1990 Meacutedard andGuibert 1990 Carmel and Safriel 1998Courtois 1998 Cosson and Meacutedard 1999Spitzenberger and Helversen 2001 Russoand Jones 2003) The foraging activity inour study occurred over aquatic habitatswhereas use of terrestrial habitats waslinked to commuting Carmel and Safriel(1998) and Russo and Jones (2003) founda slightly higher use of terrestrial habitatthan we did although their study method

162 D Almenar J Aihartza U Goiti E Salsamendi and I Garin

Category 0 plusmn SE t-statistic df P-value Preference rankHabitats

River versus Pool 60 plusmn 100 603 16 0001 RivergtgtgtPoolgtgtgtCanalRiver versus Canal 90 plusmn 046 1951 16 0001Pool versus Canal 29 plusmn 120 262 16 001

River water surfaceSmooth versus Rippled 16 plusmn 068 231 14 001 SmoothgtgtgtRippledgtCoveredSmooth versus Covered 34 plusmn 159 211 4 ndashRippled versus Covered 17 plusmn 183 090 4 ndash

TABLE 2 Relative selection of habitat and physical features of rivers by 17 M capaccinii in spring in the XuacutequerRiver basin following compositional analysis Positive comparisons (indicating preference of numerator versusdenominator) are shown with the mean and SE values for log-ratio differences The t-statistic degrees offreedom and P-values estimated by means of randomisation are also presented Rankings of preferredcategories are provided in the last column Each category written before gt is preferred to those following thesymbol The symbol gtgtgt shows significant differences in preference

did not allow the identification of the kindof activity involved

Another selection pattern cannot be dis-carded where the population status or habi-tat availability are different The high popu-lation size of M capaccinii in the study areasuggests that the habitat quality is equallyhigh (sensu Garshelis 2000) In suboptimalhabitat conditions the species would possi-bly be forced to change its behaviour (Goi-ti et al 2003) The best-known species of

Habitat of Myotis capaccinii 163

Confidence intervals ProportionCategoryLower Upper available Selection

HabitatsCanal 0 0006 0111 ndashPool 0034 0071 0023 PositiveRiver 0923 0962 0866 Positive

River water surfaceSmooth 0807 0871 0737 PositiveRippled 0120 0182 0241 NegativeCovered 0001 0015 0022 Negative

TABLE 3 Selection of aquatic habitats and physical features of rivers by 17 M capaccinii following the Neu etalrsquos (1974) method The 95 Baileyrsquos confidence intervals (CI) for the proportion of use are shown and itscomparison to proportion available allows for assessing the significance of selection CI for canals is not reliablebecause the number of locations in this habitat was less than five

trawling bats are able to exploit niches other than water using different foragingstrategies (mainly aerial hawking) ie My-otis daubentonii (Jones and Rayner 1988Arnold et al 1998) M adversus (Jonesand Rayner 1991) and Noctilio albiventris(Kalko et al 1998)

Three putative factors might explain theobserved selection behaviour of M capac-cinii the water surfacesrsquo acoustical proper-ties the distribution of different kinds of

FIG 2 Use (O) versus availability (Q) of different aquatic habitats (left) and river features (right) resulting fromthe tracking of 17 M capaccinii Percentage use is measured as accumulated foraging time for all bats in each

class Percent availability is measured in an MCP encompassing all bat locations

water surfaces and prey availability Withinrivers tracked bats preferred open waterswith smooth surfaces as reported for othertrawling Myotis (Von Frenckell and Barclay1987 Boonman et al 1998 Rydell et al1999 Warren et al 2000) Siemers et al(2001) proved that smooth surfaces act aslsquoacoustic mirrorsrsquo improving target detec-tion for different Myotis including M ca-paccinii Calm waters present less high-frequency water noise interfering less inthe batsrsquo echolocating system (Von Frenc-kell and Barclay 1987 Mackey and Bar-clay 1989) In this sense target hiding bynoise and structural clutter presumablyleads to underexploitation of rippled watersurfaces in rivers Also the tracked individ-uals clumped together in one of the two riv-er sectors that harboured almost all of theforaging activity After leaving the roostthey moved consistently there without anyapparent search phase If physical featuresof water were the only factors leading to theselection of foraging sites we should expecta higher dispersion of individuals A morelikely explanation for this aggregation couldbe a patchy distribution of the availablefood among the acoustically suitable riverstretches Barclay et al (2000) found thatMyotis moluccarum foraged at 10 km fromthe roost and suggested that available foodin overlooked nearer water surfaces was in-sufficient to meet the batsrsquo needs The ex-treme aggregation observed in our studymay also be beneficial to bats throughgrouping and associated information trans-fer (Wilkinson 1995) presence of relativesandor minimization of predation risk Ifthis pattern of aggregated spatial use is re-lated to food availability we should expectvariation from aggregation to total disper-sion depending on the conditions in otherareas andor seasons Alternatively if theaggregation is determined by social factorswe should expect a similar pattern in otherareas andor seasons

Ponds were highly used by M capac-cinii in Israel although seemingly therewere no water-running rivers during thestudy period (Carmel and Safriel 1998)Despite the smooth and usually clutter-freewater surface of ponds we found that onlyone of the 40 available ponds in our studyarea was used and it was 100 m from one ofthe most used river stretches Low use ofponds located far from rivers has been re-ported in M daubentonii (Ciechanowski2002) Searching costs along the linearlycontinuous river stretches may be lowerthan over ponds because ponds are separat-ed by large unprofitable (terrestrial) areaswhilst rivers offer a landscape elementwhere both commuting and foraging arecompatible

Although water surfaces in canals werealso smooth these were the least preferredaquatic habitats likely because of the lackof substrate and the high water speed whichmight limit the development of insect com-munities Moreover speed at water surfaceis usually high in canals and this couldmake prey detection and capture by bats difficult Only one bat was detected forag-ing over a canal but the foraging lasted forseveral minutes and was performed whilethe inividual commuted along the canal tothe feeding place in a river Avoidance ofsurfaces completely covered by vegetationhas a more obvious explanation as it is vir-tually impossible for the bats to fly throughdense reed formations

Lakes have been also noted as exten-sively occupied by the species (Russo andJones 2003) There are no lakes in ourstudy area but we observed that the largereservoir of Tous located upstream in theXuacutequer River was not used It is probablethat lower water temperature delays theemergence of aquatic insects in the reser-voir

Myotis capaccinii is able to cover longdistances to reach its foraging sites as are

164 D Almenar J Aihartza U Goiti E Salsamendi and I Garin

other trawling species (Arnold et al 1998Barclay et al 2000) Recorded average and maximum distances are high comparedto other bats with similar wing shapes(Jones et al 1995) Aquatic habitats areusually arranged in a linear pattern or scat-tered along the landscape so M capacci-nii should be able to reach distant places es-pecially if available prey is not evenly dis-tributed throughout these habitats Thisability allows the species to exploit vast ar-eas around its main caves

This work presents the first publishedresults on habitat selection and spatial usefor M capaccinii providing useful informa-tion for the management of its foragingsites Aquatic habitats around 10 km of thebatsrsquo main caves must be protected espe-cially those consisting of open calm watersTransformation of low-flowing or stagnantwaters into fast-running waters could harmbat populations Canalization of riverscould reduce the availability of prey by re-ducing substrate quality and habitat diversi-ty and increasing water speed (Brookes1988) thereby negatively affecting M ca-paccinii The strong dependence of M ca-paccinii on aquatic habitats highlights theneed for involving this species in future en-vironmental impact assessment of anylarge-scale hydraulic transformations suchas big dams or water transfers betweenbasins

Eutrophication has a variable effectamong bats that linked to aquatic habitatsWhile Myotis macropus and M daubento-nii seem not to be negatively affected bythis process (Vaughan et al 1996 Law andUrquhart 2000) M dasycneme prefers toforage in waters with a low degree of eu-trophication (Van De Sijpe et al 2004)Meacutedard and Guibert (1990) associated M capaccinii with well-preserved Mediter-ranean streams and proposed that the deg-radation of this habitat (mainly by loss ofriverbank tree cover and eutrophication) is

involved in the rarefaction of the species inFrance Nevertheless Courtois (1998) didnot find any association of this species withgood-quality waters in Corsica Our resultsfit better with those in Corsica than with thescenario proposed by Meacutedard and Guibert(1990) given that the rivers in the studyarea are usually eutrophicated It is possiblethat eutrophication does not represent a sig-nificant hazard for M capaccinii

ACKNOWLEDEGMENTS

We wish to thank Christian Leidenberger for hisvaluable help in the fieldwork and also Toni AlcocerToni Castelloacute and Miguel Angel Monsalve We thankBrock Fenton and an anonymous referee for theircomments that helped to improve the manuscript andDavid Howe and Kate Johnson for kindly reviewingthe English This work was part of a LIFE project(LIFE00NATE7337) coordinated by Carlos IbaacuteZezEstacioacuten Bioloacutegica de DoZana (CSIC) and co-fundedby the Conselleria de Territori i Habitatge of the Ge-neralitat Valenciana and the European CommissionSupport was also provided to this project by theCPEMN ldquoLa Granja de El Salerrdquo and UPVEHU(through research grant 9UPV0076310-158492004) DA and ES were supported by the CSIC and UG by the Basque Government This study wasperformed with permission of the Valencian Gov-ernment

LITERATURE CITED

AEBISCHER N J P A ROBERTSON and R E KEN-WARD 1993 Compositional analysis of habitatuse from animal radio-tracking data Ecology 741313ndash1325

AHLEacuteN I 1990 Identification of bats in flight Swed-ish Society for Conservation of Nature Stock-holm 50 pp

AIHARTZA J R U GOITI D ALMENAR and I GARIN2003 Evidences of piscivory by Myotis capac-cinii (Bonaparte 1837) in Southern Iberian Pen-insula Acta Chiropterologica 5 193ndash198

ALDRIDGE H D J N 1988 Flight kinematics and en-ergetics in the little brown bat Myotis lucifugus(Chiroptera Vespertilionidae) with reference tothe influence of ground effect Journal of Zool-ogy (London) 216 507ndash518

ALDRIDGE H D J N and R M BRIGHAM 1988 Load carrying and manoeuvrability in an

Habitat of Myotis capaccinii 165

insectivorous bat a test of the 5 lsquorulersquo of radio-telemetry Journal of Mammalogy 69 379ndash382

ARNOLD A M BRAUN N BECKER and V STORCH1998 Beitrag zur Oumlkologie der Wasserfleder-maus (Myotis daubentoni) in Nordbaden Caroli-nea 56 103ndash110

BARCLAY R M R B J CHRUSZCZ and M RHODES2000 Foraging behaviour of the large-footed my-otis Myotis moluccarum (Chiroptera Vesperti-lionidae) in south-eastern Queensland AustralianJournal of Zoology 48 385ndash392

BOONMAN A M M BOONMAN F BRETSCHNEIDERand W A VAN DE GRIND 1998 Prey detection intrawling insectivorous bats duckweed affectshunting behaviour in Daubentonrsquos bat Myotisdaubentonii Behavioral Ecology and Sociobiol-ogy 44 99ndash107

BRITTON A R C G JONES J M V RAYNER A MBOONMAN and B VERBOOM 1997 Flight per-formance echolocation and foraging behaviourin pond bats Myotis dasycneme (ChiropteraVespertilionidae) Journal of Zoology (London)241 503ndash522

BROOKES A 1988 Channelized rivers perspectivesfor environmental management John Wiley andSons Chichester 342 pp

CARMEL Y and U SAFRIEL 1998 Habitat use bybats in a Mediterranean ecosystem in Israel mdashconservation implications Biological Conserva-tion 84 245ndash250

CHERRY S 1996 A comparison of confidence intervalmethods for habitat use-availability studies Jour-nal of Wildlife Management 60 653ndash658

CIECHANOWSKI M 2002 Community structure andactivity of bats (Chiroptera) over different waterbodies Mammalian Biology 67 276ndash285

COSSON E and P MEacuteDARD 1999 Murin de Capac-cinii Myotis capaccinii (Bonaparte 1837) Pp47ndash51 in Habitats et activiteacute de chasse des chi-roptPres menaceacutes en Europe synthPse de con-naissances actuelles en vue drsquoune gestion conser-vatrice (S Y ROUEacute and M BARATAUD eds) LeRhinolophe Vol Spec 136 pp

COURTOIS J 1998 Contribution B la connaissance dela reacutepartition et des caracteacuteristiques biologiquesdu murin de Capaccini (Myotis capaccinii) enCorse Arvicola 10 42ndash46

FENTON M B and W BOGDANOWICZ 2002 Rela-tionships between external morphology and for-aging behaviour bats in the genus Myotis Cana-dian Journal of Zoology 80 1004ndash1013

FINDLEY J S 1972 Phenetic relationships amongbats of the genus Myotis Systematic Zoology 2131ndash52

FLAVIN D A S S BIGGANE C B SHIEL P SMIDDY

and J S FAIRLEY 2001 Analysis of the diet ofDaubentonrsquos bat Myotis daubentonii in IrelandActa Theriologica 46 43ndash52

GARSHELIS D L 2000 Delusions in habitat evalua-tion measuring use selection and importancePp 111ndash164 in Research techniques in animalecology controversies and consequences (LBOITANI and T K FULLER eds) Columbia Uni-versity Press New York 442 pp

GOITI U J R AIHARTZA I GARIN and J ZABALA2003 Influence of habitat on the foraging behav-iour of the Mediterranean horseshoe bat Rhino-lophus euryale Acta Chiropterologica 5 75ndash84

GUILLEacuteN A 1999 Myotis capaccinii Pp 106ndash107 inThe atlas of European mammals (A J MITCHELL-JONES G AMORI W BOGDANOWICZ B KRY-ŠTUFEK P J H REIJNDERS F SPITZENBERGER M STUBBE J B M THISSEN V VOHRALIK andJ ZIMA eds) T amp AD Poyser Ltd London 484 pp

HUTSON A M S P MICKLEBURGH and P A RACEY(comp) 2001 Microchiropteran bats Global sta-tus survey and conservation action plan IUCNGland 259 pp

JONES G and J M V RAYNER 1988 Flight perfor-mance foraging tactics and echolocation in free-living Daubentonrsquos bats Myotis daubentoni (Chi-roptera Vespertilionidae) Journal of Zoology(London) 215 113ndash132

JONES G and J M V RAYNER 1991 Flight perfor-mance foraging tactics and echolocation in thetrawling insectivorous bat Myotis adversus (Chi-roptera Vespertilionidae) Journal of Zoology(London) 225 393ndash412

JONES G L DUVERGEacute and R D RANSOME 1995Conservation biology of an endangered speciesfield studies of greater horseshoe bat Pp309ndash324 in Ecology evolution and behaviour ofbats (P A RACEY and S M SWIFT eds) Sym-posia of the Zoological Society of London 67 1ndash421

KALKO E 1990 Field study on the echolocation andhunting behaviour of the long-fingered batMyotis capaccinii Bat Research News 3142ndash43

KALKO E K V H-U SCHNITZLER I KAIPF and AD GRINNELL 1998 Echolocation and foragingbehavior of the lesser bulldog bat Noctilio albi-ventris preadaptations for piscivory BehavioralEcology and Sociobiology 42 305ndash319

KUNZ T H and A KURTA 1988 Capture methodsand holding devices Pp 1ndash29 in Ecological andbehavioral methods for the study of bats (T HKUNZ ed) Smithsonian Institution Press Wash-ington DC 533 pp

166 D Almenar J Aihartza U Goiti E Salsamendi and I Garin

Habitat of Myotis capaccinii 167

LANZA B and P AGNELLI 2000 Long fingered batMyotis capaccinii (Bonaparte 1837) Pp 57ndash58in Italian mammals (M SPAGNESI S TOSO andA M DE MARINIS eds) Ministerio dellrsquoAmbien-te Servizio Conservazione della Natura and In-stituto Nazionale per la Fauna Selvatica A GhigiOzzano dellrsquoEmmilia 204 pp

LAW B and C A URQUHART 2000 Diet of the large-footed myotis Myotis macropus at a forest streamroost in northern New South Wales AustralianMammalogy 22 121ndash124

MA J G JONES S ZHANG J SHEN W METZNER L ZHANG and B LIANG 2003 Dietary analysisconfirms that Rickettrsquos big-footed bat (Myotisricketti) is a piscivore Journal of Zoology (Lon-don) 261 245ndash248

MACKEY R L and R M R BARCLAY 1989 The influence of physical clutter and noise on the ac-tivity of bats over water Canadian Journal of Zo-ology 67 1167ndash1170

MANLY B F J L L MCDONALD D L THOMAS T L MCDONALD and W P ERICKSON 2002 Re-source selection by animals Statistical designand analysis for field studies Kluwer AcademicPublishers Dordrecht 240 pp

