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ARTICLE IN PRESSG ModelAMBIO-40595; No. of Pages 5Mammalian Biology xxx (2013) xxxxxx

Contents lists available at SciVerse ScienceDirect

Mammalian Biology

jou rna l h omepa ge: www.elsev ier .com/ locate /mambio

hort Communication

uriosity killed the bat: Domestic cats as bat predators

eonardo Ancillottoa, Maria Tiziana Serangelib, Danilo Russob,c,

Dipartimento di Biologia e Biotecnologie Charles Darwin, Universit degli Studi di Roma La Sapienza, Roma, ItalyLaboratorio di Ecologia Applicata, Dipartimento di Agraria, Universit degli Studi di Napoli Federico II, Portici (Napoli), ItalySchool of Biological Sciences, University of Bristol, Bristol, UK

r t i c l e i n f o

rticle history:eceived 14 November 2012ccepted 22 January 2013vailable online xxx

eywords:hiropteraelidaeredationrban ecosystemsildlife rehabilitation

a b s t r a c t

Domestic cats are suspected to have an impact on wild populations of birds and small mammals, butpublished reports of predation on bats are either rare or anecdotal. We based our study on 1012 recordsof bats admitted at four wildlife rescue centres in peninsular Italy in 20092011. We hypothesized that(1) cats prevalently prey on bats emerging from roosts, so newborns or non volant juveniles should beless exposed to predation; (2) because cats occur in human settlements, the bat species most frequentlyinvolved are house-roosting (3) predation is season-biased, most events being more likely to take placein summer when females congregate in roosts to reproduce; (4) predation events concentrate in sparse-urban and rural areas, where free-ranging cats occur more frequently; and (5) some individual cats mayspecialize in capturing bats. We found that predation by cats was the first cause of rescue for bats inthe study area, accounting for 28.7% of records of adult bats admitted to rehabilitation centres. Althoughmost bats caught by cats belonged to house-roosting species, at least 3 of the 11 species affected weretree- or cave-roosting. Predation affected more frequently adult females in summer and thus threatenedreproductive colonies, which were often subjected to repeated predations. As predicted, predation eventswere associated with land cover, being more abundant in rural and sparse urban areas, where cats are

more often allowed to stay outdoor, as confirmed by the results of a cat owner survey we carried out. Catsare explorative mammals, so they may be easily attracted at bat roosts by sensory cues involving sound,smell and vision. Our analysis covered a broad geographical area over a relatively long period and suggeststhat the threat posed to bats by cats may be significant and should be carefully considered in conservationplans. Strategies to mitigate this impact should encompass the control of feral cat populations and indoor

at leasellscrestriction of owned cats 2013 Deutsche Ge

In human-dominated landscapes many wild animal species fallictim to opportunistic wild (Thorington and Bowman, 2003) oromestic predators including dogs (Beck, 1973; Galetti and Sazima,006) and cats (Churcher and Lawton, 1989). In urban areas, domes-ic cats are the most abundant carnivores (Coleman and Temple,993; Lepczyk et al., 2003) and thus can prey on a large amountf wildlife every year (Woods et al., 2003). So far, most studiesave addressed the impact of cats on particular taxa (birds; e.g.an Heezik et al., 2010), in specific geographical contexts (islands,here introduced cats can become invasive, thus being a pecu-

iar and different case; Dickman, 1996; Medina and Nogales, 2009)r have regarded relatively limited periods (Woods et al., 2003;Please cite this article in press as: Ancillotto, L., et al., Curiosity killedhttp://dx.doi.org/10.1016/j.mambio.2013.01.003

epczyk et al., 2003). Other biases that affect the current knowl-dge of wildlife predation by cats originate by the heterogeneousonditions of free-ranging felines (owned or feral) considered for

Corresponding author at: Laboratorio di Ecologia Applicata, Dipartimento digraria, Universit degli Studi di Napoli Federico II, via Universit 100, I-80055ortici (Napoli), Italy. Tel.: +39 2532017.

E-mail address: danrusso@unina.it (D. Russo).

