G Model

MAMBIO-40595; No. of Pages 5

ARTICLE IN PRESS
Mammalian Biology xxx (2013) xxxxxx

Contents lists available at SciVerse ScienceDirect

Mammalian Biology
journal homepage: www.elsevier.com/locate/mambio

Short Communication

Curiosity killed the bat: Domestic cats as bat predators

Leonardo Ancillotto a , Maria Tiziana Serangeli b , Danilo Russo b,c,
a

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

a r t i c l e

i n f o

Article history:
Received 14 November 2012
Accepted 22 January 2013
Available online xxx
Keywords:
Chiroptera
Felidae
Predation
Urban ecosystems
Wildlife 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, but
published reports of predation on bats are either rare or anecdotal. We based our study on 1012 records
of 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 be
less exposed to predation; (2) because cats occur in human settlements, the bat species most frequently
involved are house-roosting (3) predation is season-biased, most events being more likely to take place
in summer when females congregate in roosts to reproduce; (4) predation events concentrate in sparseurban and rural areas, where free-ranging cats occur more frequently; and (5) some individual cats may
specialize in capturing bats. We found that predation by cats was the rst cause of rescue for bats in
the study area, accounting for 28.7% of records of adult bats admitted to rehabilitation centres. Although
most bats caught by cats belonged to house-roosting species, at least 3 of the 11 species affected were
tree- or cave-roosting. Predation affected more frequently adult females in summer and thus threatened
reproductive colonies, which were often subjected to repeated predations. As predicted, predation events
were associated with land cover, being more abundant in rural and sparse urban areas, where cats are
more often allowed to stay outdoor, as conrmed by the results of a cat owner survey we carried out. Cats
are 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 suggests
that the threat posed to bats by cats may be signicant and should be carefully considered in conservation
plans. Strategies to mitigate this impact should encompass the control of feral cat populations and indoor
restriction of owned cats at least where predation on bats is probable.
2013 Deutsche Gesellschaft fr Sugetierkunde. Published by Elsevier GmbH. All rights reserved.

In human-dominated landscapes many wild animal species fall

victim to opportunistic wild (Thorington and Bowman, 2003) or
domestic predators including dogs (Beck, 1973; Galetti and Sazima,
2006) and cats (Churcher and Lawton, 1989). In urban areas, domestic cats are the most abundant carnivores (Coleman and Temple,
1993; Lepczyk et al., 2003) and thus can prey on a large amount
of wildlife every year (Woods et al., 2003). So far, most studies
have addressed the impact of cats on particular taxa (birds; e.g.
Van Heezik et al., 2010), in specic geographical contexts (islands,
where introduced cats can become invasive, thus being a peculiar and different case; Dickman, 1996; Medina and Nogales, 2009)
or have regarded relatively limited periods (Woods et al., 2003;
Lepczyk et al., 2003). Other biases that affect the current knowledge of wildlife predation by cats originate by the heterogeneous
conditions of free-ranging felines (owned or feral) considered for

Corresponding author at: Laboratorio di Ecologia Applicata, Dipartimento di

Agraria, Universit degli Studi di Napoli Federico II, via Universit 100, I-80055
Portici (Napoli), Italy. Tel.: +39 2532017.
E-mail address: danrusso@unina.it (D. Russo).

analysis and the different methods adopted (scat/guts analysis or

prey brought home), overall making comparisons of different studies difcult (Krauze-Gryz et al., 2012).
Many bat species from temperate regions roost in human-made
structures, often houses (Barbour and Davis, 1969), for at least a
part of their life cycle (generally the reproductive phase, when
females congregate in nurseries), a habit which increases the likelihood of encountering cats. Because nursery colonies are often
composed of many individuals, predators may take a large toll on
them (Rodrguez-Durn and Lewis, 1985; Speakman, 1991; Rosina
and Shokhrin, 2011; Scrimgeour et al., 2012), particularly on adults
and volant juveniles, newborns and non-volant bats being less
exposed to the risk of predation.
Although only occasional evidence of cat predation on bats is
available (Phillips et al., 2001; Woods et al., 2003; Mastrobuoni
et al., 2005), there are reasons to believe that the impact of cats on
bats is most likely to have been underestimated (Altringham, 2011).
To help ll this knowledge gap, in this study we present a 3-year
assessment of cat predation on bats based on the analysis of records
of rescued bats available from four Italian wildlife rehabilitation
centres.