MEacuteDARD P and E GUIBERT 1990 Disparition drsquounmilieu et rareacutefaction drsquoune espPce en France lemurin de Capaccini Myotis capaccinii (Bona-parte 1837) Mammalia 54 297ndash300

NEU C W C R BYERS and J M PEEK 1974 A technique for analysis of utilization-availab-ility data Journal of Wildlife Management 38541ndash545

NORBERG U M and J M V RAYNER 1987 Ecol-ogical morphology and flight in bats (MammaliaChiroptera) wing adaptations flight perfor-mance foraging strategy and echolocation Phil-osophical Transactions of the Royal Society ofLondon 316B 335ndash427

RAYNER J M V 1991 On the aerodynamics of ani-mal flight in ground effect Philosophical Trans-actions of the Royal Society of London 334B119ndash128

ROBSON S K 1984 Myotis adversus (ChiropteraVespertilionidae) Australiarsquos fish-eating batAustralian Mammalogy 7 51ndash52

RUEDI M and A MAYER 2001 Molecular systemat-ics of bats of the genus Myotis (Vespertilionidae)suggests deterministic ecomorphological conver-gences Molecular Phylogenetics and Evolution21 436ndash448

RUSSO D and G JONES 2003 Use of foraging habi-tats by bats in a Mediterranean area determined

by acoustic surveys conservation implicationsEcography 26 197ndash209

RYDELL J L A MILLER and M E JENSEN 1999Echolocation constraints of Daubentonrsquos bat foraging over water Functional Ecology 13247ndash255

SANCHIS-MOLL E J (ed) 1988 Guiacutea de la naturale-za de la Comunidad Valenciana Edicions Alfonsel MagnBnim Valencia 664 pp

SIEMERS B M P STILZ and H-U SCHNITZLER2001 The acoustic advantage of hunting at lowheights above water behavioural experiments onthe European lsquotrawlingrsquo bats Myotis capacciniiM dasycneme and M daubentonii Journal ofExperimental Biology 204 3843ndash3854

SIEMERS B M E BAUR and H-U SCHNITZLER2005 Acoustic mirror effect increases prey de-tection distance in trawling bats Naturwissen-schaften 92 272ndash276

SPITZENBERGER F and O VON HELVERSEN 2001Myotis capaccinii (Bonaparte 1837) mdash Langfuszlig-fledermaus Pp 281ndash302 in Handbuch der Saumlu-getiere Europas Band 4I Fledertiere I (J NIET-HAMMER and F KRAPP eds) Aula-Verlag Wies-baden 602 pp

VAN DE SIJPE M B VANDENDRIESSCHE P VOET J VANDENBERGHE J DUYCK E NAEYAERT M MANHAEVE and E MARTENS 2004 Summerdistribution of the pond bat Myotis dasycne-me (Chiroptera Vespertilionidae) in the west ofFlanders (Belgium) with regard to water qualityMammalia 68 377ndash386

VAUGHAN N G JONES and S HARRIS 1996 Effectsof sewage effluent on the activity of bats (Chi-roptera Vespertilionidae) foraging along riversBiological Conservation 78 337ndash343

VON FRENCKELL B and R M R BARCLAY 1987Bat activity over calm and turbulent water Cana-dian Journal of Zoology 65 219ndash222

WARREN R D D A WATERS J D ALTRINGHAM andD J BULLOCK 2000 The distribution of Dauben-tonrsquos bats (Myotis daubentonii) and pipistrellebats (Pipistrellus pipistrellus) (Vespertilionidae)in relation to small-scale variation in riverinehabitat Biological Conservation 92 85ndash91

WHITE G C and R A GARROT 1990 Analysis ofwildlife radio-tracking data Academic PressLondon 383 pp

WILKINSON G S 1995 Information transfer in batsPp 345ndash360 in Ecology evolution and behav-iour of bats (P A RACEY and S M SWIFT eds)Symposia of the Zoological Society of London67 1ndash421

Received 29 April 2005 accepted 13 January 2006

of most of the individuals Furthermore toensure that the observed locations were notbiased toward those sites most easily acces-sible once the signal of a target individualwas lost we searched for it thoroughly overthe whole study area This prevented biasedresults but further reduced the number of observed locations On average individualswere located during 36 nights (range 1ndash8)recording an average of 467 minutes of for-aging activity (range 42ndash1270)

Habitat Use and Selection

The MCP covered 745 km2 with an al-titude range of 25ndash417 m asl (see Fig 1)None of the bats was located foraging onterrestrial habitats All the foraging activityconcentrated on aquatic environments Thetracked bats spent almost all their foragingtime over rivers (943 unweighted accu-mulated foraging time) Three bats also foraged in a pond next to a river (55) A canal was also used (12) where onlyone bat foraged for several minutes per-forming some kind of lsquoforaging while com-mutingrsquo behaviour given that this bat alsofollowed the canal to reach the foraging

place in the river where it spent the rest ofthe night The use of aquatic habitats was non-random (compositional analysisweighted mean Wilkrsquos λ = 0036 Neu etalrsquos method G = 20154 P lt 0001 for bothcomparisons) Rivers and ponds were posi-tively selected Rivers were significantlypreferred over the rest of habitats whileponds were significantly preferred to canals(Tables 2 and 3)

Concerning river features the usedstretches were mainly stagnant or had slow-flowing waters eg small reservoirs orpools usually containing eutrophic andcloudy water Bats spent most of their timeover river stretches with smooth surfaces(847 unweighted accumulated foragingtime over rivers) although some activitywas recorded over rippled waters (147)or surfaces covered by vegetation (05)Use of each class was non-random (com-positional analysis weighted mean Wilkrsquos λ = 0268 Neu et alrsquos method G = 6830for both P lt 001) Smooth surfaces werepositively and significantly selected overthe rest of the features whereas clutter-ed and covered surfaces were negatively selected (Tables 2 and 3) Differences

Habitat of Myotis capaccinii 161

FIG 1 Map of the study area showing the aggregation of the foraging locations Rivers and canals are drawn ingrey (canals as dashed lines) Black dots represent foraging locations The asterisk represents the main cave

and the triangle the secondary cave The arrow shows the direction of the water flow

between cluttered and covered surfacescould not be tested by randomisation be-cause the degrees of freedom were less thanfive (Aebischer et al 1993) due to somelsquozerorsquo values of availability

Spatial Use

After emergence bats moved directly tothe foraging places Bats commuted overterrestrial habitats rivers or canals Al-though the availability of rivers exceeded26 km in length within the MCP all the for-aging activity concentrated in two collectiveforaging sectors mdash 44 km and 33 km longrespectively mdash situated 11 km apart fromeach other The use of spotlights revealed a high density of bats hunting in the forag-ing sites within these sectors whereas notrawling bat activity was observed in othersectors of the rivers Moreover all but oneof the tagged bats foraged exclusively inone of the two foraging sectors during thestudy period Within these sectors mostbats repeatedly changed the foraging siteswhere they flew upward and downwardfrom minutes to hours

This aggregation was not determined bylow availability of suitable foraging habi-tats Calm open waters amounted to 62 of the available river surfaces and these

two sectors comprised only 19 of thesesmooth surfaces

Between individual bats size of forag-ing sites varied from 017 to 418 km inlength (0 SD 194 145) and from 035 to 1137 ha in surface (393 317) Neithervariable was independent from tracking effort (Pearsonrsquos correlation coefficientr = 063 and r = 048 for both n = 17 and P lt 001)

On average the bats moved 37 km (SD = 38) each night to their foraging sitesThe farthest foraging location reached by an individual in a single night was 99 km fromits roost

DISCUSSION

Our results pertaining to habitat selec-tion show that M capaccinii is specialisedin feeding over water as previous studieshave indicated (Kalko 1990 Meacutedard andGuibert 1990 Carmel and Safriel 1998Courtois 1998 Cosson and Meacutedard 1999Spitzenberger and Helversen 2001 Russoand Jones 2003) The foraging activity inour study occurred over aquatic habitatswhereas use of terrestrial habitats waslinked to commuting Carmel and Safriel(1998) and Russo and Jones (2003) founda slightly higher use of terrestrial habitatthan we did although their study method

162 D Almenar J Aihartza U Goiti E Salsamendi and I Garin

Category 0 plusmn SE t-statistic df P-value Preference rankHabitats

River versus Pool 60 plusmn 100 603 16 0001 RivergtgtgtPoolgtgtgtCanalRiver versus Canal 90 plusmn 046 1951 16 0001Pool versus Canal 29 plusmn 120 262 16 001

River water surfaceSmooth versus Rippled 16 plusmn 068 231 14 001 SmoothgtgtgtRippledgtCoveredSmooth versus Covered 34 plusmn 159 211 4 ndashRippled versus Covered 17 plusmn 183 090 4 ndash

TABLE 2 Relative selection of habitat and physical features of rivers by 17 M capaccinii in spring in the XuacutequerRiver basin following compositional analysis Positive comparisons (indicating preference of numerator versusdenominator) are shown with the mean and SE values for log-ratio differences The t-statistic degrees offreedom and P-values estimated by means of randomisation are also presented Rankings of preferredcategories are provided in the last column Each category written before gt is preferred to those following thesymbol The symbol gtgtgt shows significant differences in preference

did not allow the identification of the kindof activity involved

Another selection pattern cannot be dis-carded where the population status or habi-tat availability are different The high popu-lation size of M capaccinii in the study areasuggests that the habitat quality is equallyhigh (sensu Garshelis 2000) In suboptimalhabitat conditions the species would possi-bly be forced to change its behaviour (Goi-ti et al 2003) The best-known species of

Habitat of Myotis capaccinii 163

Confidence intervals ProportionCategoryLower Upper available Selection

HabitatsCanal 0 0006 0111 ndashPool 0034 0071 0023 PositiveRiver 0923 0962 0866 Positive

River water surfaceSmooth 0807 0871 0737 PositiveRippled 0120 0182 0241 NegativeCovered 0001 0015 0022 Negative

TABLE 3 Selection of aquatic habitats and physical features of rivers by 17 M capaccinii following the Neu etalrsquos (1974) method The 95 Baileyrsquos confidence intervals (CI) for the proportion of use are shown and itscomparison to proportion available allows for assessing the significance of selection CI for canals is not reliablebecause the number of locations in this habitat was less than five

trawling bats are able to exploit niches other than water using different foragingstrategies (mainly aerial hawking) ie My-otis daubentonii (Jones and Rayner 1988Arnold et al 1998) M adversus (Jonesand Rayner 1991) and Noctilio albiventris(Kalko et al 1998)

Three putative factors might explain theobserved selection behaviour of M capac-cinii the water surfacesrsquo acoustical proper-ties the distribution of different kinds of

FIG 2 Use (O) versus availability (Q) of different aquatic habitats (left) and river features (right) resulting fromthe tracking of 17 M capaccinii Percentage use is measured as accumulated foraging time for all bats in each

class Percent availability is measured in an MCP encompassing all bat locations

water surfaces and prey availability Withinrivers tracked bats preferred open waterswith smooth surfaces as reported for othertrawling Myotis (Von Frenckell and Barclay1987 Boonman et al 1998 Rydell et al1999 Warren et al 2000) Siemers et al(2001) proved that smooth surfaces act aslsquoacoustic mirrorsrsquo improving target detec-tion for different Myotis including M ca-paccinii Calm waters present less high-frequency water noise interfering less inthe batsrsquo echolocating system (Von Frenc-kell and Barclay 1987 Mackey and Bar-clay 1989) In this sense target hiding bynoise and structural clutter presumablyleads to underexploitation of rippled watersurfaces in rivers Also the tracked individ-uals clumped together in one of the two riv-er sectors that harboured almost all of theforaging activity After leaving the roostthey moved consistently there without anyapparent search phase If physical featuresof water were the only factors leading to theselection of foraging sites we should expecta higher dispersion of individuals A morelikely explanation for this aggregation couldbe a patchy distribution of the availablefood among the acoustically suitable riverstretches Barclay et al (2000) found thatMyotis moluccarum foraged at 10 km fromthe roost and suggested that available foodin overlooked nearer water surfaces was in-sufficient to meet the batsrsquo needs The ex-treme aggregation observed in our studymay also be beneficial to bats throughgrouping and associated information trans-fer (Wilkinson 1995) presence of relativesandor minimization of predation risk Ifthis pattern of aggregated spatial use is re-lated to food availability we should expectvariation from aggregation to total disper-sion depending on the conditions in otherareas andor seasons Alternatively if theaggregation is determined by social factorswe should expect a similar pattern in otherareas andor seasons

Ponds were highly used by M capac-cinii in Israel although seemingly therewere no water-running rivers during thestudy period (Carmel and Safriel 1998)Despite the smooth and usually clutter-freewater surface of ponds we found that onlyone of the 40 available ponds in our studyarea was used and it was 100 m from one ofthe most used river stretches Low use ofponds located far from rivers has been re-ported in M daubentonii (Ciechanowski2002) Searching costs along the linearlycontinuous river stretches may be lowerthan over ponds because ponds are separat-ed by large unprofitable (terrestrial) areaswhilst rivers offer a landscape elementwhere both commuting and foraging arecompatible

Although water surfaces in canals werealso smooth these were the least preferredaquatic habitats likely because of the lackof substrate and the high water speed whichmight limit the development of insect com-munities Moreover speed at water surfaceis usually high in canals and this couldmake prey detection and capture by bats difficult Only one bat was detected forag-ing over a canal but the foraging lasted forseveral minutes and was performed whilethe inividual commuted along the canal tothe feeding place in a river Avoidance ofsurfaces completely covered by vegetationhas a more obvious explanation as it is vir-tually impossible for the bats to fly throughdense reed formations

Lakes have been also noted as exten-sively occupied by the species (Russo andJones 2003) There are no lakes in ourstudy area but we observed that the largereservoir of Tous located upstream in theXuacutequer River was not used It is probablethat lower water temperature delays theemergence of aquatic insects in the reser-voir

Myotis capaccinii is able to cover longdistances to reach its foraging sites as are

164 D Almenar J Aihartza U Goiti E Salsamendi and I Garin

other trawling species (Arnold et al 1998Barclay et al 2000) Recorded average and maximum distances are high comparedto other bats with similar wing shapes(Jones et al 1995) Aquatic habitats areusually arranged in a linear pattern or scat-tered along the landscape so M capacci-nii should be able to reach distant places es-pecially if available prey is not evenly dis-tributed throughout these habitats Thisability allows the species to exploit vast ar-eas around its main caves

This work presents the first publishedresults on habitat selection and spatial usefor M capaccinii providing useful informa-tion for the management of its foragingsites Aquatic habitats around 10 km of thebatsrsquo main caves must be protected espe-cially those consisting of open calm watersTransformation of low-flowing or stagnantwaters into fast-running waters could harmbat populations Canalization of riverscould reduce the availability of prey by re-ducing substrate quality and habitat diversi-ty and increasing water speed (Brookes1988) thereby negatively affecting M ca-paccinii The strong dependence of M ca-paccinii on aquatic habitats highlights theneed for involving this species in future en-vironmental impact assessment of anylarge-scale hydraulic transformations suchas big dams or water transfers betweenbasins

Eutrophication has a variable effectamong bats that linked to aquatic habitatsWhile Myotis macropus and M daubento-nii seem not to be negatively affected bythis process (Vaughan et al 1996 Law andUrquhart 2000) M dasycneme prefers toforage in waters with a low degree of eu-trophication (Van De Sijpe et al 2004)Meacutedard and Guibert (1990) associated M capaccinii with well-preserved Mediter-ranean streams and proposed that the deg-radation of this habitat (mainly by loss ofriverbank tree cover and eutrophication) is

involved in the rarefaction of the species inFrance Nevertheless Courtois (1998) didnot find any association of this species withgood-quality waters in Corsica Our resultsfit better with those in Corsica than with thescenario proposed by Meacutedard and Guibert(1990) given that the rivers in the studyarea are usually eutrophicated It is possiblethat eutrophication does not represent a sig-nificant hazard for M capaccinii

ACKNOWLEDEGMENTS

We wish to thank Christian Leidenberger for hisvaluable help in the fieldwork and also Toni AlcocerToni Castelloacute and Miguel Angel Monsalve We thankBrock Fenton and an anonymous referee for theircomments that helped to improve the manuscript andDavid Howe and Kate Johnson for kindly reviewingthe English This work was part of a LIFE project(LIFE00NATE7337) coordinated by Carlos IbaacuteZezEstacioacuten Bioloacutegica de DoZana (CSIC) and co-fundedby the Conselleria de Territori i Habitatge of the Ge-neralitat Valenciana and the European CommissionSupport was also provided to this project by theCPEMN ldquoLa Granja de El Salerrdquo and UPVEHU(through research grant 9UPV0076310-158492004) DA and ES were supported by the CSIC and UG by the Basque Government This study wasperformed with permission of the Valencian Gov-ernment

