bs_bs_banner

Mammal Review ISSN 0305-1838

REVIEW

Bats as prey of diurnal birds: a global perspective

Peter MIKULA Department of Zoology, Faculty of Science, Charles University in Prague, Vinicn 7,
128 43 Praha 2, Czech Republic. E-mail: petomikula158@gmail.com
Federico MORELLI Biology Centre of the Czech Academy of Sciences, Institute of Hydrobiology, Na
Sdkch 7, 370 05 Cesk Budejovice, Czech Republic and INRA, AgroParisTech, UMR 1048 SADAPT, 16
rue Claude Bernard, F- 75005 Paris, France. E-mail: fmorellius@gmail.com
Radek K. LUCAN Department of Zoology, Faculty of Science, Charles University in Prague, Vinicn 7,
128 43 Praha 2, Czech Republic. E-mail: rlucan@centrum.cz
Darryl N. JONES Environmental Futures Research Institute, Griffith University, Nathan, Queensland
4111, Australia. E-mail: d.jones@griffith.edu.au
Piotr TRYJANOWSKI* Institute of Zoology, Poznan University of Life Sciences, Wojska Polskiego 71 C,
60-625 Poznan, Poland. E-mail: piotr.tryjanowski@gmail.com

Keywords
avian predation hypothesis, bats, diurnal
birds, nocturnality, predation
*Correspondence author.
Submitted: 16 September 2015
Returned for revision: 1 October 2015
Revision accepted: 16 October 2015
Editor: KH
doi:10.1111/mam.12060

ABSTRACT
1. Predation is an important selective pressure that can influence prey species in
numerous ways. Predatorprey relationships are, however, poorly understood in
taxa not typically associated with these interactions; this is especially the case
when bats (Chiroptera) are the prey.
2. The main aim here is to review and synthesise global information on the predation of bats by birds of prey (Accipitriformes and Falconiformes) and other
diurnal bird groups.
3. We compiled data on incidences of predation of bats by diurnal birds, through
an extensive multilingual study of bibliographic and Internet-based sources. Scientific papers were found mainly via the Thomson Reuters (Web of Science and
Zoological Record) and Scopus databases, Google Scholar, and Google Books.
Additional data were found through Internet searches of trip reports, images and
videos carried by Google, Google Images, Flickr, and YouTube.
4. In total, more than 1500 cases of bats being attacked by diurnal predatory birds
were obtained. Reports were documented from every continent (except Antarctica) and from 109 countries, and were thus distributed globally between 70N
and 43S. Overall, we found evidence for predation of bats by 143 species of
diurnal raptors (Accipitriformes 107 spp. and Falconiformes 36 spp.) and by 94
non-raptor bird species from 28 families. At least 124 and 50 bat species were
taken as prey by raptors and by other diurnal bird species, respectively.
5. Attacks on bats by diurnal raptors were found to be distributed globally and
were present in the majority of extant raptor lineages. Attacks on bats by other
diurnal birds were also occasionally recorded. Furthermore, the majority of extant
bat families featured as prey. These results strongly suggest that predation by birds
may act as a major factor affecting the scarcity of daytime activity in bats and as a
driver in the evolution of bat nocturnality.