1616-5047/$ see front matter 2013 Deutsche Gesellschaft fr Sugetierkunde. Published by Elsevier GmbH. All rights reserved.
http://dx.doi.org/10.1016/j.mambio.2013.01.003

Please cite this article in press as: Ancillotto, L., et al., Curiosity killed the bat: Domestic cats as bat predators. Mammal. Biol. (2013),
http://dx.doi.org/10.1016/j.mambio.2013.01.003

G Model
MAMBIO-40595; No. of Pages 5
2

ARTICLE IN PRESS
L. Ancillotto et al. / Mammalian Biology xxx (2013) xxxxxx

Fig. 1. Map of Italian regions indicating the place of origin of rescued bats. Darkgrey: >100 records; grey: 10010 records; light-grey: 15 records; white: no
records. Star-crosses indicate the position of the four wildlife rescue centres.

We hypothesize that (1) most predations concern adult bats

emerging from roosts and volant but inexperienced juveniles,
which are more easily available to cats; (2) because domestic
cats are associated with human settlements, the bat species most
frequently involved are house-roosting; (3) predation is seasonbiased, most events being more likely to take place in summer when
females congregate in roosts to reproduce so that they and their
young are more conspicuous and accessible to cats; (4) cats living in
areas characterized by varying land use may be subject to different
management and differ in the amount of time spent outdoor, thus
cat impact on bats may be inuenced by land use type, being particularly relevant in rural and sparse-urban areas; (5) individual cats
may specialize on bats and repeatedly visit the same colony, leading to locally signicant impacts. To support hypothesis 4 we also
conducted a survey among cat owners to explore the occurrence of
differences in the way domestic cats are managed in a range of land
use types. Because gardens and courtyards are commoner in rural
or sparse urban areas, we predicted that in such areas domestic cats
are more often left outdoor unguarded and thus represent a more
signicant threat to bats.
We analyzed all records of rescued bats admitted to four LIPU
(Italian League for the Protection of Birds) wildlife rehabilitation centres in 20092011. Centres were located in central (Lazio
and Toscana) and northern (Veneto and Lombardia) Italy (Fig. 1).
Records included date, species, sex (determined for 72.1% and 22.9%
of adults and juveniles, respectively), age class (i.e. adult/juvenile,
the former showing cartilage epiphyseal plates in nger bones and
more tapered nger joints; see Anthony, 1988), reproductive status, injuries (when present) and causes of rescue. The latter were