LITERATURE CITED

AEBISCHER N J P A ROBERTSON and R E KEN-WARD 1993 Compositional analysis of habitatuse from animal radio-tracking data Ecology 741313ndash1325

AHLEacuteN I 1990 Identification of bats in flight Swed-ish Society for Conservation of Nature Stock-holm 50 pp

AIHARTZA J R U GOITI D ALMENAR and I GARIN2003 Evidences of piscivory by Myotis capac-cinii (Bonaparte 1837) in Southern Iberian Pen-insula Acta Chiropterologica 5 193ndash198

ALDRIDGE H D J N 1988 Flight kinematics and en-ergetics in the little brown bat Myotis lucifugus(Chiroptera Vespertilionidae) with reference tothe influence of ground effect Journal of Zool-ogy (London) 216 507ndash518

ALDRIDGE H D J N and R M BRIGHAM 1988 Load carrying and manoeuvrability in an

Habitat of Myotis capaccinii 165

insectivorous bat a test of the 5 lsquorulersquo of radio-telemetry Journal of Mammalogy 69 379ndash382

ARNOLD A M BRAUN N BECKER and V STORCH1998 Beitrag zur Oumlkologie der Wasserfleder-maus (Myotis daubentoni) in Nordbaden Caroli-nea 56 103ndash110

BARCLAY R M R B J CHRUSZCZ and M RHODES2000 Foraging behaviour of the large-footed my-otis Myotis moluccarum (Chiroptera Vesperti-lionidae) in south-eastern Queensland AustralianJournal of Zoology 48 385ndash392

BOONMAN A M M BOONMAN F BRETSCHNEIDERand W A VAN DE GRIND 1998 Prey detection intrawling insectivorous bats duckweed affectshunting behaviour in Daubentonrsquos bat Myotisdaubentonii Behavioral Ecology and Sociobiol-ogy 44 99ndash107

BRITTON A R C G JONES J M V RAYNER A MBOONMAN and B VERBOOM 1997 Flight per-formance echolocation and foraging behaviourin pond bats Myotis dasycneme (ChiropteraVespertilionidae) Journal of Zoology (London)241 503ndash522

BROOKES A 1988 Channelized rivers perspectivesfor environmental management John Wiley andSons Chichester 342 pp

CARMEL Y and U SAFRIEL 1998 Habitat use bybats in a Mediterranean ecosystem in Israel mdashconservation implications Biological Conserva-tion 84 245ndash250

CHERRY S 1996 A comparison of confidence intervalmethods for habitat use-availability studies Jour-nal of Wildlife Management 60 653ndash658

CIECHANOWSKI M 2002 Community structure andactivity of bats (Chiroptera) over different waterbodies Mammalian Biology 67 276ndash285

COSSON E and P MEacuteDARD 1999 Murin de Capac-cinii Myotis capaccinii (Bonaparte 1837) Pp47ndash51 in Habitats et activiteacute de chasse des chi-roptPres menaceacutes en Europe synthPse de con-naissances actuelles en vue drsquoune gestion conser-vatrice (S Y ROUEacute and M BARATAUD eds) LeRhinolophe Vol Spec 136 pp

COURTOIS J 1998 Contribution B la connaissance dela reacutepartition et des caracteacuteristiques biologiquesdu murin de Capaccini (Myotis capaccinii) enCorse Arvicola 10 42ndash46

FENTON M B and W BOGDANOWICZ 2002 Rela-tionships between external morphology and for-aging behaviour bats in the genus Myotis Cana-dian Journal of Zoology 80 1004ndash1013

FINDLEY J S 1972 Phenetic relationships amongbats of the genus Myotis Systematic Zoology 2131ndash52

FLAVIN D A S S BIGGANE C B SHIEL P SMIDDY

and J S FAIRLEY 2001 Analysis of the diet ofDaubentonrsquos bat Myotis daubentonii in IrelandActa Theriologica 46 43ndash52

GARSHELIS D L 2000 Delusions in habitat evalua-tion measuring use selection and importancePp 111ndash164 in Research techniques in animalecology controversies and consequences (LBOITANI and T K FULLER eds) Columbia Uni-versity Press New York 442 pp

GOITI U J R AIHARTZA I GARIN and J ZABALA2003 Influence of habitat on the foraging behav-iour of the Mediterranean horseshoe bat Rhino-lophus euryale Acta Chiropterologica 5 75ndash84

GUILLEacuteN A 1999 Myotis capaccinii Pp 106ndash107 inThe atlas of European mammals (A J MITCHELL-JONES G AMORI W BOGDANOWICZ B KRY-ŠTUFEK P J H REIJNDERS F SPITZENBERGER M STUBBE J B M THISSEN V VOHRALIK andJ ZIMA eds) T amp AD Poyser Ltd London 484 pp

HUTSON A M S P MICKLEBURGH and P A RACEY(comp) 2001 Microchiropteran bats Global sta-tus survey and conservation action plan IUCNGland 259 pp

JONES G and J M V RAYNER 1988 Flight perfor-mance foraging tactics and echolocation in free-living Daubentonrsquos bats Myotis daubentoni (Chi-roptera Vespertilionidae) Journal of Zoology(London) 215 113ndash132

JONES G and J M V RAYNER 1991 Flight perfor-mance foraging tactics and echolocation in thetrawling insectivorous bat Myotis adversus (Chi-roptera Vespertilionidae) Journal of Zoology(London) 225 393ndash412

JONES G L DUVERGEacute and R D RANSOME 1995Conservation biology of an endangered speciesfield studies of greater horseshoe bat Pp309ndash324 in Ecology evolution and behaviour ofbats (P A RACEY and S M SWIFT eds) Sym-posia of the Zoological Society of London 67 1ndash421

KALKO E 1990 Field study on the echolocation andhunting behaviour of the long-fingered batMyotis capaccinii Bat Research News 3142ndash43

KALKO E K V H-U SCHNITZLER I KAIPF and AD GRINNELL 1998 Echolocation and foragingbehavior of the lesser bulldog bat Noctilio albi-ventris preadaptations for piscivory BehavioralEcology and Sociobiology 42 305ndash319

KUNZ T H and A KURTA 1988 Capture methodsand holding devices Pp 1ndash29 in Ecological andbehavioral methods for the study of bats (T HKUNZ ed) Smithsonian Institution Press Wash-ington DC 533 pp

166 D Almenar J Aihartza U Goiti E Salsamendi and I Garin

Habitat of Myotis capaccinii 167

LANZA B and P AGNELLI 2000 Long fingered batMyotis capaccinii (Bonaparte 1837) Pp 57ndash58in Italian mammals (M SPAGNESI S TOSO andA M DE MARINIS eds) Ministerio dellrsquoAmbien-te Servizio Conservazione della Natura and In-stituto Nazionale per la Fauna Selvatica A GhigiOzzano dellrsquoEmmilia 204 pp

LAW B and C A URQUHART 2000 Diet of the large-footed myotis Myotis macropus at a forest streamroost in northern New South Wales AustralianMammalogy 22 121ndash124

MA J G JONES S ZHANG J SHEN W METZNER L ZHANG and B LIANG 2003 Dietary analysisconfirms that Rickettrsquos big-footed bat (Myotisricketti) is a piscivore Journal of Zoology (Lon-don) 261 245ndash248

MACKEY R L and R M R BARCLAY 1989 The influence of physical clutter and noise on the ac-tivity of bats over water Canadian Journal of Zo-ology 67 1167ndash1170

MANLY B F J L L MCDONALD D L THOMAS T L MCDONALD and W P ERICKSON 2002 Re-source selection by animals Statistical designand analysis for field studies Kluwer AcademicPublishers Dordrecht 240 pp

MEacuteDARD P and E GUIBERT 1990 Disparition drsquounmilieu et rareacutefaction drsquoune espPce en France lemurin de Capaccini Myotis capaccinii (Bona-parte 1837) Mammalia 54 297ndash300

NEU C W C R BYERS and J M PEEK 1974 A technique for analysis of utilization-availab-ility data Journal of Wildlife Management 38541ndash545

NORBERG U M and J M V RAYNER 1987 Ecol-ogical morphology and flight in bats (MammaliaChiroptera) wing adaptations flight perfor-mance foraging strategy and echolocation Phil-osophical Transactions of the Royal Society ofLondon 316B 335ndash427

RAYNER J M V 1991 On the aerodynamics of ani-mal flight in ground effect Philosophical Trans-actions of the Royal Society of London 334B119ndash128

ROBSON S K 1984 Myotis adversus (ChiropteraVespertilionidae) Australiarsquos fish-eating batAustralian Mammalogy 7 51ndash52

RUEDI M and A MAYER 2001 Molecular systemat-ics of bats of the genus Myotis (Vespertilionidae)suggests deterministic ecomorphological conver-gences Molecular Phylogenetics and Evolution21 436ndash448

RUSSO D and G JONES 2003 Use of foraging habi-tats by bats in a Mediterranean area determined

by acoustic surveys conservation implicationsEcography 26 197ndash209

RYDELL J L A MILLER and M E JENSEN 1999Echolocation constraints of Daubentonrsquos bat foraging over water Functional Ecology 13247ndash255

SANCHIS-MOLL E J (ed) 1988 Guiacutea de la naturale-za de la Comunidad Valenciana Edicions Alfonsel MagnBnim Valencia 664 pp

SIEMERS B M P STILZ and H-U SCHNITZLER2001 The acoustic advantage of hunting at lowheights above water behavioural experiments onthe European lsquotrawlingrsquo bats Myotis capacciniiM dasycneme and M daubentonii Journal ofExperimental Biology 204 3843ndash3854

SIEMERS B M E BAUR and H-U SCHNITZLER2005 Acoustic mirror effect increases prey de-tection distance in trawling bats Naturwissen-schaften 92 272ndash276

SPITZENBERGER F and O VON HELVERSEN 2001Myotis capaccinii (Bonaparte 1837) mdash Langfuszlig-fledermaus Pp 281ndash302 in Handbuch der Saumlu-getiere Europas Band 4I Fledertiere I (J NIET-HAMMER and F KRAPP eds) Aula-Verlag Wies-baden 602 pp

VAN DE SIJPE M B VANDENDRIESSCHE P VOET J VANDENBERGHE J DUYCK E NAEYAERT M MANHAEVE and E MARTENS 2004 Summerdistribution of the pond bat Myotis dasycne-me (Chiroptera Vespertilionidae) in the west ofFlanders (Belgium) with regard to water qualityMammalia 68 377ndash386

VAUGHAN N G JONES and S HARRIS 1996 Effectsof sewage effluent on the activity of bats (Chi-roptera Vespertilionidae) foraging along riversBiological Conservation 78 337ndash343

VON FRENCKELL B and R M R BARCLAY 1987Bat activity over calm and turbulent water Cana-dian Journal of Zoology 65 219ndash222

WARREN R D D A WATERS J D ALTRINGHAM andD J BULLOCK 2000 The distribution of Dauben-tonrsquos bats (Myotis daubentonii) and pipistrellebats (Pipistrellus pipistrellus) (Vespertilionidae)in relation to small-scale variation in riverinehabitat Biological Conservation 92 85ndash91

WHITE G C and R A GARROT 1990 Analysis ofwildlife radio-tracking data Academic PressLondon 383 pp

WILKINSON G S 1995 Information transfer in batsPp 345ndash360 in Ecology evolution and behav-iour of bats (P A RACEY and S M SWIFT eds)Symposia of the Zoological Society of London67 1ndash421

Received 29 April 2005 accepted 13 January 2006

between cluttered and covered surfacescould not be tested by randomisation be-cause the degrees of freedom were less thanfive (Aebischer et al 1993) due to somelsquozerorsquo values of availability

Spatial Use

After emergence bats moved directly tothe foraging places Bats commuted overterrestrial habitats rivers or canals Al-though the availability of rivers exceeded26 km in length within the MCP all the for-aging activity concentrated in two collectiveforaging sectors mdash 44 km and 33 km longrespectively mdash situated 11 km apart fromeach other The use of spotlights revealed a high density of bats hunting in the forag-ing sites within these sectors whereas notrawling bat activity was observed in othersectors of the rivers Moreover all but oneof the tagged bats foraged exclusively inone of the two foraging sectors during thestudy period Within these sectors mostbats repeatedly changed the foraging siteswhere they flew upward and downwardfrom minutes to hours

This aggregation was not determined bylow availability of suitable foraging habi-tats Calm open waters amounted to 62 of the available river surfaces and these

two sectors comprised only 19 of thesesmooth surfaces

Between individual bats size of forag-ing sites varied from 017 to 418 km inlength (0 SD 194 145) and from 035 to 1137 ha in surface (393 317) Neithervariable was independent from tracking effort (Pearsonrsquos correlation coefficientr = 063 and r = 048 for both n = 17 and P lt 001)

On average the bats moved 37 km (SD = 38) each night to their foraging sitesThe farthest foraging location reached by an individual in a single night was 99 km fromits roost

DISCUSSION

Our results pertaining to habitat selec-tion show that M capaccinii is specialisedin feeding over water as previous studieshave indicated (Kalko 1990 Meacutedard andGuibert 1990 Carmel and Safriel 1998Courtois 1998 Cosson and Meacutedard 1999Spitzenberger and Helversen 2001 Russoand Jones 2003) The foraging activity inour study occurred over aquatic habitatswhereas use of terrestrial habitats waslinked to commuting Carmel and Safriel(1998) and Russo and Jones (2003) founda slightly higher use of terrestrial habitatthan we did although their study method

162 D Almenar J Aihartza U Goiti E Salsamendi and I Garin

Category 0 plusmn SE t-statistic df P-value Preference rankHabitats

River versus Pool 60 plusmn 100 603 16 0001 RivergtgtgtPoolgtgtgtCanalRiver versus Canal 90 plusmn 046 1951 16 0001Pool versus Canal 29 plusmn 120 262 16 001

River water surfaceSmooth versus Rippled 16 plusmn 068 231 14 001 SmoothgtgtgtRippledgtCoveredSmooth versus Covered 34 plusmn 159 211 4 ndashRippled versus Covered 17 plusmn 183 090 4 ndash

TABLE 2 Relative selection of habitat and physical features of rivers by 17 M capaccinii in spring in the XuacutequerRiver basin following compositional analysis Positive comparisons (indicating preference of numerator versusdenominator) are shown with the mean and SE values for log-ratio differences The t-statistic degrees offreedom and P-values estimated by means of randomisation are also presented Rankings of preferredcategories are provided in the last column Each category written before gt is preferred to those following thesymbol The symbol gtgtgt shows significant differences in preference

did not allow the identification of the kindof activity involved

Another selection pattern cannot be dis-carded where the population status or habi-tat availability are different The high popu-lation size of M capaccinii in the study areasuggests that the habitat quality is equallyhigh (sensu Garshelis 2000) In suboptimalhabitat conditions the species would possi-bly be forced to change its behaviour (Goi-ti et al 2003) The best-known species of

Habitat of Myotis capaccinii 163

Confidence intervals ProportionCategoryLower Upper available Selection

HabitatsCanal 0 0006 0111 ndashPool 0034 0071 0023 PositiveRiver 0923 0962 0866 Positive

River water surfaceSmooth 0807 0871 0737 PositiveRippled 0120 0182 0241 NegativeCovered 0001 0015 0022 Negative

TABLE 3 Selection of aquatic habitats and physical features of rivers by 17 M capaccinii following the Neu etalrsquos (1974) method The 95 Baileyrsquos confidence intervals (CI) for the proportion of use are shown and itscomparison to proportion available allows for assessing the significance of selection CI for canals is not reliablebecause the number of locations in this habitat was less than five

trawling bats are able to exploit niches other than water using different foragingstrategies (mainly aerial hawking) ie My-otis daubentonii (Jones and Rayner 1988Arnold et al 1998) M adversus (Jonesand Rayner 1991) and Noctilio albiventris(Kalko et al 1998)

Three putative factors might explain theobserved selection behaviour of M capac-cinii the water surfacesrsquo acoustical proper-ties the distribution of different kinds of