616-5047/$ see front matter 2013 Deutsche Gesellschaft fr Sugetierkunde. Publisttp://dx.doi.org/10.1016/j.mambio.2013.01.003st where predation on bats is probable.haft fr Sugetierkunde. Published by Elsevier GmbH. All rights reserved.

analysis and the different methods adopted (scat/guts analysis orprey brought home), overall making comparisons of different stud-ies difficult (Krauze-Gryz et al., 2012).

Many bat species from temperate regions roost in human-madestructures, often houses (Barbour and Davis, 1969), for at least apart of their life cycle (generally the reproductive phase, whenfemales congregate in nurseries), a habit which increases the like-lihood of encountering cats. Because nursery colonies are oftencomposed of many individuals, predators may take a large toll onthem (Rodrguez-Durn and Lewis, 1985; Speakman, 1991; Rosinaand Shokhrin, 2011; Scrimgeour et al., 2012), particularly on adultsand volant juveniles, newborns and non-volant bats being lessexposed to the risk of predation.

Although only occasional evidence of cat predation on bats isavailable (Phillips et al., 2001; Woods et al., 2003; Mastrobuoniet al., 2005), there are reasons to believe that the impact of cats onbats is most likely to have been underestimated (Altringham, 2011). the bat: Domestic cats as bat predators. Mammal. Biol. (2013),

To help fill this knowledge gap, in this study we present a 3-yearassessment of cat predation on bats based on the analysis of recordsof rescued bats available from four Italian wildlife rehabilitationcentres.

hed by Elsevier GmbH. All rights reserved.

dx.doi.org/10.1016/j.mambio.2013.01.003dx.doi.org/10.1016/j.mambio.2013.01.003http://www.sciencedirect.com/science/journal/16165047http://www.elsevier.com/locate/mambiomailto:danrusso@unina.itdx.doi.org/10.1016/j.mambio.2013.01.003

ARTICLE ING ModelMAMBIO-40595; No. of Pages 52 L. Ancillotto et al. / Mammalian B

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ig. 1. Map of Italian regions indicating the place of origin of rescued bats. Dark-rey: >100 records; grey: 10010 records; light-grey: 15 records; white: noecords. Star-crosses indicate the position of the four wildlife rescue centres.

We hypothesize that (1) most predations concern adult batsmerging from roosts and volant but inexperienced juveniles,hich are more easily available to cats; (2) because domesticats are associated with human settlements, the bat species mostrequently involved are house-roosting; (3) predation is season-iased, most events being more likely to take place in summer whenemales congregate in roosts to reproduce so that they and theiroung are more conspicuous and accessible to cats; (4) cats living inreas characterized by varying land use may be subject to differentanagement and differ in the amount of time spent outdoor, thusat impact on bats may be influenced by land use type, being partic-larly relevant in rural and sparse-urban areas; (5) individual catsay specialize on bats and repeatedly visit the same colony, lead-

ng to locally significant impacts. To support hypothesis 4 we alsoonducted a survey among cat owners to explore the occurrence ofifferences in the way domestic cats are managed in a range of landse types. Because gardens and courtyards are commoner in ruralr sparse urban areas, we predicted that in such areas domestic catsre more often left outdoor unguarded and thus represent a moreignificant threat to bats.

We analyzed all records of rescued bats admitted to four LIPUItalian League for the Protection of Birds) wildlife rehabilita-ion centres in 20092011. Centres were located in central (Laziond Toscana) and northern (Veneto and Lombardia) Italy (Fig. 1).ecords included date, species, sex (determined for 72.1% and 22.9%Please cite this article in press as: Ancillotto, L., et al., Curiosity killedhttp://dx.doi.org/10.1016/j.mambio.2013.01.003

f adults and juveniles, respectively), age class (i.e. adult/juvenile,he former showing cartilage epiphyseal plates in finger bones andore tapered finger joints; see Anthony, 1988), reproductive sta-

us, injuries (when present) and causes of rescue. The latter were PRESSiology xxx (2013) xxxxxx

categorized as follows: (1) impact, i.e. bats which collided withbuildings, vehicles and other human-made structures; (2) debili-tation, i.e. animals found starving or dehydrated; (3) cat predation;(4) removal from roost, i.e. bats found in a roost and purposelytaken from it by people; (5) unknown reason, and (6) fallen fromroost (newborns or non-volant juveniles found on the floor beneatha roost). The people bringing bats were also briefly interviewed bythe centres staff to record the cause of rescue. In all selected cen-tres, most admitted bats were reliably identified by trained staff;bats whose identity was uncertain were categorized as undeter-mined. The cryptic Pipistrellus pipistrellus and P. pygmaeus, whosedistinction may be confidently done only by acoustic or molecularanalysis (Jones and Parjis, 1993), were pooled together.