Mammal Review (2016) 2016 The Mammal Society and John Wiley & Sons Ltd

Predation of bats by diurnal birds

INTRODUCTION
Predation is an important selective pressure that can affect
individuals, populations, and communities of prey species
(Begon et al. 2005). Although predatorprey interactions
are well known for a wide variety of animals, for certain
taxa the diversity of their predators and the effects that predation impose on their populations are poorly understood.
This is especially the case for bats (Chiroptera; Lima & Dill
1990, Caro 2005). This taxon is the second largest mammalian order, comprising more than 1200 species (Gunnell &
Simmons 2012), and is among the most taxonomically and
ecologically diverse groups of mammals (Kunz & Fenton
2003).
In general, bats are K-strategists with long life spans and
small litter sizes (Kunz & Fenton 2003), and life-history
traits directly related to effective avoidance of predation
(Speakman 1991a, 1995, Rydell et al. 1996). Because of their
life history and agile flight, bats have traditionally been
regarded as having relatively few natural predators and
rarely experiencing predation; some authors have even
described bats as UFOs: uncatchable flying objects
(Jedrzejewska & Jedrzejewski 1998). This perspective is
probably associated with the scattered nature of bat predation records; reports are typically published in local journals
that may be difficult to access or in monographs and species
accounts from specific geographic regions (Sparks et al.
2000). Moreover, many relevant reports are limited to nonEnglish sources (reducing accessibility for readers from different language backgrounds), and even those accounts may
sometimes contain comprehensive reviews of predation of
bats at only national levels (e.g. Bekker & Mostert 1991,
Haensel & Smmer 2002, Duquet & Nadal 2012). Furthermore, because of their nocturnality, bats are relatively difficult to observe and study, exacerbating the low chances of
observing behavioural events. An accumulation of reports
on predation of bats is, however, gradually changing traditional knowledge on the topic, suggesting that bats may
have a wide suite of natural predators including arthropods,
fishes, amphibians, reptiles, and mammals (Sparks et al.
2000, Molinari et al. 2005, Esbrard & Vrcibradic 2007,
Nyffeler & Knrnschild 2013, de Noronha et al. 2015).
Available literature indicates that the most common and
effective predators of bats are, however, birds, especially
owls (Strigiformes; Speakman 1991a, Obuch 1998, del Hoyo
et al. 1999, Lesinski et al. 2009a, b), which prey upon them
during their coincidental nocturnal activities. Perhaps unexpectedly, due to the apparently differing activity schedules,
another important group of potential predators of bats are
diurnal avian raptors falcons, hawks, and eagles that
hunt predominantly during the day.
Previous researchers have found that predation by birds
could be responsible for about 10% of annual mortality of
2

P. Mikula et al.

bats in the temperate zone (Speakman 1991a). Over 90%

of bats taken by predators are apparently killed by owls,
whereas only 5% fall prey to diurnal raptors (Speakman
1991a). In the temperate zone, however, daytime flying by
bats is typically very unusual. Speakman (1990), for
example, mentioned observations of only 420 cases of
diurnal activity by bats (collected during a British national
survey in 19861987) and suggested that daytime flying is
at least 100 times less frequent than flying during the
night. However, supported by releasing experiments, the
diurnal predation rate on bats was estimated to be 100
1000 times higher than the nocturnal predation rate when
standardised for the duration of bat activity at night and
during the day (Speakman 1991b, Speakman et al. 1994,
Speakman 1995). Despite their limited diurnal activity
(Speakman 1990), numerous reports of predation of bats
by numerous raptors have been found worldwide
(del Hoyo et al. 1994, Sparks et al. 2000, Ferguson-Lees &
Christie 2001, Haensel & Smmer 2002). This seemingly
widespread phenomenon has, however, received very
little attention even though it may influence many
aspects of bat behaviour. Indeed, the interaction between
bats and diurnal predatory birds could act as an
important adaptive pressure potentially influencing the
evolution of bat primary activity into nocturnality
(Rydell & Speakman 1995, Speakman 1995, Lima &
OKeefe 2013).
The aim of the present study is to review and synthesise
global data on predation of bats by diurnal birds of prey
(Accipitriformes and Falconiformes) and other diurnal bird
taxa. Although predation of bats by diurnal avian predators
appears to be underestimated and insufficiently studied, we
hypothesized that it is likely to be more frequent and more
widely distributed than previously thought.

METHODS
Data searching and exclusions
An extensive study of bibliographic sources was conducted
in order to detect all relevant information on predation of
bats by diurnal birds. We defined predation as consumption
of one organism (the prey) by another organism (the predator), in which the prey is alive when the predator first
attacks it (Begon et al. 2005). Therefore, we used only
records that could be considered, on the basis of dietary and
morphological characteristics of birds, as attacks for the
purpose of capture and subsequent prey consumption,
although in many cases, the prey was not successfully
caught or when caught it was not consumed by the predator. Furthermore, we focused mainly on predation events
that took place during the day: as daytime predation we
Mammal Review (2016) 2016 The Mammal Society and John Wiley & Sons Ltd