FIG 2 Use (O) versus availability (Q) of different aquatic habitats (left) and river features (right) resulting fromthe tracking of 17 M capaccinii Percentage use is measured as accumulated foraging time for all bats in each

class Percent availability is measured in an MCP encompassing all bat locations

water surfaces and prey availability Withinrivers tracked bats preferred open waterswith smooth surfaces as reported for othertrawling Myotis (Von Frenckell and Barclay1987 Boonman et al 1998 Rydell et al1999 Warren et al 2000) Siemers et al(2001) proved that smooth surfaces act aslsquoacoustic mirrorsrsquo improving target detec-tion for different Myotis including M ca-paccinii Calm waters present less high-frequency water noise interfering less inthe batsrsquo echolocating system (Von Frenc-kell and Barclay 1987 Mackey and Bar-clay 1989) In this sense target hiding bynoise and structural clutter presumablyleads to underexploitation of rippled watersurfaces in rivers Also the tracked individ-uals clumped together in one of the two riv-er sectors that harboured almost all of theforaging activity After leaving the roostthey moved consistently there without anyapparent search phase If physical featuresof water were the only factors leading to theselection of foraging sites we should expecta higher dispersion of individuals A morelikely explanation for this aggregation couldbe a patchy distribution of the availablefood among the acoustically suitable riverstretches Barclay et al (2000) found thatMyotis moluccarum foraged at 10 km fromthe roost and suggested that available foodin overlooked nearer water surfaces was in-sufficient to meet the batsrsquo needs The ex-treme aggregation observed in our studymay also be beneficial to bats throughgrouping and associated information trans-fer (Wilkinson 1995) presence of relativesandor minimization of predation risk Ifthis pattern of aggregated spatial use is re-lated to food availability we should expectvariation from aggregation to total disper-sion depending on the conditions in otherareas andor seasons Alternatively if theaggregation is determined by social factorswe should expect a similar pattern in otherareas andor seasons

Ponds were highly used by M capac-cinii in Israel although seemingly therewere no water-running rivers during thestudy period (Carmel and Safriel 1998)Despite the smooth and usually clutter-freewater surface of ponds we found that onlyone of the 40 available ponds in our studyarea was used and it was 100 m from one ofthe most used river stretches Low use ofponds located far from rivers has been re-ported in M daubentonii (Ciechanowski2002) Searching costs along the linearlycontinuous river stretches may be lowerthan over ponds because ponds are separat-ed by large unprofitable (terrestrial) areaswhilst rivers offer a landscape elementwhere both commuting and foraging arecompatible

Although water surfaces in canals werealso smooth these were the least preferredaquatic habitats likely because of the lackof substrate and the high water speed whichmight limit the development of insect com-munities Moreover speed at water surfaceis usually high in canals and this couldmake prey detection and capture by bats difficult Only one bat was detected forag-ing over a canal but the foraging lasted forseveral minutes and was performed whilethe inividual commuted along the canal tothe feeding place in a river Avoidance ofsurfaces completely covered by vegetationhas a more obvious explanation as it is vir-tually impossible for the bats to fly throughdense reed formations

Lakes have been also noted as exten-sively occupied by the species (Russo andJones 2003) There are no lakes in ourstudy area but we observed that the largereservoir of Tous located upstream in theXuacutequer River was not used It is probablethat lower water temperature delays theemergence of aquatic insects in the reser-voir

Myotis capaccinii is able to cover longdistances to reach its foraging sites as are

164 D Almenar J Aihartza U Goiti E Salsamendi and I Garin

other trawling species (Arnold et al 1998Barclay et al 2000) Recorded average and maximum distances are high comparedto other bats with similar wing shapes(Jones et al 1995) Aquatic habitats areusually arranged in a linear pattern or scat-tered along the landscape so M capacci-nii should be able to reach distant places es-pecially if available prey is not evenly dis-tributed throughout these habitats Thisability allows the species to exploit vast ar-eas around its main caves

This work presents the first publishedresults on habitat selection and spatial usefor M capaccinii providing useful informa-tion for the management of its foragingsites Aquatic habitats around 10 km of thebatsrsquo main caves must be protected espe-cially those consisting of open calm watersTransformation of low-flowing or stagnantwaters into fast-running waters could harmbat populations Canalization of riverscould reduce the availability of prey by re-ducing substrate quality and habitat diversi-ty and increasing water speed (Brookes1988) thereby negatively affecting M ca-paccinii The strong dependence of M ca-paccinii on aquatic habitats highlights theneed for involving this species in future en-vironmental impact assessment of anylarge-scale hydraulic transformations suchas big dams or water transfers betweenbasins

Eutrophication has a variable effectamong bats that linked to aquatic habitatsWhile Myotis macropus and M daubento-nii seem not to be negatively affected bythis process (Vaughan et al 1996 Law andUrquhart 2000) M dasycneme prefers toforage in waters with a low degree of eu-trophication (Van De Sijpe et al 2004)Meacutedard and Guibert (1990) associated M capaccinii with well-preserved Mediter-ranean streams and proposed that the deg-radation of this habitat (mainly by loss ofriverbank tree cover and eutrophication) is

involved in the rarefaction of the species inFrance Nevertheless Courtois (1998) didnot find any association of this species withgood-quality waters in Corsica Our resultsfit better with those in Corsica than with thescenario proposed by Meacutedard and Guibert(1990) given that the rivers in the studyarea are usually eutrophicated It is possiblethat eutrophication does not represent a sig-nificant hazard for M capaccinii

ACKNOWLEDEGMENTS

We wish to thank Christian Leidenberger for hisvaluable help in the fieldwork and also Toni AlcocerToni Castelloacute and Miguel Angel Monsalve We thankBrock Fenton and an anonymous referee for theircomments that helped to improve the manuscript andDavid Howe and Kate Johnson for kindly reviewingthe English This work was part of a LIFE project(LIFE00NATE7337) coordinated by Carlos IbaacuteZezEstacioacuten Bioloacutegica de DoZana (CSIC) and co-fundedby the Conselleria de Territori i Habitatge of the Ge-neralitat Valenciana and the European CommissionSupport was also provided to this project by theCPEMN ldquoLa Granja de El Salerrdquo and UPVEHU(through research grant 9UPV0076310-158492004) DA and ES were supported by the CSIC and UG by the Basque Government This study wasperformed with permission of the Valencian Gov-ernment

LITERATURE CITED

AEBISCHER N J P A ROBERTSON and R E KEN-WARD 1993 Compositional analysis of habitatuse from animal radio-tracking data Ecology 741313ndash1325

AHLEacuteN I 1990 Identification of bats in flight Swed-ish Society for Conservation of Nature Stock-holm 50 pp

AIHARTZA J R U GOITI D ALMENAR and I GARIN2003 Evidences of piscivory by Myotis capac-cinii (Bonaparte 1837) in Southern Iberian Pen-insula Acta Chiropterologica 5 193ndash198

ALDRIDGE H D J N 1988 Flight kinematics and en-ergetics in the little brown bat Myotis lucifugus(Chiroptera Vespertilionidae) with reference tothe influence of ground effect Journal of Zool-ogy (London) 216 507ndash518

ALDRIDGE H D J N and R M BRIGHAM 1988 Load carrying and manoeuvrability in an

Habitat of Myotis capaccinii 165

insectivorous bat a test of the 5 lsquorulersquo of radio-telemetry Journal of Mammalogy 69 379ndash382

ARNOLD A M BRAUN N BECKER and V STORCH1998 Beitrag zur Oumlkologie der Wasserfleder-maus (Myotis daubentoni) in Nordbaden Caroli-nea 56 103ndash110

BARCLAY R M R B J CHRUSZCZ and M RHODES2000 Foraging behaviour of the large-footed my-otis Myotis moluccarum (Chiroptera Vesperti-lionidae) in south-eastern Queensland AustralianJournal of Zoology 48 385ndash392

BOONMAN A M M BOONMAN F BRETSCHNEIDERand W A VAN DE GRIND 1998 Prey detection intrawling insectivorous bats duckweed affectshunting behaviour in Daubentonrsquos bat Myotisdaubentonii Behavioral Ecology and Sociobiol-ogy 44 99ndash107

BRITTON A R C G JONES J M V RAYNER A MBOONMAN and B VERBOOM 1997 Flight per-formance echolocation and foraging behaviourin pond bats Myotis dasycneme (ChiropteraVespertilionidae) Journal of Zoology (London)241 503ndash522

BROOKES A 1988 Channelized rivers perspectivesfor environmental management John Wiley andSons Chichester 342 pp

CARMEL Y and U SAFRIEL 1998 Habitat use bybats in a Mediterranean ecosystem in Israel mdashconservation implications Biological Conserva-tion 84 245ndash250

CHERRY S 1996 A comparison of confidence intervalmethods for habitat use-availability studies Jour-nal of Wildlife Management 60 653ndash658

CIECHANOWSKI M 2002 Community structure andactivity of bats (Chiroptera) over different waterbodies Mammalian Biology 67 276ndash285

COSSON E and P MEacuteDARD 1999 Murin de Capac-cinii Myotis capaccinii (Bonaparte 1837) Pp47ndash51 in Habitats et activiteacute de chasse des chi-roptPres menaceacutes en Europe synthPse de con-naissances actuelles en vue drsquoune gestion conser-vatrice (S Y ROUEacute and M BARATAUD eds) LeRhinolophe Vol Spec 136 pp

COURTOIS J 1998 Contribution B la connaissance dela reacutepartition et des caracteacuteristiques biologiquesdu murin de Capaccini (Myotis capaccinii) enCorse Arvicola 10 42ndash46

FENTON M B and W BOGDANOWICZ 2002 Rela-tionships between external morphology and for-aging behaviour bats in the genus Myotis Cana-dian Journal of Zoology 80 1004ndash1013

FINDLEY J S 1972 Phenetic relationships amongbats of the genus Myotis Systematic Zoology 2131ndash52

FLAVIN D A S S BIGGANE C B SHIEL P SMIDDY

and J S FAIRLEY 2001 Analysis of the diet ofDaubentonrsquos bat Myotis daubentonii in IrelandActa Theriologica 46 43ndash52

GARSHELIS D L 2000 Delusions in habitat evalua-tion measuring use selection and importancePp 111ndash164 in Research techniques in animalecology controversies and consequences (LBOITANI and T K FULLER eds) Columbia Uni-versity Press New York 442 pp

GOITI U J R AIHARTZA I GARIN and J ZABALA2003 Influence of habitat on the foraging behav-iour of the Mediterranean horseshoe bat Rhino-lophus euryale Acta Chiropterologica 5 75ndash84

GUILLEacuteN A 1999 Myotis capaccinii Pp 106ndash107 inThe atlas of European mammals (A J MITCHELL-JONES G AMORI W BOGDANOWICZ B KRY-ŠTUFEK P J H REIJNDERS F SPITZENBERGER M STUBBE J B M THISSEN V VOHRALIK andJ ZIMA eds) T amp AD Poyser Ltd London 484 pp

HUTSON A M S P MICKLEBURGH and P A RACEY(comp) 2001 Microchiropteran bats Global sta-tus survey and conservation action plan IUCNGland 259 pp

JONES G and J M V RAYNER 1988 Flight perfor-mance foraging tactics and echolocation in free-living Daubentonrsquos bats Myotis daubentoni (Chi-roptera Vespertilionidae) Journal of Zoology(London) 215 113ndash132

JONES G and J M V RAYNER 1991 Flight perfor-mance foraging tactics and echolocation in thetrawling insectivorous bat Myotis adversus (Chi-roptera Vespertilionidae) Journal of Zoology(London) 225 393ndash412

JONES G L DUVERGEacute and R D RANSOME 1995Conservation biology of an endangered speciesfield studies of greater horseshoe bat Pp309ndash324 in Ecology evolution and behaviour ofbats (P A RACEY and S M SWIFT eds) Sym-posia of the Zoological Society of London 67 1ndash421

KALKO E 1990 Field study on the echolocation andhunting behaviour of the long-fingered batMyotis capaccinii Bat Research News 3142ndash43

KALKO E K V H-U SCHNITZLER I KAIPF and AD GRINNELL 1998 Echolocation and foragingbehavior of the lesser bulldog bat Noctilio albi-ventris preadaptations for piscivory BehavioralEcology and Sociobiology 42 305ndash319

KUNZ T H and A KURTA 1988 Capture methodsand holding devices Pp 1ndash29 in Ecological andbehavioral methods for the study of bats (T HKUNZ ed) Smithsonian Institution Press Wash-ington DC 533 pp

166 D Almenar J Aihartza U Goiti E Salsamendi and I Garin

Habitat of Myotis capaccinii 167

LANZA B and P AGNELLI 2000 Long fingered batMyotis capaccinii (Bonaparte 1837) Pp 57ndash58in Italian mammals (M SPAGNESI S TOSO andA M DE MARINIS eds) Ministerio dellrsquoAmbien-te Servizio Conservazione della Natura and In-stituto Nazionale per la Fauna Selvatica A GhigiOzzano dellrsquoEmmilia 204 pp

LAW B and C A URQUHART 2000 Diet of the large-footed myotis Myotis macropus at a forest streamroost in northern New South Wales AustralianMammalogy 22 121ndash124

MA J G JONES S ZHANG J SHEN W METZNER L ZHANG and B LIANG 2003 Dietary analysisconfirms that Rickettrsquos big-footed bat (Myotisricketti) is a piscivore Journal of Zoology (Lon-don) 261 245ndash248

MACKEY R L and R M R BARCLAY 1989 The influence of physical clutter and noise on the ac-tivity of bats over water Canadian Journal of Zo-ology 67 1167ndash1170

MANLY B F J L L MCDONALD D L THOMAS T L MCDONALD and W P ERICKSON 2002 Re-source selection by animals Statistical designand analysis for field studies Kluwer AcademicPublishers Dordrecht 240 pp

MEacuteDARD P and E GUIBERT 1990 Disparition drsquounmilieu et rareacutefaction drsquoune espPce en France lemurin de Capaccini Myotis capaccinii (Bona-parte 1837) Mammalia 54 297ndash300

NEU C W C R BYERS and J M PEEK 1974 A technique for analysis of utilization-availab-ility data Journal of Wildlife Management 38541ndash545

NORBERG U M and J M V RAYNER 1987 Ecol-ogical morphology and flight in bats (MammaliaChiroptera) wing adaptations flight perfor-mance foraging strategy and echolocation Phil-osophical Transactions of the Royal Society ofLondon 316B 335ndash427

RAYNER J M V 1991 On the aerodynamics of ani-mal flight in ground effect Philosophical Trans-actions of the Royal Society of London 334B119ndash128

ROBSON S K 1984 Myotis adversus (ChiropteraVespertilionidae) Australiarsquos fish-eating batAustralian Mammalogy 7 51ndash52

RUEDI M and A MAYER 2001 Molecular systemat-ics of bats of the genus Myotis (Vespertilionidae)suggests deterministic ecomorphological conver-gences Molecular Phylogenetics and Evolution21 436ndash448

RUSSO D and G JONES 2003 Use of foraging habi-tats by bats in a Mediterranean area determined

by acoustic surveys conservation implicationsEcography 26 197ndash209

RYDELL J L A MILLER and M E JENSEN 1999Echolocation constraints of Daubentonrsquos bat foraging over water Functional Ecology 13247ndash255

SANCHIS-MOLL E J (ed) 1988 Guiacutea de la naturale-za de la Comunidad Valenciana Edicions Alfonsel MagnBnim Valencia 664 pp

SIEMERS B M P STILZ and H-U SCHNITZLER2001 The acoustic advantage of hunting at lowheights above water behavioural experiments onthe European lsquotrawlingrsquo bats Myotis capacciniiM dasycneme and M daubentonii Journal ofExperimental Biology 204 3843ndash3854

SIEMERS B M E BAUR and H-U SCHNITZLER2005 Acoustic mirror effect increases prey de-tection distance in trawling bats Naturwissen-schaften 92 272ndash276

SPITZENBERGER F and O VON HELVERSEN 2001Myotis capaccinii (Bonaparte 1837) mdash Langfuszlig-fledermaus Pp 281ndash302 in Handbuch der Saumlu-getiere Europas Band 4I Fledertiere I (J NIET-HAMMER and F KRAPP eds) Aula-Verlag Wies-baden 602 pp

VAN DE SIJPE M B VANDENDRIESSCHE P VOET J VANDENBERGHE J DUYCK E NAEYAERT M MANHAEVE and E MARTENS 2004 Summerdistribution of the pond bat Myotis dasycne-me (Chiroptera Vespertilionidae) in the west ofFlanders (Belgium) with regard to water qualityMammalia 68 377ndash386

VAUGHAN N G JONES and S HARRIS 1996 Effectsof sewage effluent on the activity of bats (Chi-roptera Vespertilionidae) foraging along riversBiological Conservation 78 337ndash343

VON FRENCKELL B and R M R BARCLAY 1987Bat activity over calm and turbulent water Cana-dian Journal of Zoology 65 219ndash222

WARREN R D D A WATERS J D ALTRINGHAM andD J BULLOCK 2000 The distribution of Dauben-tonrsquos bats (Myotis daubentonii) and pipistrellebats (Pipistrellus pipistrellus) (Vespertilionidae)in relation to small-scale variation in riverinehabitat Biological Conservation 92 85ndash91