categorized as follows: (1) impact, i.e. bats which collided with

buildings, vehicles and other human-made structures; (2) debilitation, i.e. animals found starving or dehydrated; (3) cat predation;
(4) removal from roost, i.e. bats found in a roost and purposely
taken from it by people; (5) unknown reason, and (6) fallen from
roost (newborns or non-volant juveniles found on the oor beneath
a roost). The people bringing bats were also briey interviewed by
the centres staff to record the cause of rescue. In all selected centres, most admitted bats were reliably identied by trained staff;
bats whose identity was uncertain were categorized as undetermined. The cryptic Pipistrellus pipistrellus and P. pygmaeus, whose
distinction may be condently done only by acoustic or molecular
analysis (Jones and Parjis, 1993), were pooled together.
We assessed landscape composition within a 1-km radius circle
surrounding each rescue site by photo interpretation of orthophotos (Italian Ministry for the Environment, Land and Sea) and using
the gvSIG open-source GIS software (Iver, Generalitat Valencia,
Universidad Jaume I and Prodevelop, Spain). The chosen radius
covers the average home range size generally shown by freeranging domestic cats (Kays and DeWan, 2004). Land cover was
classied as: dense urban areas (continuous urban matrix, with
multi-storey buildings and vegetation cover <20%); sparse urban
areas (built-up areas dominated by single-storey buildings and vegetation cover between 20 and 50%); rural areas (single buildings
or small groups, vegetation cover between 50 and 80%); scattered
buildings (isolated buildings in a landscape whose vegetation cover
was 8095%); and non-urban areas (vegetation cover >95%).
To explore differences in cat management by owners in different land use types, we submitted a questionnaire to cat owners
in veterinary clinics, pet shops as well as door-to-door in different
urban conditions in three regions of Central Italy (Lazio, Umbria and
Abruzzo), covering the previously described categories. Cat owners
were asked to communicate (a) whether their cat was allowed to
get access to outdoor spaces, (b) the amount of time spent outside in
daytime (c) whether cats had access to outdoor during night time,
and (d) how many bats their cat brought home in the last three
years.
We determined the association between rescue causes of bats
and respectively month, season, gender and land cover by chisquare tests on contingency tables. The same test was applied to the
cats owner surveys data: in that case we tested whether the numbers of bats brought home by cats were associated respectively with
the time spent outdoor in daytime and with outdoor access during
the night, as well as with land cover. Signicant (p < 0.05) chi-square
tests were followed by an analysis of residuals to determine the
contribution of each category to the result (Haberman, 1973). All
tests were performed with R rel. 2.14.0 (http://www.R-project.org).
We obtained records of bats admitted to four wildlife rescue
centres originating from 13 Italian regions, but mainly from Lazio,
Toscana (central Italy) and Veneto (northern Italy; Fig. 1). Of 1012
bats admitted to the centres, 115 (11.3%) had been preyed upon by
a cat. Rescued bats belonged to 12 species, for all of which except
Myotis bechsteinii cases of predation by cats had been recorded
(Table 1).
In agreement with our hypothesis, only 2.4% of newborns or
juveniles admitted had been caught by cats whereas adults were
much more frequent: of 341 adult records, cat predation was
the most frequent rescue cause (28.7%), followed by debilitation
(23.3%), impact (18.2%), and removal from roost (4.6%); unknown
causes accounted for 25.2% of cases. Of 671 records of newborns
and juveniles, 90.0% were young bats accidentally fallen from roost,
other causes accounting for 10% of the dataset. Because the method
we adopted for ageing bats does not permit to distinguish between
ying and non-ying juveniles, as epiphyseal plates can be seen
on young bats for a few weeks after they become volant (BrunetRossini and Wilkinson, 2009), at least a few records from our

Please cite this article in press as: Ancillotto, L., et al., Curiosity killed the bat: Domestic cats as bat predators. Mammal. Biol. (2013),
http://dx.doi.org/10.1016/j.mambio.2013.01.003

G Model
MAMBIO-40595; No. of Pages 5

ARTICLE IN PRESS
L. Ancillotto et al. / Mammalian Biology xxx (2013) xxxxxx

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

Adults

Kuhls pipistrelle (Pipistrellus kuhlii)

Savis pipistrelle (Hypsugo savii)
European free-tailed bat (Tadarida teniotis)
Common/soprano pipistrelle (Pipistrellus
pipistrellus and P. pygmaeus)
Common serotine (Eptesicus serotinus)
Leislers bat (Nyctalus leisleri)
Brown long-eared bat (Plecotus auritus)
Schreibers bat (Miniopterus schreibersii)
Nathusius pipistrelle (Pipistrellus nathusii)
Bechsteins bat (Myotis bechsteinii)
Natterers bat (Myotis nattereri)
Lesser horseshoe bat (Rhinolophus hipposideros)
Unidentied

150 (38)
120 (35)
14 (2)
5 (5)