P. Mikula et al.

considered all cases where diurnal bird predators were able

to orientate themselves and hunt bats visually (including at
dawn and dusk); we also included night attacks on bats illuminated by artificial light.
In our search for scientific papers, we utilised mainly the
Thomson Reuters (Web of Science and Zoological Record)
and Scopus databases, Google Scholar, and Google Books.
When a suitable paper was found, its references (backward
search) and citation records (forward search) were used to
search for other relevant articles. Additionally, we searched
for bat predation records in previously undescribed sources
by conducting Internet searches for trip reports, images, and
videos carried by Google, Google Images, Flickr, and
YouTube. We searched not only in the English language and
Latin script but also in numerous other national languages
(for language index see Appendix S1) and scripts, although
the search was restricted by our knowledge of particular
languages.
For analyses we included only reports where birds had
been observed unequivocally attacking or catching bats or
where such an event could be inferred from the text. We did
not distinguish between successful and unsuccessful
hunting attempts when the observation satisfied this criterion. To avoid pseudoreplication, we excluded records from
secondary sources (mainly books and reviews) if we were
able to obtain the original source; if we were not, records
from secondary sources were treated similarly to the
primary records. Where multiple sources mentioned the
same predation event, we used only the oldest source in the
data set for analysis. Our data set did not include cases
where birds were observed sitting or flying near roosting
places of bats if no direct observations of bat hunting had
been recorded.
Some recorded attacks on bats by diurnal birds did not
satisfy our criteria of being predation attempts. Nonpredatory attacks usually belonged to the class of defensive
bird behaviour known as mobbing. Thus, we excluded cases
of birds chasing bats, or where bat deaths were caused by
birds unlikely to consume the bats as food, such as peafowl
Pavo sp., swifts (Apodidae), swallow Hirundo rustica, blackbirds Agelaius spp., and some tits (Paridae; Tugendhat 1966,
Kervyn 1998, Sparks et al. 2000, Suzuki 2012). We also
excluded cases where birds attacked bats or killed them
during competition for nesting holes, as observed in woodpeckers (Picidae), starling Sturnus vulgaris, and parakeet
Psittacula krameri (Mason et al. 1972, Bekker & Mostert
1991, Kervyn 1998, Menchetti et al. 2014). In addition, we
excluded cases where small diurnal raptors were observed
chasing large bat species, when successful predation was
highly unlikely (e.g. falcons Falco amurensis and Falco
subbuteo were observed chasing Eidolon helvum bats in
Kasanka forest, Zambia; Anonymous 2010, Willems pers.
comm.).
Mammal Review (2016) 2016 The Mammal Society and John Wiley & Sons Ltd

Predation of bats by diurnal birds

Maps and relevant geographical information

In the preparation of maps and in our geographical assessment, we excluded records with highly generalised and geographically unspecified information on predation of bats.
We included only records with specified geographical positions at the spatial level of country or smaller geographical
areas (e.g. Guatemala or Malay Peninsula). In each case we
specified the exact geographical position with restrictions
detailing how this was known. When the country or geographical location was not specified in greater detail, geographical coordinates were marked in the centre of the area
that resulted from superimposing the position given and the
bird and/or bat species distribution (if available) at the site.
Because of the various methods used by the people collecting the data (such as direct observations, pellet analyses,
collection of prey remnants under nests, prey delivery
observations to nest, excavations on nesting sites, trip
reports, photo and video records, and personal communications) and the differing status of the records (from anecdotal notes to systematic long-term research), the number
of attacks by birds on bats was counted as the number of
sources (papers, trip reports etc.) in which hunting attempts
were described, even when predation events were temporally separated and included more than one individual bat
or bats of several taxa. However, if one source contained
records for more diurnal avian species or from multiple
localities (natural habitats: distance of at least a few
kilometres; urban areas: on the level of different cities), we
noted each locality as a separate case.
All country and bat records for each avian species are
included in Appendices S2 and S3; for references see Appendix S4. Data on global diversity, body lengths and weights of
the raptors and other diurnal birds were obtained from the
ornithological literature (del Hoyo et al. 2015).

Geographical Information System analyses

All information collected was transformed into maps using
ArcGIS 10.1 (ESRI, Redlands, California, USA), an integrated collection of Geographical Information System software products (Anonymous 2012). In this way, all recorded
sites were georeferenced.
In order to investigate the pattern of records, we
employed the kernel density function, a non-parametric
technique used to estimate the probability density function
of a random variable. Kernel density estimation is a fundamental data-smoothing technique, where inferences about
the population of data are made, based on a finite data
sample. This estimates the density function directly from
the data without making any assumptions about the underlying distribution and constitutes a general density estimation tool, applied directly in ArcGIS. In this study, kernel
3

Predation of bats by diurnal birds

density estimation was applied to map the distribution of

records, on the basis of the type of source (scientific literature or Internet search).