WHITE G C and R A GARROT 1990 Analysis ofwildlife radio-tracking data Academic PressLondon 383 pp

WILKINSON G S 1995 Information transfer in batsPp 345ndash360 in Ecology evolution and behav-iour of bats (P A RACEY and S M SWIFT eds)Symposia of the Zoological Society of London67 1ndash421

Received 29 April 2005 accepted 13 January 2006

did not allow the identification of the kindof activity involved

Another selection pattern cannot be dis-carded where the population status or habi-tat availability are different The high popu-lation size of M capaccinii in the study areasuggests that the habitat quality is equallyhigh (sensu Garshelis 2000) In suboptimalhabitat conditions the species would possi-bly be forced to change its behaviour (Goi-ti et al 2003) The best-known species of

Habitat of Myotis capaccinii 163

Confidence intervals ProportionCategoryLower Upper available Selection

HabitatsCanal 0 0006 0111 ndashPool 0034 0071 0023 PositiveRiver 0923 0962 0866 Positive

River water surfaceSmooth 0807 0871 0737 PositiveRippled 0120 0182 0241 NegativeCovered 0001 0015 0022 Negative

TABLE 3 Selection of aquatic habitats and physical features of rivers by 17 M capaccinii following the Neu etalrsquos (1974) method The 95 Baileyrsquos confidence intervals (CI) for the proportion of use are shown and itscomparison to proportion available allows for assessing the significance of selection CI for canals is not reliablebecause the number of locations in this habitat was less than five

trawling bats are able to exploit niches other than water using different foragingstrategies (mainly aerial hawking) ie My-otis daubentonii (Jones and Rayner 1988Arnold et al 1998) M adversus (Jonesand Rayner 1991) and Noctilio albiventris(Kalko et al 1998)

Three putative factors might explain theobserved selection behaviour of M capac-cinii the water surfacesrsquo acoustical proper-ties the distribution of different kinds of

FIG 2 Use (O) versus availability (Q) of different aquatic habitats (left) and river features (right) resulting fromthe tracking of 17 M capaccinii Percentage use is measured as accumulated foraging time for all bats in each

class Percent availability is measured in an MCP encompassing all bat locations

water surfaces and prey availability Withinrivers tracked bats preferred open waterswith smooth surfaces as reported for othertrawling Myotis (Von Frenckell and Barclay1987 Boonman et al 1998 Rydell et al1999 Warren et al 2000) Siemers et al(2001) proved that smooth surfaces act aslsquoacoustic mirrorsrsquo improving target detec-tion for different Myotis including M ca-paccinii Calm waters present less high-frequency water noise interfering less inthe batsrsquo echolocating system (Von Frenc-kell and Barclay 1987 Mackey and Bar-clay 1989) In this sense target hiding bynoise and structural clutter presumablyleads to underexploitation of rippled watersurfaces in rivers Also the tracked individ-uals clumped together in one of the two riv-er sectors that harboured almost all of theforaging activity After leaving the roostthey moved consistently there without anyapparent search phase If physical featuresof water were the only factors leading to theselection of foraging sites we should expecta higher dispersion of individuals A morelikely explanation for this aggregation couldbe a patchy distribution of the availablefood among the acoustically suitable riverstretches Barclay et al (2000) found thatMyotis moluccarum foraged at 10 km fromthe roost and suggested that available foodin overlooked nearer water surfaces was in-sufficient to meet the batsrsquo needs The ex-treme aggregation observed in our studymay also be beneficial to bats throughgrouping and associated information trans-fer (Wilkinson 1995) presence of relativesandor minimization of predation risk Ifthis pattern of aggregated spatial use is re-lated to food availability we should expectvariation from aggregation to total disper-sion depending on the conditions in otherareas andor seasons Alternatively if theaggregation is determined by social factorswe should expect a similar pattern in otherareas andor seasons

Ponds were highly used by M capac-cinii in Israel although seemingly therewere no water-running rivers during thestudy period (Carmel and Safriel 1998)Despite the smooth and usually clutter-freewater surface of ponds we found that onlyone of the 40 available ponds in our studyarea was used and it was 100 m from one ofthe most used river stretches Low use ofponds located far from rivers has been re-ported in M daubentonii (Ciechanowski2002) Searching costs along the linearlycontinuous river stretches may be lowerthan over ponds because ponds are separat-ed by large unprofitable (terrestrial) areaswhilst rivers offer a landscape elementwhere both commuting and foraging arecompatible

Although water surfaces in canals werealso smooth these were the least preferredaquatic habitats likely because of the lackof substrate and the high water speed whichmight limit the development of insect com-munities Moreover speed at water surfaceis usually high in canals and this couldmake prey detection and capture by bats difficult Only one bat was detected forag-ing over a canal but the foraging lasted forseveral minutes and was performed whilethe inividual commuted along the canal tothe feeding place in a river Avoidance ofsurfaces completely covered by vegetationhas a more obvious explanation as it is vir-tually impossible for the bats to fly throughdense reed formations

Lakes have been also noted as exten-sively occupied by the species (Russo andJones 2003) There are no lakes in ourstudy area but we observed that the largereservoir of Tous located upstream in theXuacutequer River was not used It is probablethat lower water temperature delays theemergence of aquatic insects in the reser-voir

Myotis capaccinii is able to cover longdistances to reach its foraging sites as are

164 D Almenar J Aihartza U Goiti E Salsamendi and I Garin

other trawling species (Arnold et al 1998Barclay et al 2000) Recorded average and maximum distances are high comparedto other bats with similar wing shapes(Jones et al 1995) Aquatic habitats areusually arranged in a linear pattern or scat-tered along the landscape so M capacci-nii should be able to reach distant places es-pecially if available prey is not evenly dis-tributed throughout these habitats Thisability allows the species to exploit vast ar-eas around its main caves

This work presents the first publishedresults on habitat selection and spatial usefor M capaccinii providing useful informa-tion for the management of its foragingsites Aquatic habitats around 10 km of thebatsrsquo main caves must be protected espe-cially those consisting of open calm watersTransformation of low-flowing or stagnantwaters into fast-running waters could harmbat populations Canalization of riverscould reduce the availability of prey by re-ducing substrate quality and habitat diversi-ty and increasing water speed (Brookes1988) thereby negatively affecting M ca-paccinii The strong dependence of M ca-paccinii on aquatic habitats highlights theneed for involving this species in future en-vironmental impact assessment of anylarge-scale hydraulic transformations suchas big dams or water transfers betweenbasins

Eutrophication has a variable effectamong bats that linked to aquatic habitatsWhile Myotis macropus and M daubento-nii seem not to be negatively affected bythis process (Vaughan et al 1996 Law andUrquhart 2000) M dasycneme prefers toforage in waters with a low degree of eu-trophication (Van De Sijpe et al 2004)Meacutedard and Guibert (1990) associated M capaccinii with well-preserved Mediter-ranean streams and proposed that the deg-radation of this habitat (mainly by loss ofriverbank tree cover and eutrophication) is

involved in the rarefaction of the species inFrance Nevertheless Courtois (1998) didnot find any association of this species withgood-quality waters in Corsica Our resultsfit better with those in Corsica than with thescenario proposed by Meacutedard and Guibert(1990) given that the rivers in the studyarea are usually eutrophicated It is possiblethat eutrophication does not represent a sig-nificant hazard for M capaccinii

ACKNOWLEDEGMENTS

We wish to thank Christian Leidenberger for hisvaluable help in the fieldwork and also Toni AlcocerToni Castelloacute and Miguel Angel Monsalve We thankBrock Fenton and an anonymous referee for theircomments that helped to improve the manuscript andDavid Howe and Kate Johnson for kindly reviewingthe English This work was part of a LIFE project(LIFE00NATE7337) coordinated by Carlos IbaacuteZezEstacioacuten Bioloacutegica de DoZana (CSIC) and co-fundedby the Conselleria de Territori i Habitatge of the Ge-neralitat Valenciana and the European CommissionSupport was also provided to this project by theCPEMN ldquoLa Granja de El Salerrdquo and UPVEHU(through research grant 9UPV0076310-158492004) DA and ES were supported by the CSIC and UG by the Basque Government This study wasperformed with permission of the Valencian Gov-ernment

LITERATURE CITED

AEBISCHER N J P A ROBERTSON and R E KEN-WARD 1993 Compositional analysis of habitatuse from animal radio-tracking data Ecology 741313ndash1325

AHLEacuteN I 1990 Identification of bats in flight Swed-ish Society for Conservation of Nature Stock-holm 50 pp

AIHARTZA J R U GOITI D ALMENAR and I GARIN2003 Evidences of piscivory by Myotis capac-cinii (Bonaparte 1837) in Southern Iberian Pen-insula Acta Chiropterologica 5 193ndash198

ALDRIDGE H D J N 1988 Flight kinematics and en-ergetics in the little brown bat Myotis lucifugus(Chiroptera Vespertilionidae) with reference tothe influence of ground effect Journal of Zool-ogy (London) 216 507ndash518

ALDRIDGE H D J N and R M BRIGHAM 1988 Load carrying and manoeuvrability in an

Habitat of Myotis capaccinii 165

insectivorous bat a test of the 5 lsquorulersquo of radio-telemetry Journal of Mammalogy 69 379ndash382

ARNOLD A M BRAUN N BECKER and V STORCH1998 Beitrag zur Oumlkologie der Wasserfleder-maus (Myotis daubentoni) in Nordbaden Caroli-nea 56 103ndash110

BARCLAY R M R B J CHRUSZCZ and M RHODES2000 Foraging behaviour of the large-footed my-otis Myotis moluccarum (Chiroptera Vesperti-lionidae) in south-eastern Queensland AustralianJournal of Zoology 48 385ndash392

BOONMAN A M M BOONMAN F BRETSCHNEIDERand W A VAN DE GRIND 1998 Prey detection intrawling insectivorous bats duckweed affectshunting behaviour in Daubentonrsquos bat Myotisdaubentonii Behavioral Ecology and Sociobiol-ogy 44 99ndash107

BRITTON A R C G JONES J M V RAYNER A MBOONMAN and B VERBOOM 1997 Flight per-formance echolocation and foraging behaviourin pond bats Myotis dasycneme (ChiropteraVespertilionidae) Journal of Zoology (London)241 503ndash522

BROOKES A 1988 Channelized rivers perspectivesfor environmental management John Wiley andSons Chichester 342 pp

CARMEL Y and U SAFRIEL 1998 Habitat use bybats in a Mediterranean ecosystem in Israel mdashconservation implications Biological Conserva-tion 84 245ndash250

CHERRY S 1996 A comparison of confidence intervalmethods for habitat use-availability studies Jour-nal of Wildlife Management 60 653ndash658

CIECHANOWSKI M 2002 Community structure andactivity of bats (Chiroptera) over different waterbodies Mammalian Biology 67 276ndash285

COSSON E and P MEacuteDARD 1999 Murin de Capac-cinii Myotis capaccinii (Bonaparte 1837) Pp47ndash51 in Habitats et activiteacute de chasse des chi-roptPres menaceacutes en Europe synthPse de con-naissances actuelles en vue drsquoune gestion conser-vatrice (S Y ROUEacute and M BARATAUD eds) LeRhinolophe Vol Spec 136 pp

COURTOIS J 1998 Contribution B la connaissance dela reacutepartition et des caracteacuteristiques biologiquesdu murin de Capaccini (Myotis capaccinii) enCorse Arvicola 10 42ndash46

FENTON M B and W BOGDANOWICZ 2002 Rela-tionships between external morphology and for-aging behaviour bats in the genus Myotis Cana-dian Journal of Zoology 80 1004ndash1013

FINDLEY J S 1972 Phenetic relationships amongbats of the genus Myotis Systematic Zoology 2131ndash52

FLAVIN D A S S BIGGANE C B SHIEL P SMIDDY

and J S FAIRLEY 2001 Analysis of the diet ofDaubentonrsquos bat Myotis daubentonii in IrelandActa Theriologica 46 43ndash52

GARSHELIS D L 2000 Delusions in habitat evalua-tion measuring use selection and importancePp 111ndash164 in Research techniques in animalecology controversies and consequences (LBOITANI and T K FULLER eds) Columbia Uni-versity Press New York 442 pp

GOITI U J R AIHARTZA I GARIN and J ZABALA2003 Influence of habitat on the foraging behav-iour of the Mediterranean horseshoe bat Rhino-lophus euryale Acta Chiropterologica 5 75ndash84

GUILLEacuteN A 1999 Myotis capaccinii Pp 106ndash107 inThe atlas of European mammals (A J MITCHELL-JONES G AMORI W BOGDANOWICZ B KRY-ŠTUFEK P J H REIJNDERS F SPITZENBERGER M STUBBE J B M THISSEN V VOHRALIK andJ ZIMA eds) T amp AD Poyser Ltd London 484 pp

HUTSON A M S P MICKLEBURGH and P A RACEY(comp) 2001 Microchiropteran bats Global sta-tus survey and conservation action plan IUCNGland 259 pp

JONES G and J M V RAYNER 1988 Flight perfor-mance foraging tactics and echolocation in free-living Daubentonrsquos bats Myotis daubentoni (Chi-roptera Vespertilionidae) Journal of Zoology(London) 215 113ndash132

JONES G and J M V RAYNER 1991 Flight perfor-mance foraging tactics and echolocation in thetrawling insectivorous bat Myotis adversus (Chi-roptera Vespertilionidae) Journal of Zoology(London) 225 393ndash412

JONES G L DUVERGEacute and R D RANSOME 1995Conservation biology of an endangered speciesfield studies of greater horseshoe bat Pp309ndash324 in Ecology evolution and behaviour ofbats (P A RACEY and S M SWIFT eds) Sym-posia of the Zoological Society of London 67 1ndash421

KALKO E 1990 Field study on the echolocation andhunting behaviour of the long-fingered batMyotis capaccinii Bat Research News 3142ndash43

KALKO E K V H-U SCHNITZLER I KAIPF and AD GRINNELL 1998 Echolocation and foragingbehavior of the lesser bulldog bat Noctilio albi-ventris preadaptations for piscivory BehavioralEcology and Sociobiology 42 305ndash319

KUNZ T H and A KURTA 1988 Capture methodsand holding devices Pp 1ndash29 in Ecological andbehavioral methods for the study of bats (T HKUNZ ed) Smithsonian Institution Press Wash-ington DC 533 pp

166 D Almenar J Aihartza U Goiti E Salsamendi and I Garin

Habitat of Myotis capaccinii 167

LANZA B and P AGNELLI 2000 Long fingered batMyotis capaccinii (Bonaparte 1837) Pp 57ndash58in Italian mammals (M SPAGNESI S TOSO andA M DE MARINIS eds) Ministerio dellrsquoAmbien-te Servizio Conservazione della Natura and In-stituto Nazionale per la Fauna Selvatica A GhigiOzzano dellrsquoEmmilia 204 pp

LAW B and C A URQUHART 2000 Diet of the large-footed myotis Myotis macropus at a forest streamroost in northern New South Wales AustralianMammalogy 22 121ndash124

MA J G JONES S ZHANG J SHEN W METZNER L ZHANG and B LIANG 2003 Dietary analysisconfirms that Rickettrsquos big-footed bat (Myotisricketti) is a piscivore Journal of Zoology (Lon-don) 261 245ndash248

MACKEY R L and R M R BARCLAY 1989 The influence of physical clutter and noise on the ac-tivity of bats over water Canadian Journal of Zo-ology 67 1167ndash1170

MANLY B F J L L MCDONALD D L THOMAS T L MCDONALD and W P ERICKSON 2002 Re-source selection by animals Statistical designand analysis for field studies Kluwer AcademicPublishers Dordrecht 240 pp

MEacuteDARD P and E GUIBERT 1990 Disparition drsquounmilieu et rareacutefaction drsquoune espPce en France lemurin de Capaccini Myotis capaccinii (Bona-parte 1837) Mammalia 54 297ndash300

NEU C W C R BYERS and J M PEEK 1974 A technique for analysis of utilization-availab-ility data Journal of Wildlife Management 38541ndash545

NORBERG U M and J M V RAYNER 1987 Ecol-ogical morphology and flight in bats (MammaliaChiroptera) wing adaptations flight perfor-mance foraging strategy and echolocation Phil-osophical Transactions of the Royal Society ofLondon 316B 335ndash427

RAYNER J M V 1991 On the aerodynamics of ani-mal flight in ground effect Philosophical Trans-actions of the Royal Society of London 334B119ndash128

ROBSON S K 1984 Myotis adversus (ChiropteraVespertilionidae) Australiarsquos fish-eating batAustralian Mammalogy 7 51ndash52