Juveniles
331 (4)
171 (7)
22 (0)
37 (2)

4 (1)
3 (2)
2 (1)
2 (2)
1 (1)
0
1 (1)
1 (1)
38 (10)

1 (0)
2 (1)
2 (0)
0
0
1 (0)
0
0
104 (2)

TOT

341 (99)

671 (16)

category juvenile may have been volant young bats: thus, the difference we detected between adults and juveniles preyed by cats
may be even stronger. Such young bats were admitted exclusively
during summer months, i.e. in the reproductive season (from June
to September, see Fig. 2af), so that the following results refer to
the adult-sample only.
As predicted, house-roosting bats, including Savis pipistrelles
(Hypsugo savii), European free-tailed bats (Tadarida teniotis) and
Kuhls and common/soprano pipistrelles (Pipistrellus kuhlii and P.
pipistrellus/pygmaeus) approached 80% of the total sample of predation on adults, but this also featured, albeit occasionally, species
more often roosting in caves (Miniopterus schreibersii) or trees (Pipistrellus nathusii, Nyctalus leisleri).
The relative frequencies of the rescue causes recorded differed across months (2 = 39.667, df = 12, p = 0.012) and seasons
(2 = 15.082, df = 3, p = 0.013), i.e. spring (AprilJune), summer (JulySeptember), autumn (OctoberDecember) and winter
(JanuaryMarch). Although predation by cats was recorded

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 rescue
peaked in mid-summer months, except deliberate removal from
roost which mainly occurred in winter and probably concerned
torpid subjects which people easily took from roosts (Fig. 2d).
Female adult bats were preyed upon by cats more frequently
than males, (2 = 5.444, df = 1, p = 0.019) accounting for 68.7% of
the sample, whereas no inuence of gender was detected on
the 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 bats
was associated with land cover (2 = 39.096, df = 16, p = 0.001),
being higher in rural areas as well as in areas characterized by
scattered buildings, whereas impact events and unknown causes of
rescue were more frequent in dense urban areas; all other causes
showed no land cover association.
Land cover also inuenced the distribution of rescue events for
newborns or non-volant juveniles (2 = 38.315, df = 16, p = 0.001):
debilitation, impact and unknown cause records were more frequent in dense urban areas, whereas predation by cats and
juveniles fallen from roost were more frequent in sparse and rural
areas.
According to what reported by people who rescued the bats and
brought them to the rehabilitation centres, at least ten roosts (H.
savii, n = 4; P. pipistrellus/pygmaeus, n = 2; P. kuhlii, = n = 2; P. kuhlii
and H. savii mixed colonies, n = 2) were targeted by repeated cat
predation over 13 years. All cats involved had an owner and were
allowed to access terraces or gardens; they caught, on average,
7.3 4.3 bats per year (mean SD; n = 10). The highest predation
rate affected a P. kuhlii nursery (where in three years at least 47
bats were caught during roost emergence) which was accessible
to the cat at night; in another case (an H. savii nursery colony)
the colony disappeared after undergoing two years of numerous
predation events (Table 2).
Our cat owner survey (n = 106) showed that the time domestic cats were allowed to spend outside decreased along a

Fig. 2. Monthly relative frequencies of bat rescue causes from wildlife rescue centres in the 2009/2011 period. Black bars: adults; grey bars: juveniles; a = cat predation;
b = impact; c = debilitation; d = removal from roost; e = unknown reason; f = fallen from roost.