We assessed landscape composition within a 1-km radius circlesurrounding each rescue site by photo interpretation of orthopho-tos (Italian Ministry for the Environment, Land and Sea) and usingthe gvSIG open-source GIS software (Iver, Generalitat Valencia,Universidad Jaume I and Prodevelop, Spain). The chosen radiuscovers the average home range size generally shown by free-ranging domestic cats (Kays and DeWan, 2004). Land cover wasclassified as: dense urban areas (continuous urban matrix, withmulti-storey buildings and vegetation cover 95%).

To explore differences in cat management by owners in differ-ent land use types, we submitted a questionnaire to cat ownersin veterinary clinics, pet shops as well as door-to-door in differenturban conditions in three regions of Central Italy (Lazio, Umbria andAbruzzo), covering the previously described categories. Cat ownerswere asked to communicate (a) whether their cat was allowed toget access to outdoor spaces, (b) the amount of time spent outside indaytime (c) whether cats had access to outdoor during night time,and (d) how many bats their cat brought home in the last threeyears.

We determined the association between rescue causes of batsand respectively month, season, gender and land cover by chi-square tests on contingency tables. The same test was applied to thecats owner surveys data: in that case we tested whether the num-bers of bats brought home by cats were associated respectively withthe time spent outdoor in daytime and with outdoor access duringthe night, as well as with land cover. Significant (p < 0.05) chi-squaretests were followed by an analysis of residuals to determine thecontribution of each category to the result (Haberman, 1973). Alltests were performed with R rel. 2.14.0 (http://www.R-project.org).

We obtained records of bats admitted to four wildlife rescuecentres originating from 13 Italian regions, but mainly from Lazio,Toscana (central Italy) and Veneto (northern Italy; Fig. 1). Of 1012bats admitted to the centres, 115 (11.3%) had been preyed upon bya cat. Rescued bats belonged to 12 species, for all of which exceptMyotis bechsteinii cases of predation by cats had been recorded(Table 1).

In agreement with our hypothesis, only 2.4% of newborns orjuveniles admitted had been caught by cats whereas adults weremuch more frequent: of 341 adult records, cat predation wasthe most frequent rescue cause (28.7%), followed by debilitation(23.3%), impact (18.2%), and removal from roost (4.6%); unknowncauses accounted for 25.2% of cases. Of 671 records of newbornsand juveniles, 90.0% were young bats accidentally fallen from roost,other causes accounting for 10% of the dataset. Because the method the bat: Domestic cats as bat predators. Mammal. Biol. (2013),

we adopted for ageing bats does not permit to distinguish betweenflying and non-flying juveniles, as epiphyseal plates can be seenon young bats for a few weeks after they become volant (Brunet-Rossini and Wilkinson, 2009), at least a few records from our

dx.doi.org/10.1016/j.mambio.2013.01.003http://www.r-project.org/

ARTICLE ING ModelMAMBIO-40595; No. of Pages 5L. Ancillotto et al. / Mammalian B

Table 1Numbers and species of adult and juvenile bats admitted at four wildlife rescuecentres in peninsular Italy between 2009 and 2011. Numbers in parentheses indicatebats caught by cats.