RUEDI M and A MAYER 2001 Molecular systemat-ics of bats of the genus Myotis (Vespertilionidae)suggests deterministic ecomorphological conver-gences Molecular Phylogenetics and Evolution21 436ndash448

RUSSO D and G JONES 2003 Use of foraging habi-tats by bats in a Mediterranean area determined

by acoustic surveys conservation implicationsEcography 26 197ndash209

RYDELL J L A MILLER and M E JENSEN 1999Echolocation constraints of Daubentonrsquos bat foraging over water Functional Ecology 13247ndash255

SANCHIS-MOLL E J (ed) 1988 Guiacutea de la naturale-za de la Comunidad Valenciana Edicions Alfonsel MagnBnim Valencia 664 pp

SIEMERS B M P STILZ and H-U SCHNITZLER2001 The acoustic advantage of hunting at lowheights above water behavioural experiments onthe European lsquotrawlingrsquo bats Myotis capacciniiM dasycneme and M daubentonii Journal ofExperimental Biology 204 3843ndash3854

SIEMERS B M E BAUR and H-U SCHNITZLER2005 Acoustic mirror effect increases prey de-tection distance in trawling bats Naturwissen-schaften 92 272ndash276

SPITZENBERGER F and O VON HELVERSEN 2001Myotis capaccinii (Bonaparte 1837) mdash Langfuszlig-fledermaus Pp 281ndash302 in Handbuch der Saumlu-getiere Europas Band 4I Fledertiere I (J NIET-HAMMER and F KRAPP eds) Aula-Verlag Wies-baden 602 pp

VAN DE SIJPE M B VANDENDRIESSCHE P VOET J VANDENBERGHE J DUYCK E NAEYAERT M MANHAEVE and E MARTENS 2004 Summerdistribution of the pond bat Myotis dasycne-me (Chiroptera Vespertilionidae) in the west ofFlanders (Belgium) with regard to water qualityMammalia 68 377ndash386

VAUGHAN N G JONES and S HARRIS 1996 Effectsof sewage effluent on the activity of bats (Chi-roptera Vespertilionidae) foraging along riversBiological Conservation 78 337ndash343

VON FRENCKELL B and R M R BARCLAY 1987Bat activity over calm and turbulent water Cana-dian Journal of Zoology 65 219ndash222

WARREN R D D A WATERS J D ALTRINGHAM andD J BULLOCK 2000 The distribution of Dauben-tonrsquos bats (Myotis daubentonii) and pipistrellebats (Pipistrellus pipistrellus) (Vespertilionidae)in relation to small-scale variation in riverinehabitat Biological Conservation 92 85ndash91

WHITE G C and R A GARROT 1990 Analysis ofwildlife radio-tracking data Academic PressLondon 383 pp

WILKINSON G S 1995 Information transfer in batsPp 345ndash360 in Ecology evolution and behav-iour of bats (P A RACEY and S M SWIFT eds)Symposia of the Zoological Society of London67 1ndash421

Received 29 April 2005 accepted 13 January 2006

water surfaces and prey availability Withinrivers tracked bats preferred open waterswith smooth surfaces as reported for othertrawling Myotis (Von Frenckell and Barclay1987 Boonman et al 1998 Rydell et al1999 Warren et al 2000) Siemers et al(2001) proved that smooth surfaces act aslsquoacoustic mirrorsrsquo improving target detec-tion for different Myotis including M ca-paccinii Calm waters present less high-frequency water noise interfering less inthe batsrsquo echolocating system (Von Frenc-kell and Barclay 1987 Mackey and Bar-clay 1989) In this sense target hiding bynoise and structural clutter presumablyleads to underexploitation of rippled watersurfaces in rivers Also the tracked individ-uals clumped together in one of the two riv-er sectors that harboured almost all of theforaging activity After leaving the roostthey moved consistently there without anyapparent search phase If physical featuresof water were the only factors leading to theselection of foraging sites we should expecta higher dispersion of individuals A morelikely explanation for this aggregation couldbe a patchy distribution of the availablefood among the acoustically suitable riverstretches Barclay et al (2000) found thatMyotis moluccarum foraged at 10 km fromthe roost and suggested that available foodin overlooked nearer water surfaces was in-sufficient to meet the batsrsquo needs The ex-treme aggregation observed in our studymay also be beneficial to bats throughgrouping and associated information trans-fer (Wilkinson 1995) presence of relativesandor minimization of predation risk Ifthis pattern of aggregated spatial use is re-lated to food availability we should expectvariation from aggregation to total disper-sion depending on the conditions in otherareas andor seasons Alternatively if theaggregation is determined by social factorswe should expect a similar pattern in otherareas andor seasons

Ponds were highly used by M capac-cinii in Israel although seemingly therewere no water-running rivers during thestudy period (Carmel and Safriel 1998)Despite the smooth and usually clutter-freewater surface of ponds we found that onlyone of the 40 available ponds in our studyarea was used and it was 100 m from one ofthe most used river stretches Low use ofponds located far from rivers has been re-ported in M daubentonii (Ciechanowski2002) Searching costs along the linearlycontinuous river stretches may be lowerthan over ponds because ponds are separat-ed by large unprofitable (terrestrial) areaswhilst rivers offer a landscape elementwhere both commuting and foraging arecompatible

Although water surfaces in canals werealso smooth these were the least preferredaquatic habitats likely because of the lackof substrate and the high water speed whichmight limit the development of insect com-munities Moreover speed at water surfaceis usually high in canals and this couldmake prey detection and capture by bats difficult Only one bat was detected forag-ing over a canal but the foraging lasted forseveral minutes and was performed whilethe inividual commuted along the canal tothe feeding place in a river Avoidance ofsurfaces completely covered by vegetationhas a more obvious explanation as it is vir-tually impossible for the bats to fly throughdense reed formations

Lakes have been also noted as exten-sively occupied by the species (Russo andJones 2003) There are no lakes in ourstudy area but we observed that the largereservoir of Tous located upstream in theXuacutequer River was not used It is probablethat lower water temperature delays theemergence of aquatic insects in the reser-voir

Myotis capaccinii is able to cover longdistances to reach its foraging sites as are

164 D Almenar J Aihartza U Goiti E Salsamendi and I Garin

other trawling species (Arnold et al 1998Barclay et al 2000) Recorded average and maximum distances are high comparedto other bats with similar wing shapes(Jones et al 1995) Aquatic habitats areusually arranged in a linear pattern or scat-tered along the landscape so M capacci-nii should be able to reach distant places es-pecially if available prey is not evenly dis-tributed throughout these habitats Thisability allows the species to exploit vast ar-eas around its main caves

This work presents the first publishedresults on habitat selection and spatial usefor M capaccinii providing useful informa-tion for the management of its foragingsites Aquatic habitats around 10 km of thebatsrsquo main caves must be protected espe-cially those consisting of open calm watersTransformation of low-flowing or stagnantwaters into fast-running waters could harmbat populations Canalization of riverscould reduce the availability of prey by re-ducing substrate quality and habitat diversi-ty and increasing water speed (Brookes1988) thereby negatively affecting M ca-paccinii The strong dependence of M ca-paccinii on aquatic habitats highlights theneed for involving this species in future en-vironmental impact assessment of anylarge-scale hydraulic transformations suchas big dams or water transfers betweenbasins

Eutrophication has a variable effectamong bats that linked to aquatic habitatsWhile Myotis macropus and M daubento-nii seem not to be negatively affected bythis process (Vaughan et al 1996 Law andUrquhart 2000) M dasycneme prefers toforage in waters with a low degree of eu-trophication (Van De Sijpe et al 2004)Meacutedard and Guibert (1990) associated M capaccinii with well-preserved Mediter-ranean streams and proposed that the deg-radation of this habitat (mainly by loss ofriverbank tree cover and eutrophication) is

involved in the rarefaction of the species inFrance Nevertheless Courtois (1998) didnot find any association of this species withgood-quality waters in Corsica Our resultsfit better with those in Corsica than with thescenario proposed by Meacutedard and Guibert(1990) given that the rivers in the studyarea are usually eutrophicated It is possiblethat eutrophication does not represent a sig-nificant hazard for M capaccinii

ACKNOWLEDEGMENTS

We wish to thank Christian Leidenberger for hisvaluable help in the fieldwork and also Toni AlcocerToni Castelloacute and Miguel Angel Monsalve We thankBrock Fenton and an anonymous referee for theircomments that helped to improve the manuscript andDavid Howe and Kate Johnson for kindly reviewingthe English This work was part of a LIFE project(LIFE00NATE7337) coordinated by Carlos IbaacuteZezEstacioacuten Bioloacutegica de DoZana (CSIC) and co-fundedby the Conselleria de Territori i Habitatge of the Ge-neralitat Valenciana and the European CommissionSupport was also provided to this project by theCPEMN ldquoLa Granja de El Salerrdquo and UPVEHU(through research grant 9UPV0076310-158492004) DA and ES were supported by the CSIC and UG by the Basque Government This study wasperformed with permission of the Valencian Gov-ernment

LITERATURE CITED

AEBISCHER N J P A ROBERTSON and R E KEN-WARD 1993 Compositional analysis of habitatuse from animal radio-tracking data Ecology 741313ndash1325

AHLEacuteN I 1990 Identification of bats in flight Swed-ish Society for Conservation of Nature Stock-holm 50 pp

AIHARTZA J R U GOITI D ALMENAR and I GARIN2003 Evidences of piscivory by Myotis capac-cinii (Bonaparte 1837) in Southern Iberian Pen-insula Acta Chiropterologica 5 193ndash198

ALDRIDGE H D J N 1988 Flight kinematics and en-ergetics in the little brown bat Myotis lucifugus(Chiroptera Vespertilionidae) with reference tothe influence of ground effect Journal of Zool-ogy (London) 216 507ndash518

ALDRIDGE H D J N and R M BRIGHAM 1988 Load carrying and manoeuvrability in an

Habitat of Myotis capaccinii 165

insectivorous bat a test of the 5 lsquorulersquo of radio-telemetry Journal of Mammalogy 69 379ndash382

ARNOLD A M BRAUN N BECKER and V STORCH1998 Beitrag zur Oumlkologie der Wasserfleder-maus (Myotis daubentoni) in Nordbaden Caroli-nea 56 103ndash110

BARCLAY R M R B J CHRUSZCZ and M RHODES2000 Foraging behaviour of the large-footed my-otis Myotis moluccarum (Chiroptera Vesperti-lionidae) in south-eastern Queensland AustralianJournal of Zoology 48 385ndash392

BOONMAN A M M BOONMAN F BRETSCHNEIDERand W A VAN DE GRIND 1998 Prey detection intrawling insectivorous bats duckweed affectshunting behaviour in Daubentonrsquos bat Myotisdaubentonii Behavioral Ecology and Sociobiol-ogy 44 99ndash107

BRITTON A R C G JONES J M V RAYNER A MBOONMAN and B VERBOOM 1997 Flight per-formance echolocation and foraging behaviourin pond bats Myotis dasycneme (ChiropteraVespertilionidae) Journal of Zoology (London)241 503ndash522

BROOKES A 1988 Channelized rivers perspectivesfor environmental management John Wiley andSons Chichester 342 pp

CARMEL Y and U SAFRIEL 1998 Habitat use bybats in a Mediterranean ecosystem in Israel mdashconservation implications Biological Conserva-tion 84 245ndash250

CHERRY S 1996 A comparison of confidence intervalmethods for habitat use-availability studies Jour-nal of Wildlife Management 60 653ndash658

CIECHANOWSKI M 2002 Community structure andactivity of bats (Chiroptera) over different waterbodies Mammalian Biology 67 276ndash285

COSSON E and P MEacuteDARD 1999 Murin de Capac-cinii Myotis capaccinii (Bonaparte 1837) Pp47ndash51 in Habitats et activiteacute de chasse des chi-roptPres menaceacutes en Europe synthPse de con-naissances actuelles en vue drsquoune gestion conser-vatrice (S Y ROUEacute and M BARATAUD eds) LeRhinolophe Vol Spec 136 pp

COURTOIS J 1998 Contribution B la connaissance dela reacutepartition et des caracteacuteristiques biologiquesdu murin de Capaccini (Myotis capaccinii) enCorse Arvicola 10 42ndash46

FENTON M B and W BOGDANOWICZ 2002 Rela-tionships between external morphology and for-aging behaviour bats in the genus Myotis Cana-dian Journal of Zoology 80 1004ndash1013

FINDLEY J S 1972 Phenetic relationships amongbats of the genus Myotis Systematic Zoology 2131ndash52

FLAVIN D A S S BIGGANE C B SHIEL P SMIDDY

and J S FAIRLEY 2001 Analysis of the diet ofDaubentonrsquos bat Myotis daubentonii in IrelandActa Theriologica 46 43ndash52

GARSHELIS D L 2000 Delusions in habitat evalua-tion measuring use selection and importancePp 111ndash164 in Research techniques in animalecology controversies and consequences (LBOITANI and T K FULLER eds) Columbia Uni-versity Press New York 442 pp

GOITI U J R AIHARTZA I GARIN and J ZABALA2003 Influence of habitat on the foraging behav-iour of the Mediterranean horseshoe bat Rhino-lophus euryale Acta Chiropterologica 5 75ndash84

GUILLEacuteN A 1999 Myotis capaccinii Pp 106ndash107 inThe atlas of European mammals (A J MITCHELL-JONES G AMORI W BOGDANOWICZ B KRY-ŠTUFEK P J H REIJNDERS F SPITZENBERGER M STUBBE J B M THISSEN V VOHRALIK andJ ZIMA eds) T amp AD Poyser Ltd London 484 pp

HUTSON A M S P MICKLEBURGH and P A RACEY(comp) 2001 Microchiropteran bats Global sta-tus survey and conservation action plan IUCNGland 259 pp

JONES G and J M V RAYNER 1988 Flight perfor-mance foraging tactics and echolocation in free-living Daubentonrsquos bats Myotis daubentoni (Chi-roptera Vespertilionidae) Journal of Zoology(London) 215 113ndash132

JONES G and J M V RAYNER 1991 Flight perfor-mance foraging tactics and echolocation in thetrawling insectivorous bat Myotis adversus (Chi-roptera Vespertilionidae) Journal of Zoology(London) 225 393ndash412

JONES G L DUVERGEacute and R D RANSOME 1995Conservation biology of an endangered speciesfield studies of greater horseshoe bat Pp309ndash324 in Ecology evolution and behaviour ofbats (P A RACEY and S M SWIFT eds) Sym-posia of the Zoological Society of London 67 1ndash421

KALKO E 1990 Field study on the echolocation andhunting behaviour of the long-fingered batMyotis capaccinii Bat Research News 3142ndash43

KALKO E K V H-U SCHNITZLER I KAIPF and AD GRINNELL 1998 Echolocation and foragingbehavior of the lesser bulldog bat Noctilio albi-ventris preadaptations for piscivory BehavioralEcology and Sociobiology 42 305ndash319

KUNZ T H and A KURTA 1988 Capture methodsand holding devices Pp 1ndash29 in Ecological andbehavioral methods for the study of bats (T HKUNZ ed) Smithsonian Institution Press Wash-ington DC 533 pp

166 D Almenar J Aihartza U Goiti E Salsamendi and I Garin

Habitat of Myotis capaccinii 167

LANZA B and P AGNELLI 2000 Long fingered batMyotis capaccinii (Bonaparte 1837) Pp 57ndash58in Italian mammals (M SPAGNESI S TOSO andA M DE MARINIS eds) Ministerio dellrsquoAmbien-te Servizio Conservazione della Natura and In-stituto Nazionale per la Fauna Selvatica A GhigiOzzano dellrsquoEmmilia 204 pp

LAW B and C A URQUHART 2000 Diet of the large-footed myotis Myotis macropus at a forest streamroost in northern New South Wales AustralianMammalogy 22 121ndash124

MA J G JONES S ZHANG J SHEN W METZNER L ZHANG and B LIANG 2003 Dietary analysisconfirms that Rickettrsquos big-footed bat (Myotisricketti) is a piscivore Journal of Zoology (Lon-don) 261 245ndash248

MACKEY R L and R M R BARCLAY 1989 The influence of physical clutter and noise on the ac-tivity of bats over water Canadian Journal of Zo-ology 67 1167ndash1170

MANLY B F J L L MCDONALD D L THOMAS T L MCDONALD and W P ERICKSON 2002 Re-source selection by animals Statistical designand analysis for field studies Kluwer AcademicPublishers Dordrecht 240 pp

MEacuteDARD P and E GUIBERT 1990 Disparition drsquounmilieu et rareacutefaction drsquoune espPce en France lemurin de Capaccini Myotis capaccinii (Bona-parte 1837) Mammalia 54 297ndash300

NEU C W C R BYERS and J M PEEK 1974 A technique for analysis of utilization-availab-ility data Journal of Wildlife Management 38541ndash545