Please cite this article in press as: Ancillotto, L., et al., Curiosity killed the bat: Domestic cats as bat predators. Mammal. Biol. (2013),
http://dx.doi.org/10.1016/j.mambio.2013.01.003

G Model
MAMBIO-40595; No. of Pages 5

ARTICLE IN PRESS
L. Ancillotto et al. / Mammalian Biology xxx (2013) xxxxxx

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

Species

2009

2010

2011

Rome
Castelnuovo di Porto
Cassino
Rome
Rome
Rome
Rome
Rome
Villa Latina
Rome

Hypsugo savii
Pipistrellus pipistrellus/pygmaeus
Pipistrellus kuhlii
Pipistrellus pipistrellus/pygmaeus
Pipistrellus kuhlii & Hypsugo savii
Hypsugo savii
Hypsugo savii
Hypsugo savii
Pipistrellus kuhlii
Pipistrellus kuhlii & Hypsugo savii

12
7
15

8
6
20
4
5
5
7
5

4
12
4
8
6
6
4
2
7

rural-to-urban gradient. Cats in rural and sparse urban areas were

allowed to stay outdoor for longer than cats in denser urban areas
(2 = 52.735, df = 12, p < 0.001) with 34.4% of cats from rural and
urban-sparse areas getting outdoor access for 612 h during daytime versus only 7% of cats from dense urban settlements. More
than 41% of cats from rural and sparse-urban areas had regular
outdoor access at night-time, while this was possible only for the
7% of cats from dense urban areas (2 = 37.647, df = 9, p < 0.001).
Predation on bats was more common in rural and sparse urban
areas (2 = 8.923, df = 3, p = 0.030), involving 16.4% and 7% of cats
from rural/urban-sparse areas and urban-dense respectively; it
was more frequently performed by cats regularly staying outside
overnight (2 = 9.963, df = 3, p = 0.020), which were reported to prey
on average 2.0 1.3 bats/year (n = 13).
Our study shows that in the Italian peninsula cat predation
may at least locally represent a signicant threat to bats, being
the rst cause of admission of adult bats to rehabilitation centres.
Clearly, the actual impact of predation by cats on bat populations
is still unknown and our study was not designed to answer this
question. Species roosting in buildings were more frequent in our
sample, yet this was expected since our analysis concerned subjects
rescued by people, an event denitely more likely to occur near
houses.
As hypothesized, juveniles were less frequently caught than
adults despite being smaller and easier to subdue and handle for a
cat. In most cases the bat species we found to be affected by predation roost in crevices or ssures, where non-volant roosting bats
are effectively sheltered from cats unless they fall accidentally from
the roost (Serangeli et al., 2012). However, because newborns may
be more easily dismembered or eaten by a cat, or more frequently
overlooked by rescuers, this may have at least partly inuenced
our results leading to an underestimation of such small prey items
(Krauze-Gryz et al., 2012).
We also found that predation on bats mainly occurred in rural
areas, or areas characterized by single buildings interspersed with
large vegetation patches. Our cat owner survey conrmed that cats
are more frequently allowed to stay outside in rural or sparse urban
areas, where bat roosts are also common. Instead, owned cats in
denser urban settlements were kept indoor more often, a fact which
may account for the lower numbers of bats caught in such land
use situations, despite the possibly higher absolute density of cats
(Natoli, 1985).
The type of analysis we did inevitably underestimated the
impact on bats using roosts other than buildings, which may be
not negligible especially if feral cats are considered. Despite our
probable underestimation, 3 of the 11 species found to be affected
by cat predation typically roost in caves or trees (e.g. Schober
and Grimmberger, 1997). Altogether, these data are alarming
because they suggest that free-ranging cats may have a significant impact on bats (including those from species of special