Species Adults Juveniles

Kuhls pipistrelle (Pipistrellus kuhlii) 150 (38) 331 (4)Savis pipistrelle (Hypsugo savii) 120 (35) 171 (7)European free-tailed bat (Tadarida teniotis) 14 (2) 22 (0)Common/soprano pipistrelle (Pipistrelluspipistrellus and P. pygmaeus)

5 (5) 37 (2)

Common serotine (Eptesicus serotinus) 4 (1) 1 (0)Leislers bat (Nyctalus leisleri) 3 (2) 2 (1)Brown long-eared bat (Plecotus auritus) 2 (1) 2 (0)Schreibers bat (Miniopterus schreibersii) 2 (2) 0Nathusius pipistrelle (Pipistrellus nathusii) 1 (1) 0Bechsteins bat (Myotis bechsteinii) 0 1 (0)Natterers bat (Myotis nattereri) 1 (1) 0Lesser horseshoe bat (Rhinolophus hipposideros) 1 (1) 0Unidentified 38 (10) 104 (2)

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the colony disappeared after undergoing two years of numerous

Fb

TOT 341 (99) 671 (16)

ategory juvenile may have been volant young bats: thus, the dif-erence we detected between adults and juveniles preyed by catsay be even stronger. Such young bats were admitted exclusivelyuring summer months, i.e. in the reproductive season (from Juneo September, see Fig. 2af), so that the following results refer tohe adult-sample only.

As predicted, house-roosting bats, including Savis pipistrellesHypsugo savii), European free-tailed bats (Tadarida teniotis) anduhls and common/soprano pipistrelles (Pipistrellus kuhlii and P.ipistrellus/pygmaeus) approached 80% of the total sample of pre-ation on adults, but this also featured, albeit occasionally, speciesore often roosting in caves (Miniopterus schreibersii) or trees (Pip-

strellus nathusii, Nyctalus leisleri).The relative frequencies of the rescue causes recorded dif-

ered across months (2 = 39.667, df = 12, p = 0.012) and seasonsPlease cite this article in press as: Ancillotto, L., et al., Curiosity killedhttp://dx.doi.org/10.1016/j.mambio.2013.01.003

2 = 15.082, df = 3, p = 0.013), i.e. spring (AprilJune), sum-er (JulySeptember), autumn (OctoberDecember) and winter

JanuaryMarch). Although predation by cats was recorded

ig. 2. Monthly relative frequencies of bat rescue causes from wildlife rescue centres in = impact; c = debilitation; d = removal from roost; e = unknown reason; f = fallen from ro PRESSiology xxx (2013) xxxxxx 3

year-round, it varied between months (2 = 53.197, df = 11,p < 0.0001) and ranged between 5.3% (January) and 43.5% (May),being highest from May to September. The other causes of rescuepeaked in mid-summer months, except deliberate removal fromroost which mainly occurred in winter and probably concernedtorpid subjects which people easily took from roosts (Fig. 2d).

Female adult bats were preyed upon by cats more frequentlythan males, (2 = 5.444, df = 1, p = 0.019) accounting for 68.7% ofthe sample, whereas no influence of gender was detected onthe remaining causes (debilitation: 2 = 2.131, df = 1, n.s.; impact:2 = 1.316, df = 1, n.s.; removal: 2 = 1, df = 1, n.s.; unknown: 2 = 1.4,df = 1, n.s.).

The frequency of occurrence of predation events on adult batswas associated with land cover (2 = 39.096, df = 16, p = 0.001),being higher in rural areas as well as in areas characterized byscattered buildings, whereas impact events and unknown causes ofrescue were more frequent in dense urban areas; all other causesshowed no land cover association.

Land cover also influenced the distribution of rescue events fornewborns or non-volant juveniles (2 = 38.315, df = 16, p = 0.001):debilitation, impact and unknown cause records were more fre-quent in dense urban areas, whereas predation by cats andjuveniles fallen from roost were more frequent in sparse and ruralareas.

According to what reported by people who rescued the bats andbrought them to the rehabilitation centres, at least ten roosts (H.savii, n = 4; P. pipistrellus/pygmaeus, n = 2; P. kuhlii, = n = 2; P. kuhliiand H. savii mixed colonies, n = 2) were targeted by repeated catpredation over 13 years. All cats involved had an owner and wereallowed to access terraces or gardens; they caught, on average,7.3 4.3 bats per year (mean SD; n = 10). The highest predationrate affected a P. kuhlii nursery (where in three years at least 47bats were caught during roost emergence) which was accessibleto the cat at night; in another case (an H. savii nursery colony) the bat: Domestic cats as bat predators. Mammal. Biol. (2013),

predation events (Table 2).Our cat owner survey (n = 106) showed that the time domes-

tic cats were allowed to spend outside decreased along a

the 2009/2011 period. Black bars: adults; grey bars: juveniles; a = cat predation;ost.