NORBERG U M and J M V RAYNER 1987 Ecol-ogical morphology and flight in bats (MammaliaChiroptera) wing adaptations flight perfor-mance foraging strategy and echolocation Phil-osophical Transactions of the Royal Society ofLondon 316B 335ndash427

RAYNER J M V 1991 On the aerodynamics of ani-mal flight in ground effect Philosophical Trans-actions of the Royal Society of London 334B119ndash128

ROBSON S K 1984 Myotis adversus (ChiropteraVespertilionidae) Australiarsquos fish-eating batAustralian Mammalogy 7 51ndash52

RUEDI M and A MAYER 2001 Molecular systemat-ics of bats of the genus Myotis (Vespertilionidae)suggests deterministic ecomorphological conver-gences Molecular Phylogenetics and Evolution21 436ndash448

RUSSO D and G JONES 2003 Use of foraging habi-tats by bats in a Mediterranean area determined

by acoustic surveys conservation implicationsEcography 26 197ndash209

RYDELL J L A MILLER and M E JENSEN 1999Echolocation constraints of Daubentonrsquos bat foraging over water Functional Ecology 13247ndash255

SANCHIS-MOLL E J (ed) 1988 Guiacutea de la naturale-za de la Comunidad Valenciana Edicions Alfonsel MagnBnim Valencia 664 pp

SIEMERS B M P STILZ and H-U SCHNITZLER2001 The acoustic advantage of hunting at lowheights above water behavioural experiments onthe European lsquotrawlingrsquo bats Myotis capacciniiM dasycneme and M daubentonii Journal ofExperimental Biology 204 3843ndash3854

SIEMERS B M E BAUR and H-U SCHNITZLER2005 Acoustic mirror effect increases prey de-tection distance in trawling bats Naturwissen-schaften 92 272ndash276

SPITZENBERGER F and O VON HELVERSEN 2001Myotis capaccinii (Bonaparte 1837) mdash Langfuszlig-fledermaus Pp 281ndash302 in Handbuch der Saumlu-getiere Europas Band 4I Fledertiere I (J NIET-HAMMER and F KRAPP eds) Aula-Verlag Wies-baden 602 pp

VAN DE SIJPE M B VANDENDRIESSCHE P VOET J VANDENBERGHE J DUYCK E NAEYAERT M MANHAEVE and E MARTENS 2004 Summerdistribution of the pond bat Myotis dasycne-me (Chiroptera Vespertilionidae) in the west ofFlanders (Belgium) with regard to water qualityMammalia 68 377ndash386

VAUGHAN N G JONES and S HARRIS 1996 Effectsof sewage effluent on the activity of bats (Chi-roptera Vespertilionidae) foraging along riversBiological Conservation 78 337ndash343

VON FRENCKELL B and R M R BARCLAY 1987Bat activity over calm and turbulent water Cana-dian Journal of Zoology 65 219ndash222

WARREN R D D A WATERS J D ALTRINGHAM andD J BULLOCK 2000 The distribution of Dauben-tonrsquos bats (Myotis daubentonii) and pipistrellebats (Pipistrellus pipistrellus) (Vespertilionidae)in relation to small-scale variation in riverinehabitat Biological Conservation 92 85ndash91

WHITE G C and R A GARROT 1990 Analysis ofwildlife radio-tracking data Academic PressLondon 383 pp

WILKINSON G S 1995 Information transfer in batsPp 345ndash360 in Ecology evolution and behav-iour of bats (P A RACEY and S M SWIFT eds)Symposia of the Zoological Society of London67 1ndash421

Received 29 April 2005 accepted 13 January 2006

other trawling species (Arnold et al 1998Barclay et al 2000) Recorded average and maximum distances are high comparedto other bats with similar wing shapes(Jones et al 1995) Aquatic habitats areusually arranged in a linear pattern or scat-tered along the landscape so M capacci-nii should be able to reach distant places es-pecially if available prey is not evenly dis-tributed throughout these habitats Thisability allows the species to exploit vast ar-eas around its main caves

This work presents the first publishedresults on habitat selection and spatial usefor M capaccinii providing useful informa-tion for the management of its foragingsites Aquatic habitats around 10 km of thebatsrsquo main caves must be protected espe-cially those consisting of open calm watersTransformation of low-flowing or stagnantwaters into fast-running waters could harmbat populations Canalization of riverscould reduce the availability of prey by re-ducing substrate quality and habitat diversi-ty and increasing water speed (Brookes1988) thereby negatively affecting M ca-paccinii The strong dependence of M ca-paccinii on aquatic habitats highlights theneed for involving this species in future en-vironmental impact assessment of anylarge-scale hydraulic transformations suchas big dams or water transfers betweenbasins

Eutrophication has a variable effectamong bats that linked to aquatic habitatsWhile Myotis macropus and M daubento-nii seem not to be negatively affected bythis process (Vaughan et al 1996 Law andUrquhart 2000) M dasycneme prefers toforage in waters with a low degree of eu-trophication (Van De Sijpe et al 2004)Meacutedard and Guibert (1990) associated M capaccinii with well-preserved Mediter-ranean streams and proposed that the deg-radation of this habitat (mainly by loss ofriverbank tree cover and eutrophication) is

involved in the rarefaction of the species inFrance Nevertheless Courtois (1998) didnot find any association of this species withgood-quality waters in Corsica Our resultsfit better with those in Corsica than with thescenario proposed by Meacutedard and Guibert(1990) given that the rivers in the studyarea are usually eutrophicated It is possiblethat eutrophication does not represent a sig-nificant hazard for M capaccinii

ACKNOWLEDEGMENTS

We wish to thank Christian Leidenberger for hisvaluable help in the fieldwork and also Toni AlcocerToni Castelloacute and Miguel Angel Monsalve We thankBrock Fenton and an anonymous referee for theircomments that helped to improve the manuscript andDavid Howe and Kate Johnson for kindly reviewingthe English This work was part of a LIFE project(LIFE00NATE7337) coordinated by Carlos IbaacuteZezEstacioacuten Bioloacutegica de DoZana (CSIC) and co-fundedby the Conselleria de Territori i Habitatge of the Ge-neralitat Valenciana and the European CommissionSupport was also provided to this project by theCPEMN ldquoLa Granja de El Salerrdquo and UPVEHU(through research grant 9UPV0076310-158492004) DA and ES were supported by the CSIC and UG by the Basque Government This study wasperformed with permission of the Valencian Gov-ernment

LITERATURE CITED

AEBISCHER N J P A ROBERTSON and R E KEN-WARD 1993 Compositional analysis of habitatuse from animal radio-tracking data Ecology 741313ndash1325

AHLEacuteN I 1990 Identification of bats in flight Swed-ish Society for Conservation of Nature Stock-holm 50 pp

AIHARTZA J R U GOITI D ALMENAR and I GARIN2003 Evidences of piscivory by Myotis capac-cinii (Bonaparte 1837) in Southern Iberian Pen-insula Acta Chiropterologica 5 193ndash198

ALDRIDGE H D J N 1988 Flight kinematics and en-ergetics in the little brown bat Myotis lucifugus(Chiroptera Vespertilionidae) with reference tothe influence of ground effect Journal of Zool-ogy (London) 216 507ndash518

ALDRIDGE H D J N and R M BRIGHAM 1988 Load carrying and manoeuvrability in an

Habitat of Myotis capaccinii 165

insectivorous bat a test of the 5 lsquorulersquo of radio-telemetry Journal of Mammalogy 69 379ndash382

ARNOLD A M BRAUN N BECKER and V STORCH1998 Beitrag zur Oumlkologie der Wasserfleder-maus (Myotis daubentoni) in Nordbaden Caroli-nea 56 103ndash110

BARCLAY R M R B J CHRUSZCZ and M RHODES2000 Foraging behaviour of the large-footed my-otis Myotis moluccarum (Chiroptera Vesperti-lionidae) in south-eastern Queensland AustralianJournal of Zoology 48 385ndash392

BOONMAN A M M BOONMAN F BRETSCHNEIDERand W A VAN DE GRIND 1998 Prey detection intrawling insectivorous bats duckweed affectshunting behaviour in Daubentonrsquos bat Myotisdaubentonii Behavioral Ecology and Sociobiol-ogy 44 99ndash107

BRITTON A R C G JONES J M V RAYNER A MBOONMAN and B VERBOOM 1997 Flight per-formance echolocation and foraging behaviourin pond bats Myotis dasycneme (ChiropteraVespertilionidae) Journal of Zoology (London)241 503ndash522

BROOKES A 1988 Channelized rivers perspectivesfor environmental management John Wiley andSons Chichester 342 pp

CARMEL Y and U SAFRIEL 1998 Habitat use bybats in a Mediterranean ecosystem in Israel mdashconservation implications Biological Conserva-tion 84 245ndash250

CHERRY S 1996 A comparison of confidence intervalmethods for habitat use-availability studies Jour-nal of Wildlife Management 60 653ndash658

CIECHANOWSKI M 2002 Community structure andactivity of bats (Chiroptera) over different waterbodies Mammalian Biology 67 276ndash285

COSSON E and P MEacuteDARD 1999 Murin de Capac-cinii Myotis capaccinii (Bonaparte 1837) Pp47ndash51 in Habitats et activiteacute de chasse des chi-roptPres menaceacutes en Europe synthPse de con-naissances actuelles en vue drsquoune gestion conser-vatrice (S Y ROUEacute and M BARATAUD eds) LeRhinolophe Vol Spec 136 pp

COURTOIS J 1998 Contribution B la connaissance dela reacutepartition et des caracteacuteristiques biologiquesdu murin de Capaccini (Myotis capaccinii) enCorse Arvicola 10 42ndash46

FENTON M B and W BOGDANOWICZ 2002 Rela-tionships between external morphology and for-aging behaviour bats in the genus Myotis Cana-dian Journal of Zoology 80 1004ndash1013

FINDLEY J S 1972 Phenetic relationships amongbats of the genus Myotis Systematic Zoology 2131ndash52

FLAVIN D A S S BIGGANE C B SHIEL P SMIDDY

and J S FAIRLEY 2001 Analysis of the diet ofDaubentonrsquos bat Myotis daubentonii in IrelandActa Theriologica 46 43ndash52

GARSHELIS D L 2000 Delusions in habitat evalua-tion measuring use selection and importancePp 111ndash164 in Research techniques in animalecology controversies and consequences (LBOITANI and T K FULLER eds) Columbia Uni-versity Press New York 442 pp

GOITI U J R AIHARTZA I GARIN and J ZABALA2003 Influence of habitat on the foraging behav-iour of the Mediterranean horseshoe bat Rhino-lophus euryale Acta Chiropterologica 5 75ndash84

GUILLEacuteN A 1999 Myotis capaccinii Pp 106ndash107 inThe atlas of European mammals (A J MITCHELL-JONES G AMORI W BOGDANOWICZ B KRY-ŠTUFEK P J H REIJNDERS F SPITZENBERGER M STUBBE J B M THISSEN V VOHRALIK andJ ZIMA eds) T amp AD Poyser Ltd London 484 pp

HUTSON A M S P MICKLEBURGH and P A RACEY(comp) 2001 Microchiropteran bats Global sta-tus survey and conservation action plan IUCNGland 259 pp

JONES G and J M V RAYNER 1988 Flight perfor-mance foraging tactics and echolocation in free-living Daubentonrsquos bats Myotis daubentoni (Chi-roptera Vespertilionidae) Journal of Zoology(London) 215 113ndash132

JONES G and J M V RAYNER 1991 Flight perfor-mance foraging tactics and echolocation in thetrawling insectivorous bat Myotis adversus (Chi-roptera Vespertilionidae) Journal of Zoology(London) 225 393ndash412

JONES G L DUVERGEacute and R D RANSOME 1995Conservation biology of an endangered speciesfield studies of greater horseshoe bat Pp309ndash324 in Ecology evolution and behaviour ofbats (P A RACEY and S M SWIFT eds) Sym-posia of the Zoological Society of London 67 1ndash421

KALKO E 1990 Field study on the echolocation andhunting behaviour of the long-fingered batMyotis capaccinii Bat Research News 3142ndash43

KALKO E K V H-U SCHNITZLER I KAIPF and AD GRINNELL 1998 Echolocation and foragingbehavior of the lesser bulldog bat Noctilio albi-ventris preadaptations for piscivory BehavioralEcology and Sociobiology 42 305ndash319

KUNZ T H and A KURTA 1988 Capture methodsand holding devices Pp 1ndash29 in Ecological andbehavioral methods for the study of bats (T HKUNZ ed) Smithsonian Institution Press Wash-ington DC 533 pp

166 D Almenar J Aihartza U Goiti E Salsamendi and I Garin

Habitat of Myotis capaccinii 167

LANZA B and P AGNELLI 2000 Long fingered batMyotis capaccinii (Bonaparte 1837) Pp 57ndash58in Italian mammals (M SPAGNESI S TOSO andA M DE MARINIS eds) Ministerio dellrsquoAmbien-te Servizio Conservazione della Natura and In-stituto Nazionale per la Fauna Selvatica A GhigiOzzano dellrsquoEmmilia 204 pp

LAW B and C A URQUHART 2000 Diet of the large-footed myotis Myotis macropus at a forest streamroost in northern New South Wales AustralianMammalogy 22 121ndash124

MA J G JONES S ZHANG J SHEN W METZNER L ZHANG and B LIANG 2003 Dietary analysisconfirms that Rickettrsquos big-footed bat (Myotisricketti) is a piscivore Journal of Zoology (Lon-don) 261 245ndash248

MACKEY R L and R M R BARCLAY 1989 The influence of physical clutter and noise on the ac-tivity of bats over water Canadian Journal of Zo-ology 67 1167ndash1170

MANLY B F J L L MCDONALD D L THOMAS T L MCDONALD and W P ERICKSON 2002 Re-source selection by animals Statistical designand analysis for field studies Kluwer AcademicPublishers Dordrecht 240 pp

MEacuteDARD P and E GUIBERT 1990 Disparition drsquounmilieu et rareacutefaction drsquoune espPce en France lemurin de Capaccini Myotis capaccinii (Bona-parte 1837) Mammalia 54 297ndash300

NEU C W C R BYERS and J M PEEK 1974 A technique for analysis of utilization-availab-ility data Journal of Wildlife Management 38541ndash545

NORBERG U M and J M V RAYNER 1987 Ecol-ogical morphology and flight in bats (MammaliaChiroptera) wing adaptations flight perfor-mance foraging strategy and echolocation Phil-osophical Transactions of the Royal Society ofLondon 316B 335ndash427

RAYNER J M V 1991 On the aerodynamics of ani-mal flight in ground effect Philosophical Trans-actions of the Royal Society of London 334B119ndash128

ROBSON S K 1984 Myotis adversus (ChiropteraVespertilionidae) Australiarsquos fish-eating batAustralian Mammalogy 7 51ndash52

RUEDI M and A MAYER 2001 Molecular systemat-ics of bats of the genus Myotis (Vespertilionidae)suggests deterministic ecomorphological conver-gences Molecular Phylogenetics and Evolution21 436ndash448

RUSSO D and G JONES 2003 Use of foraging habi-tats by bats in a Mediterranean area determined

by acoustic surveys conservation implicationsEcography 26 197ndash209

RYDELL J L A MILLER and M E JENSEN 1999Echolocation constraints of Daubentonrsquos bat foraging over water Functional Ecology 13247ndash255

SANCHIS-MOLL E J (ed) 1988 Guiacutea de la naturale-za de la Comunidad Valenciana Edicions Alfonsel MagnBnim Valencia 664 pp

SIEMERS B M P STILZ and H-U SCHNITZLER2001 The acoustic advantage of hunting at lowheights above water behavioural experiments onthe European lsquotrawlingrsquo bats Myotis capacciniiM dasycneme and M daubentonii Journal ofExperimental Biology 204 3843ndash3854

SIEMERS B M E BAUR and H-U SCHNITZLER2005 Acoustic mirror effect increases prey de-tection distance in trawling bats Naturwissen-schaften 92 272ndash276

SPITZENBERGER F and O VON HELVERSEN 2001Myotis capaccinii (Bonaparte 1837) mdash Langfuszlig-fledermaus Pp 281ndash302 in Handbuch der Saumlu-getiere Europas Band 4I Fledertiere I (J NIET-HAMMER and F KRAPP eds) Aula-Verlag Wies-baden 602 pp

VAN DE SIJPE M B VANDENDRIESSCHE P VOET J VANDENBERGHE J DUYCK E NAEYAERT M MANHAEVE and E MARTENS 2004 Summerdistribution of the pond bat Myotis dasycne-me (Chiroptera Vespertilionidae) in the west ofFlanders (Belgium) with regard to water qualityMammalia 68 377ndash386