conservation concern) both in human-modied and natural landscapes where their rescue, or the retrieval of their remains, is much
less probable and may go unnoticed.
Predation was more likely to concern adult females and occurred
in summer (i.e. in the reproductive season) when nurseries congregate. Cats are explorative mammals (Machado and Genaro,
2010), so they may be easily attracted by sensory cues including sound emitted by bats, the smell of droppings accumulated
near roost sites or the observation of ying bats at its entrance.
Bats seem to be particularly vulnerable to predation by cats when
leaving roosts (Irwin and Speakman, 2003; Scrimgeour et al.,
2012); the broad frequency range heard by cats (between 48 and
85,000 Hz) enable them to detect most bat echolocation and social
calls (Heffner and Heffner, 1985). In fact sound, and smell too,
are conspicuous cues helping predators to locate roosts (Gillette
and Kimbrough, 1970; Fenton, 1995; Irwin and Speakman, 2003)
and once a roost is found, repeated predation may cause signicant mortality (Scrimgeour et al., 2012, this work) not effectively
compensated by new births due to bats slow reproduction rate
(Racey, 1982). Moreover, pregnant females may also be easier targets to predators due to their limited manoeuvrability in ight
(Russo et al., 2007). Repeated predation is commonly performed
by individual cats which specialize both on bats (Scrimgeour
et al., 2012) and other prey (Churcher and Lawton, 1989; Tschanz
et al., 2011). Foraging bats too may be caught by cats when ying close to the ground (see DelPietro et al., 1994 for Desmodus
rotundus in central America), or gleaning prey from substrate a
hunting 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 and
DeWan, 2004; Medina and Nogales, 2009; Krauze-Gryz et al., 2012)
or extremely rare (Woods et al., 2003; Krauze-Gryz et al., 2012) in
the prey datasets of studies from different regions. This may be
due to the small size of most bats which make them easy to ingest,
so their remains are rarely found (as demonstrated by Chapman
and Kitchener in 1978, who rarely found bat remains among cat
prey items, but often detected them in stomach samples). However, bats were also rarely found in scat-gut contents analyzed by
Krauze-Gryz et al. (2012).
We are aware that our sample was biased in terms of habitats
and species considered because it only featured bats brought to
rescue centres. Nonetheless, data from such sources are still valuable in assessing the threats to urban wildlife and their ecological
correlates (Koenig et al., 2002; Shine and Koenig, 2001). Overall,
our assessment of the phenomenon, covering a broad geographical
area over a relatively long period, suggests that the threat posed by
domestic cats may be signicant especially on a local scale: recidivist cats systematically targeting the same site may prompt a colony
to move to a new roost, thus exposing the bats to higher predation risks or reproductive failure, or even eradicate it. Certainly,
this threat should be considered in conservation plans. Our view is
supported by Altringham (2011) who inferred from data by Woods
et al. (2003) that every year approximately 250,000 bats are killed
by cats in the UK.
Practices such as the demographic control of feral cats by sterilization and night-time indoor restriction of domestic cats, at least
during the reproductive season of bats, may certainly reduce their
impact. At least for birds and mice, prey deterrents such as collarmounted bells have proven useful to warn potential victims of the
presence of a cat (Ruxton et al., 2002; Woods et al., 2003): whether
they might also be useful to reduce predation on bats e.g. at roosts
(by delaying bat emergence when the predator is nearby) has yet
to be tested.

Please cite this article in press as: Ancillotto, L., et al., Curiosity killed the bat: Domestic cats as bat predators. Mammal. Biol. (2013),
http://dx.doi.org/10.1016/j.mambio.2013.01.003