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ARTICLE ING ModelMAMBIO-40595; No. of Pages 54 L. Ancillotto et al. / Mammalian B

Table 2Numbers of adult bats preyed upon by single owned cats at different building roostsin consecutive years between 2009 and 2011.

Colony location Species 2009 2010 2011

Rome Hypsugo savii 12 8 Castelnuovo di Porto Pipistrellus pipistrellus/pygmaeus 7 6 4Cassino Pipistrellus kuhlii 15 20 12Rome Pipistrellus pipistrellus/pygmaeus 4 4Rome Pipistrellus kuhlii & Hypsugo savii 5 8Rome Hypsugo savii 5 6Rome Hypsugo savii 7 6Rome Hypsugo savii 5 4Villa Latina Pipistrellus kuhlii 2Rome Pipistrellus kuhlii & Hypsugo savii 7

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ural-to-urban gradient. Cats in rural and sparse urban areas werellowed to stay outdoor for longer than cats in denser urban areas2 = 52.735, df = 12, p < 0.001) with 34.4% of cats from rural andrban-sparse areas getting outdoor access for 612 h during day-ime versus only 7% of cats from dense urban settlements. Morehan 41% of cats from rural and sparse-urban areas had regularutdoor access at night-time, while this was possible only for the% of cats from dense urban areas (2 = 37.647, df = 9, p < 0.001).redation on bats was more common in rural and sparse urbanreas (2 = 8.923, df = 3, p = 0.030), involving 16.4% and 7% of catsrom rural/urban-sparse areas and urban-dense respectively; itas more frequently performed by cats regularly staying outsidevernight (2 = 9.963, df = 3, p = 0.020), which were reported to preyn average 2.0 1.3 bats/year (n = 13).Our study shows that in the Italian peninsula cat predation

ay at least locally represent a significant threat to bats, beinghe first cause of admission of adult bats to rehabilitation centres.learly, the actual impact of predation by cats on bat populationss still unknown and our study was not designed to answer thisuestion. Species roosting in buildings were more frequent in ourample, yet this was expected since our analysis concerned subjectsescued by people, an event definitely more likely to occur nearouses.As hypothesized, juveniles were less frequently caught than

dults despite being smaller and easier to subdue and handle for aat. In most cases the bat species we found to be affected by pre-ation roost in crevices or fissures, where non-volant roosting batsre effectively sheltered from cats unless they fall accidentally fromhe roost (Serangeli et al., 2012). However, because newborns maye more easily dismembered or eaten by a cat, or more frequentlyverlooked by rescuers, this may have at least partly influencedur results leading to an underestimation of such small prey itemsKrauze-Gryz et al., 2012).

We also found that predation on bats mainly occurred in ruralreas, or areas characterized by single buildings interspersed witharge vegetation patches. Our cat owner survey confirmed that catsre more frequently allowed to stay outside in rural or sparse urbanreas, where bat roosts are also common. Instead, owned cats inenser urban settlements were kept indoor more often, a fact whichay account for the lower numbers of bats caught in such landse situations, despite the possibly higher absolute density of catsNatoli, 1985).

The type of analysis we did inevitably underestimated thempact on bats using roosts other than buildings, which may beot negligible especially if feral cats are considered. Despite ourrobable underestimation, 3 of the 11 species found to be affectedPlease cite this article in press as: Ancillotto, L., et al., Curiosity killedhttp://dx.doi.org/10.1016/j.mambio.2013.01.003

y cat predation typically roost in caves or trees (e.g. Schobernd Grimmberger, 1997). Altogether, these data are alarmingecause they suggest that free-ranging cats may have a signif-cant impact on bats (including those from species of special PRESSiology xxx (2013) xxxxxx

conservation concern) both in human-modified and natural land-scapes where their rescue, or the retrieval of their remains, is muchless probable and may go unnoticed.