VAUGHAN N G JONES and S HARRIS 1996 Effectsof sewage effluent on the activity of bats (Chi-roptera Vespertilionidae) foraging along riversBiological Conservation 78 337ndash343

VON FRENCKELL B and R M R BARCLAY 1987Bat activity over calm and turbulent water Cana-dian Journal of Zoology 65 219ndash222

WARREN R D D A WATERS J D ALTRINGHAM andD J BULLOCK 2000 The distribution of Dauben-tonrsquos bats (Myotis daubentonii) and pipistrellebats (Pipistrellus pipistrellus) (Vespertilionidae)in relation to small-scale variation in riverinehabitat Biological Conservation 92 85ndash91

WHITE G C and R A GARROT 1990 Analysis ofwildlife radio-tracking data Academic PressLondon 383 pp

WILKINSON G S 1995 Information transfer in batsPp 345ndash360 in Ecology evolution and behav-iour of bats (P A RACEY and S M SWIFT eds)Symposia of the Zoological Society of London67 1ndash421

Received 29 April 2005 accepted 13 January 2006

insectivorous bat a test of the 5 lsquorulersquo of radio-telemetry Journal of Mammalogy 69 379ndash382

ARNOLD A M BRAUN N BECKER and V STORCH1998 Beitrag zur Oumlkologie der Wasserfleder-maus (Myotis daubentoni) in Nordbaden Caroli-nea 56 103ndash110

BARCLAY R M R B J CHRUSZCZ and M RHODES2000 Foraging behaviour of the large-footed my-otis Myotis moluccarum (Chiroptera Vesperti-lionidae) in south-eastern Queensland AustralianJournal of Zoology 48 385ndash392

BOONMAN A M M BOONMAN F BRETSCHNEIDERand W A VAN DE GRIND 1998 Prey detection intrawling insectivorous bats duckweed affectshunting behaviour in Daubentonrsquos bat Myotisdaubentonii Behavioral Ecology and Sociobiol-ogy 44 99ndash107

BRITTON A R C G JONES J M V RAYNER A MBOONMAN and B VERBOOM 1997 Flight per-formance echolocation and foraging behaviourin pond bats Myotis dasycneme (ChiropteraVespertilionidae) Journal of Zoology (London)241 503ndash522

BROOKES A 1988 Channelized rivers perspectivesfor environmental management John Wiley andSons Chichester 342 pp

CARMEL Y and U SAFRIEL 1998 Habitat use bybats in a Mediterranean ecosystem in Israel mdashconservation implications Biological Conserva-tion 84 245ndash250

CHERRY S 1996 A comparison of confidence intervalmethods for habitat use-availability studies Jour-nal of Wildlife Management 60 653ndash658

CIECHANOWSKI M 2002 Community structure andactivity of bats (Chiroptera) over different waterbodies Mammalian Biology 67 276ndash285

COSSON E and P MEacuteDARD 1999 Murin de Capac-cinii Myotis capaccinii (Bonaparte 1837) Pp47ndash51 in Habitats et activiteacute de chasse des chi-roptPres menaceacutes en Europe synthPse de con-naissances actuelles en vue drsquoune gestion conser-vatrice (S Y ROUEacute and M BARATAUD eds) LeRhinolophe Vol Spec 136 pp

COURTOIS J 1998 Contribution B la connaissance dela reacutepartition et des caracteacuteristiques biologiquesdu murin de Capaccini (Myotis capaccinii) enCorse Arvicola 10 42ndash46

FENTON M B and W BOGDANOWICZ 2002 Rela-tionships between external morphology and for-aging behaviour bats in the genus Myotis Cana-dian Journal of Zoology 80 1004ndash1013

FINDLEY J S 1972 Phenetic relationships amongbats of the genus Myotis Systematic Zoology 2131ndash52

FLAVIN D A S S BIGGANE C B SHIEL P SMIDDY

and J S FAIRLEY 2001 Analysis of the diet ofDaubentonrsquos bat Myotis daubentonii in IrelandActa Theriologica 46 43ndash52

GARSHELIS D L 2000 Delusions in habitat evalua-tion measuring use selection and importancePp 111ndash164 in Research techniques in animalecology controversies and consequences (LBOITANI and T K FULLER eds) Columbia Uni-versity Press New York 442 pp

GOITI U J R AIHARTZA I GARIN and J ZABALA2003 Influence of habitat on the foraging behav-iour of the Mediterranean horseshoe bat Rhino-lophus euryale Acta Chiropterologica 5 75ndash84

GUILLEacuteN A 1999 Myotis capaccinii Pp 106ndash107 inThe atlas of European mammals (A J MITCHELL-JONES G AMORI W BOGDANOWICZ B KRY-ŠTUFEK P J H REIJNDERS F SPITZENBERGER M STUBBE J B M THISSEN V VOHRALIK andJ ZIMA eds) T amp AD Poyser Ltd London 484 pp

HUTSON A M S P MICKLEBURGH and P A RACEY(comp) 2001 Microchiropteran bats Global sta-tus survey and conservation action plan IUCNGland 259 pp

JONES G and J M V RAYNER 1988 Flight perfor-mance foraging tactics and echolocation in free-living Daubentonrsquos bats Myotis daubentoni (Chi-roptera Vespertilionidae) Journal of Zoology(London) 215 113ndash132

JONES G and J M V RAYNER 1991 Flight perfor-mance foraging tactics and echolocation in thetrawling insectivorous bat Myotis adversus (Chi-roptera Vespertilionidae) Journal of Zoology(London) 225 393ndash412

JONES G L DUVERGEacute and R D RANSOME 1995Conservation biology of an endangered speciesfield studies of greater horseshoe bat Pp309ndash324 in Ecology evolution and behaviour ofbats (P A RACEY and S M SWIFT eds) Sym-posia of the Zoological Society of London 67 1ndash421

KALKO E 1990 Field study on the echolocation andhunting behaviour of the long-fingered batMyotis capaccinii Bat Research News 3142ndash43

KALKO E K V H-U SCHNITZLER I KAIPF and AD GRINNELL 1998 Echolocation and foragingbehavior of the lesser bulldog bat Noctilio albi-ventris preadaptations for piscivory BehavioralEcology and Sociobiology 42 305ndash319

KUNZ T H and A KURTA 1988 Capture methodsand holding devices Pp 1ndash29 in Ecological andbehavioral methods for the study of bats (T HKUNZ ed) Smithsonian Institution Press Wash-ington DC 533 pp

166 D Almenar J Aihartza U Goiti E Salsamendi and I Garin

Habitat of Myotis capaccinii 167

LANZA B and P AGNELLI 2000 Long fingered batMyotis capaccinii (Bonaparte 1837) Pp 57ndash58in Italian mammals (M SPAGNESI S TOSO andA M DE MARINIS eds) Ministerio dellrsquoAmbien-te Servizio Conservazione della Natura and In-stituto Nazionale per la Fauna Selvatica A GhigiOzzano dellrsquoEmmilia 204 pp

LAW B and C A URQUHART 2000 Diet of the large-footed myotis Myotis macropus at a forest streamroost in northern New South Wales AustralianMammalogy 22 121ndash124

MA J G JONES S ZHANG J SHEN W METZNER L ZHANG and B LIANG 2003 Dietary analysisconfirms that Rickettrsquos big-footed bat (Myotisricketti) is a piscivore Journal of Zoology (Lon-don) 261 245ndash248

MACKEY R L and R M R BARCLAY 1989 The influence of physical clutter and noise on the ac-tivity of bats over water Canadian Journal of Zo-ology 67 1167ndash1170

MANLY B F J L L MCDONALD D L THOMAS T L MCDONALD and W P ERICKSON 2002 Re-source selection by animals Statistical designand analysis for field studies Kluwer AcademicPublishers Dordrecht 240 pp

MEacuteDARD P and E GUIBERT 1990 Disparition drsquounmilieu et rareacutefaction drsquoune espPce en France lemurin de Capaccini Myotis capaccinii (Bona-parte 1837) Mammalia 54 297ndash300

NEU C W C R BYERS and J M PEEK 1974 A technique for analysis of utilization-availab-ility data Journal of Wildlife Management 38541ndash545

NORBERG U M and J M V RAYNER 1987 Ecol-ogical morphology and flight in bats (MammaliaChiroptera) wing adaptations flight perfor-mance foraging strategy and echolocation Phil-osophical Transactions of the Royal Society ofLondon 316B 335ndash427

RAYNER J M V 1991 On the aerodynamics of ani-mal flight in ground effect Philosophical Trans-actions of the Royal Society of London 334B119ndash128

ROBSON S K 1984 Myotis adversus (ChiropteraVespertilionidae) Australiarsquos fish-eating batAustralian Mammalogy 7 51ndash52

RUEDI M and A MAYER 2001 Molecular systemat-ics of bats of the genus Myotis (Vespertilionidae)suggests deterministic ecomorphological conver-gences Molecular Phylogenetics and Evolution21 436ndash448

RUSSO D and G JONES 2003 Use of foraging habi-tats by bats in a Mediterranean area determined

by acoustic surveys conservation implicationsEcography 26 197ndash209

RYDELL J L A MILLER and M E JENSEN 1999Echolocation constraints of Daubentonrsquos bat foraging over water Functional Ecology 13247ndash255

SANCHIS-MOLL E J (ed) 1988 Guiacutea de la naturale-za de la Comunidad Valenciana Edicions Alfonsel MagnBnim Valencia 664 pp

SIEMERS B M P STILZ and H-U SCHNITZLER2001 The acoustic advantage of hunting at lowheights above water behavioural experiments onthe European lsquotrawlingrsquo bats Myotis capacciniiM dasycneme and M daubentonii Journal ofExperimental Biology 204 3843ndash3854

SIEMERS B M E BAUR and H-U SCHNITZLER2005 Acoustic mirror effect increases prey de-tection distance in trawling bats Naturwissen-schaften 92 272ndash276

SPITZENBERGER F and O VON HELVERSEN 2001Myotis capaccinii (Bonaparte 1837) mdash Langfuszlig-fledermaus Pp 281ndash302 in Handbuch der Saumlu-getiere Europas Band 4I Fledertiere I (J NIET-HAMMER and F KRAPP eds) Aula-Verlag Wies-baden 602 pp

VAN DE SIJPE M B VANDENDRIESSCHE P VOET J VANDENBERGHE J DUYCK E NAEYAERT M MANHAEVE and E MARTENS 2004 Summerdistribution of the pond bat Myotis dasycne-me (Chiroptera Vespertilionidae) in the west ofFlanders (Belgium) with regard to water qualityMammalia 68 377ndash386

VAUGHAN N G JONES and S HARRIS 1996 Effectsof sewage effluent on the activity of bats (Chi-roptera Vespertilionidae) foraging along riversBiological Conservation 78 337ndash343

VON FRENCKELL B and R M R BARCLAY 1987Bat activity over calm and turbulent water Cana-dian Journal of Zoology 65 219ndash222

WARREN R D D A WATERS J D ALTRINGHAM andD J BULLOCK 2000 The distribution of Dauben-tonrsquos bats (Myotis daubentonii) and pipistrellebats (Pipistrellus pipistrellus) (Vespertilionidae)in relation to small-scale variation in riverinehabitat Biological Conservation 92 85ndash91

WHITE G C and R A GARROT 1990 Analysis ofwildlife radio-tracking data Academic PressLondon 383 pp

WILKINSON G S 1995 Information transfer in batsPp 345ndash360 in Ecology evolution and behav-iour of bats (P A RACEY and S M SWIFT eds)Symposia of the Zoological Society of London67 1ndash421

Received 29 April 2005 accepted 13 January 2006

Habitat of Myotis capaccinii 167

LANZA B and P AGNELLI 2000 Long fingered batMyotis capaccinii (Bonaparte 1837) Pp 57ndash58in Italian mammals (M SPAGNESI S TOSO andA M DE MARINIS eds) Ministerio dellrsquoAmbien-te Servizio Conservazione della Natura and In-stituto Nazionale per la Fauna Selvatica A GhigiOzzano dellrsquoEmmilia 204 pp

LAW B and C A URQUHART 2000 Diet of the large-footed myotis Myotis macropus at a forest streamroost in northern New South Wales AustralianMammalogy 22 121ndash124

MA J G JONES S ZHANG J SHEN W METZNER L ZHANG and B LIANG 2003 Dietary analysisconfirms that Rickettrsquos big-footed bat (Myotisricketti) is a piscivore Journal of Zoology (Lon-don) 261 245ndash248

MACKEY R L and R M R BARCLAY 1989 The influence of physical clutter and noise on the ac-tivity of bats over water Canadian Journal of Zo-ology 67 1167ndash1170

MANLY B F J L L MCDONALD D L THOMAS T L MCDONALD and W P ERICKSON 2002 Re-source selection by animals Statistical designand analysis for field studies Kluwer AcademicPublishers Dordrecht 240 pp

MEacuteDARD P and E GUIBERT 1990 Disparition drsquounmilieu et rareacutefaction drsquoune espPce en France lemurin de Capaccini Myotis capaccinii (Bona-parte 1837) Mammalia 54 297ndash300

NEU C W C R BYERS and J M PEEK 1974 A technique for analysis of utilization-availab-ility data Journal of Wildlife Management 38541ndash545

NORBERG U M and J M V RAYNER 1987 Ecol-ogical morphology and flight in bats (MammaliaChiroptera) wing adaptations flight perfor-mance foraging strategy and echolocation Phil-osophical Transactions of the Royal Society ofLondon 316B 335ndash427

RAYNER J M V 1991 On the aerodynamics of ani-mal flight in ground effect Philosophical Trans-actions of the Royal Society of London 334B119ndash128

ROBSON S K 1984 Myotis adversus (ChiropteraVespertilionidae) Australiarsquos fish-eating batAustralian Mammalogy 7 51ndash52

RUEDI M and A MAYER 2001 Molecular systemat-ics of bats of the genus Myotis (Vespertilionidae)suggests deterministic ecomorphological conver-gences Molecular Phylogenetics and Evolution21 436ndash448

RUSSO D and G JONES 2003 Use of foraging habi-tats by bats in a Mediterranean area determined

by acoustic surveys conservation implicationsEcography 26 197ndash209

RYDELL J L A MILLER and M E JENSEN 1999Echolocation constraints of Daubentonrsquos bat foraging over water Functional Ecology 13247ndash255

SANCHIS-MOLL E J (ed) 1988 Guiacutea de la naturale-za de la Comunidad Valenciana Edicions Alfonsel MagnBnim Valencia 664 pp

SIEMERS B M P STILZ and H-U SCHNITZLER2001 The acoustic advantage of hunting at lowheights above water behavioural experiments onthe European lsquotrawlingrsquo bats Myotis capacciniiM dasycneme and M daubentonii Journal ofExperimental Biology 204 3843ndash3854

SIEMERS B M E BAUR and H-U SCHNITZLER2005 Acoustic mirror effect increases prey de-tection distance in trawling bats Naturwissen-schaften 92 272ndash276

SPITZENBERGER F and O VON HELVERSEN 2001Myotis capaccinii (Bonaparte 1837) mdash Langfuszlig-fledermaus Pp 281ndash302 in Handbuch der Saumlu-getiere Europas Band 4I Fledertiere I (J NIET-HAMMER and F KRAPP eds) Aula-Verlag Wies-baden 602 pp

VAN DE SIJPE M B VANDENDRIESSCHE P VOET J VANDENBERGHE J DUYCK E NAEYAERT M MANHAEVE and E MARTENS 2004 Summerdistribution of the pond bat Myotis dasycne-me (Chiroptera Vespertilionidae) in the west ofFlanders (Belgium) with regard to water qualityMammalia 68 377ndash386

VAUGHAN N G JONES and S HARRIS 1996 Effectsof sewage effluent on the activity of bats (Chi-roptera Vespertilionidae) foraging along riversBiological Conservation 78 337ndash343

VON FRENCKELL B and R M R BARCLAY 1987Bat activity over calm and turbulent water Cana-dian Journal of Zoology 65 219ndash222

WARREN R D D A WATERS J D ALTRINGHAM andD J BULLOCK 2000 The distribution of Dauben-tonrsquos bats (Myotis daubentonii) and pipistrellebats (Pipistrellus pipistrellus) (Vespertilionidae)in relation to small-scale variation in riverinehabitat Biological Conservation 92 85ndash91

WHITE G C and R A GARROT 1990 Analysis ofwildlife radio-tracking data Academic PressLondon 383 pp

WILKINSON G S 1995 Information transfer in batsPp 345ndash360 in Ecology evolution and behav-iour of bats (P A RACEY and S M SWIFT eds)Symposia of the Zoological Society of London67 1ndash421

Received 29 April 2005 accepted 13 January 2006