G Model
MAMBIO-40595; No. of Pages 5

ARTICLE IN PRESS
L. Ancillotto et al. / Mammalian Biology xxx (2013) xxxxxx

Acknowledgements
We are grateful to the Italian League for the Protection of Birds
(LIPU) and particularly to the staff and volunteers of the wildlife
rescue centres that readily provided us with valuable data and
support: Centro Recupero Fauna Selvatica (Roma), Centro Recupero Uccelli Marini (Livorno), Centro Recupero Fauna Selvatica La
Fagiana (Pontevecchio di Magenta Milano) and Centro Recupero
Animali Selvatici (Padova). We also thank Fabio Bontadina, Xavier
Puig and one anonymous reviewer for their valuable comments on
a previous ms version.
References
Altringham, J.D., 2011. Bats from Evolution to Conservation. Oxford University
Press, Oxford, UK.
Anthony, E.L.P., 1988. Age determination in bats. In: Kunz, T.H. (Ed.), Ecological
and Behavioral Methods for the Study of Bats. Smithsonian Institution Press,
Washington DC/London, pp. 4758.
Barbour, R.W., Davis, W.H., 1969. Bats of America. University Press of Kentucky,
Lexington.
Beck, A.M., 1973. The Ecology of Stray Dogs. Purdue University Press, West Lafayette,
IN.
Brunet-Rossini, A.K., Wilkinson, G.S., 2009. Methods for Age Estimation and the
Study of Senescence in Bats. In: Kunz, T.H., Parsons, S. (Eds.), Ecological and
Behavioral Methods for the Study of Bats. Johns Hopkins University Press, Baltimore, pp. 15325.
Chapman, A., Kitchener, D.J., 1978. Mammals of Durokoppin and Kodj Kodjin nature
reserves. Records W. Aust. Mus. suppl. 7, 4954.
Churcher, P.B., Lawton, J.H., 1989. Beware of well-fed felines. Nat. Hist. 7, 4047.
Coleman, J.S., Temple, S.A., 1993. Rural residents free-ranging domestic cats: a survey. Wildl. Soc. B 21, 381390.
Delpietro, H., Konolsaisen, F., Marchevsky, N., Russo, G., 1994. Domestic cat predation
on vampire bats (Desmodus rotundus) while foraging on goats, pigs, cows and
human beings. Appl. Anim. Behav. Sci. 39, 141150.
Dickman, C.R., 1996. Overview of the Impacts of Feral Cats on Australian Native
Fauna. Australian Nature Conservation Agency, Canberra.
Dietz, C., von Helversen, O., Nill, D., 2009. Bats of Britain Europe and Northwest
Africa. A and C Black, London, UK.
Fenton, M.B., 1995. Constraints and exibility; bats as predators, bats as preys. Symp.
Zool. Soc. Lond. 67, 277290.
Galetti, M., Sazima, I., 2006. Impact of feral dogs in an urban Atlantic forest fragment
in southeastern Brazil. Natureza Conservaco 5, 146151.
Gillette, D.D., Kimbrough, J.D., 1970. Chiropteran mortality. In: Slaughter, B.H., Walton, D.W. (Eds.), About Bats. Southern Methodist University Press, Dallas.
Haberman, S.J., 1973. The analysis of residuals in cross-classied tables. Biometrics
29, 205220.
Heffner, R.S., Heffner, H.E., 1985. Hearing range of the domestic cat. Hear. Res. 19,
8588.
Irwin, N.R., Speakman, J., 2003. Azorean bats Nyctalus azoreum, cluster as they
emerge from roost, despite the lack of avian predators. Acta Chiropt. 5, 185192.
Jones, G., Van Parijs, S.M., 1993. Bimodal echolocation in pipistrelle bats: are cryptic
species present? Proc. R. Soc. Lond. Biol. Sci. 251B, 119125.