Predation was more likely to concern adult females and occurredin summer (i.e. in the reproductive season) when nurseries con-gregate. Cats are explorative mammals (Machado and Genaro,2010), so they may be easily attracted by sensory cues includ-ing sound emitted by bats, the smell of droppings accumulatednear roost sites or the observation of flying bats at its entrance.Bats seem to be particularly vulnerable to predation by cats whenleaving roosts (Irwin and Speakman, 2003; Scrimgeour et al.,2012); the broad frequency range heard by cats (between 48 and85,000 Hz) enable them to detect most bat echolocation and socialcalls (Heffner and Heffner, 1985). In fact sound, and smell too,are conspicuous cues helping predators to locate roosts (Gilletteand Kimbrough, 1970; Fenton, 1995; Irwin and Speakman, 2003)and once a roost is found, repeated predation may cause signifi-cant mortality (Scrimgeour et al., 2012, this work) not effectivelycompensated by new births due to bats slow reproduction rate(Racey, 1982). Moreover, pregnant females may also be easier tar-gets to predators due to their limited manoeuvrability in flight(Russo et al., 2007). Repeated predation is commonly performedby individual cats which specialize both on bats (Scrimgeouret al., 2012) and other prey (Churcher and Lawton, 1989; Tschanzet al., 2011). Foraging bats too may be caught by cats when fly-ing close to the ground (see DelPietro et al., 1994 for Desmodusrotundus in central America), or gleaning prey from substrate ahunting technique used by several European species (Dietz et al.,2009).

Although small mammals are common cat prey (Liberg, 1984;Woods et al., 2003), bat predation is only accidentally recorded(Dickman, 1996; Phillips et al., 2001; Mastrobuoni et al., 2005).It is important to highlight that bats are also absent (Kays andDeWan, 2004; Medina and Nogales, 2009; Krauze-Gryz et al., 2012)or extremely rare (Woods et al., 2003; Krauze-Gryz et al., 2012) inthe prey datasets of studies from different regions. This may bedue to the small size of most bats which make them easy to ingest,so their remains are rarely found (as demonstrated by Chapmanand Kitchener in 1978, who rarely found bat remains among catprey items, but often detected them in stomach samples). How-ever, bats were also rarely found in scat-gut contents analyzed byKrauze-Gryz et al. (2012).

We are aware that our sample was biased in terms of habitatsand species considered because it only featured bats brought torescue centres. Nonetheless, data from such sources are still valu-able in assessing the threats to urban wildlife and their ecologicalcorrelates (Koenig et al., 2002; Shine and Koenig, 2001). Overall,our assessment of the phenomenon, covering a broad geographicalarea over a relatively long period, suggests that the threat posed bydomestic cats may be significant especially on a local scale: recidi-vist cats systematically targeting the same site may prompt a colonyto move to a new roost, thus exposing the bats to higher preda-tion risks or reproductive failure, or even eradicate it. Certainly,this threat should be considered in conservation plans. Our view issupported by Altringham (2011) who inferred from data by Woodset al. (2003) that every year approximately 250,000 bats are killedby cats in the UK.

Practices such as the demographic control of feral cats by steril-ization and night-time indoor restriction of domestic cats, at leastduring the reproductive season of bats, may certainly reduce theirimpact. At least for birds and mice, prey deterrents such as collar-mounted bells have proven useful to warn potential victims of the the bat: Domestic cats as bat predators. Mammal. Biol. (2013),

presence of a cat (Ruxton et al., 2002; Woods et al., 2003): whetherthey might also be useful to reduce predation on bats e.g. at roosts(by delaying bat emergence when the predator is nearby) has yetto be tested.

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ARTICLEAMBIO-40595; No. of Pages 5L. Ancillotto et al. / Mamma

cknowledgements

We are grateful to the Italian League for the Protection of BirdsLIPU) and particularly to the staff and volunteers of the wildlifeescue centres that readily provided us with valuable data andupport: Centro Recupero Fauna Selvatica (Roma), Centro Recu-ero Uccelli Marini (Livorno), Centro Recupero Fauna Selvatica Laagiana (Pontevecchio di Magenta Milano) and Centro Recuperonimali Selvatici (Padova). We also thank Fabio Bontadina, Xavieruig and one anonymous reviewer for their valuable comments on

previous ms version.

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Curiosity killed the bat: Domestic cats as bat predatorsAcknowledgementsReferences