Kays, R.W., DeWan, A.A., 2004. Ecological impact of inside/outside house cats around
a suburban nature preserve. Anim. Conserv. 7, 111.
Koenig, J., Shine, R., Shea, G., 2002. The dangers of Life in the city: patterns of activity, injury and mortality in suburban lizards (Tiliqua scincoides). J. Herpetol. 36,
6268.
Krauze-Gryz, D., Gryz, J., Goszczynski, J., 2012. Predation by domestic cats in rural
areas of central Poland: an assessment based on two methods. J. Zool. (Lond.),
http://dx.doi.org/10.1111/j.1469-7998.2012.00950.x.
Lepczyk, C.A., Mertig, A.G., Liu, J., 2003. Landowners and cat predation across ruralto-urban landscapes. Biol. Conserv. 115, 191201.
Liberg, O., 1984. Food habits and prey impact by feral and house-based domestic
cats in a rural area in southern Sweden. J. Mammal. 65, 424432.
Machado, J.C., Genaro, G., 2010. Comportamento exploratrio em gatos domsticos (Felis silvestris catus Linnaeus, 1758): uma reviso. Archit. Vet. Sci. 15,
107117.
Mastrobuoni, G., Gaiba, G., Ragno, R., 2005. Prima segnalazione per il Lazio (Italia
Centrale) di Vespertilio di Bechstein, Myotis bechsteinii (Chiroptera: Vespertilionidae). Boll. Mus. Reg. Sci. Nat. Torino 22, 525530.
Medina, F.M., Nogales, M., 2009. A review on the impacts of feral cats (Felis silvestris
catus) in the Canary Islands: implication for the conservation of its endangered
fauna. Biodivers. Conserv. 18, 829846.
Natoli, E., 1985. Spacing pattern in a colony of urban stray cats (Felis catus L.) in the
historic center of Rome. Appl. Anim. Behav. Sci. 14, 289304.
Phillips, S., Coburn, D.K., James, R., 2001. An observation of cat predation upon an
eastern blossom bat Syconycteris australis. Aust. Mammal. 23, 5758.
Racey, P.A., 1982. Ecology of bat reproduction. In: Kunz, T.H. (Ed.), Ecology of Bats.
Plenum Press, New York.
Rodrguez-Durn, A., Lewis, A.R., 1985. Seasonal predation by Merlins on sooty mustached bats in Western Puerto Rico. Biotropica 17, 7174.
Rosina, V.V., Shokhrin, V.P., 2011. Bats in the diet of owls from the Russian far East,
southern Sikhote Alin. Hystrix It. J. Mammal. 22, 205213.
Russo, D., Cistrone, L., Jones, G., 2007. Emergence time in forest bats: the inuence
of canopy closure. Acta Oecol. 31, 119126.
Ruxton, G.D., Thomas, S., Wright, J.W., 2002. Bells reduce predation of wildlife by
domestic cats (Felis catus). J. Zool. (Lond.) 56, 8183.
Schober, W., Grimmberger, E., 1997. The Bats of Europe and North America. T.F.H.
publications, Neptune.
Scrimgeour, J., Beath, A., Swanney, M., 2012. Cat predation of short-tailed
bats (Mystacina tuberculata rhyocobia) in Rangataua forest, Mount
Ruapehu, Central North Island, New Zealand. New Zeal. J. Zool. (Lond.),
http://dx.doi.org/10.1080/03014223.2011.649770.
Serangeli, M.T., Cistrone, L., Ancillotto, L., Tomassini, A., Russo, D., 2012. The postrelease fate of hand-reared orphaned bats: survival and habitat selection. Anim.
Welf. 21, 918.
Shine, R., Koenig, J., 2001. Snakes in the garden: an analysis of reptiles rescued by
community-based wildlife carers. Biol. Conserv. 102, 271283.
Speakman, J.R., 1991. The impact of predation by birds on bat populations in the
British Isles. Mammal. Rev. 21, 123142.
Thorington, K.K., Bowman, R., 2003. Predation rate on articial nests increases with
human housing density in suburban habitats. Ecography 26, 188196.
Tschanz, B., Hegglin, D., Gloor, S., Bontadina, F., 2011. Hunters and non-hunters:
skewed predation rate by domestic cats in a rural village. Eur. J. Wildl. Res. 57,
597602.
Van Heezik, Y., Smyth, A., Adams, A., Gordon, J., 2010. Do domestic cats impose an
unsustainable harvest on urban bird populations? Biol. Conserv. 143, 121130.
Woods, M., McDonald, R.A., Harris, S., 2003. Predation of wildlife by domestic cats
Felis catus in Great Britain. Mammal. Rev. 33, 174188.

Please cite this article in press as: Ancillotto, L., et al., Curiosity killed the bat: Domestic cats as bat predators. Mammal. Biol. (2013),
http://dx.doi.org/10.1016/j.mambio.2013.01.003