Visualisation of bat predators in the raptor

phylogenetic tree
To visualise bat predation activity in diurnal raptors, a
phylogenetic tree of extant raptor genera was constructed
based on consensus avian phylogenetic tool available at
http://birdtree.org/ (Jetz et al. 2012). As an outgroup we
used gulls (Laridae). As the source of our consensus tree
we used the Hackett All Species tree with 1000 randomly
generated trees. The most credible tree was then determined using the tool TreeAnnotator v1.8.2 in the software
BEAST v1.8.2 (Drummond & Rambaut 2007). The
consensus tree was then graphically adjusted in FigTree
v1.4.2 (Andrew Rambaut, University of Edinburgh, UK;
http://tree.bio.ed.ac.uk/software/figtree/). We excluded the
sister group of Accipitriformes, new world vultures
(Cathartiformes), because of an absence of bat hunting
behaviour in whole lineage. In addition, other diurnal
non-raptorial bird groups were not visualised due to fragmentary records.

RESULTS
Geographic distribution of attacks on bats
by diurnal birds
Altogether we obtained 1530 cases of predation of
bats by diurnal raptors and other diurnal birds (711
Accipitriformes, 608 Falconiformes, and 211 non-raptor
birds). Attacks on bats by diurnal birds were reported from
109 countries from all continents with the exception of Antarctica. Records of raptors attacking bats were obtained
from 107 countries (26 countries in Africa, 27 in Asia, five
in Australia and Oceania, 26 in Europe, 13 in North and
Central America, 10 in South America; Fig. 1a, b). Records
of non-raptors attacking bats were obtained from 45 countries (eight countries in Africa, 16 in Asia, one in Australia
and Oceania, 14 in Europe, four in North and Central
America, two in South America; Fig. 1c). The kernel density
function revealed that the density of scientific literature
records was highest in developed countries (Fig. 2a),
whereas Internet-derived records reached high densities also
outside these countries (Fig. 2b).
The northernmost observations of predation on bats by
Falconiformes, Accipitriformes, and other birds were from
Troms county (northern Norway) at 6900N, 1900E,
Lule (northern Sweden) at 6530N, 2140E and Tafjorden
(western Norway) at 6214N, 0725E, respectively. In the
first case, Falco columbarius attacked Eptesicus nilssonii
4

P. Mikula et al.

(Frafjord 2012); in the second case, Accipiter nisus attacked

the same species (Rydell 1992). Larus argentatus, Larus
canus, and Corvus cornix were observed sitting near maternity roosts of Pipistrellus pygmaeus or trying to catch them
near Tafjorden (Michaelsen et al. 2014, Michaelsen pers.
comm.).
The most poleward records for Falconiformes from the
Southern Hemisphere were from Puerto Madryn (Patagonia, Argentina) at 4246S, 652W and Mount Cookson
(South Island, New Zealand) at 4233S, 1738E. In the first
location, Falco sparverius was preying probably upon
Tadarida brasiliensis (Pagnoni 2013). In the second case,
archaeological excavations revealed skeletal remains of
Mystacina robusta and Mystacina tuberculata at predatoraccumulated deposits attributed to Falco novaeseelandiae
(Worthy & Holdaway 1995). For Accipitriformes, two individuals of Eptesicus vulturnus were found in the nonbreeding diet of Accipiter novaehollandiae in Tasmania
(Australia) approximately at 4140S, 14737E (Olsen et al.
1990). Of non-raptors, Ojeda and Chazarreta (2006)
observed Campephilus magellanicus delivering a bat to its
nestlings in the Nahuel Huapi National Park (Patagonia,
Argentina) at 4108S, 7112W. In the Australian subtropics, Strepera graculina and Dacelo novaeguineae were
observed attacking released Chalinolobus morio, Eptesicus
regulus, Nyctophilus gouldi, and Vespadelus darlingtoni in the
Central Highlands of Victoria (Australia) at 3730S,
14345E (Speakman et al. 1994).

Distribution of attacks on bats within

groups of diurnal birds
We found evidence of predation on bats in 143 species of
diurnal raptors from 47 genera and three families
(Accipitridae 106 spp., Falconidae 36 spp., Pandionidae 1
sp.; for details, see Appendix S2). Bat-hunting by raptors
was found in about 42% of all species and in 61% of genera
in the family Accipitridae, and in more than 56% and 36%,
respectively, in the family Falconidae. This suggests that predation of bats is a widespread phenomenon present in the
majority of extant raptor groups (Fig. 3).
We found evidence of bat hunting in 94 species of nonraptorial diurnal birds from 28 families (Alcedinidae 9
spp., Anatidae 1 sp., Ardeidae 5 spp., Bucerotidae 11 spp.,
Cariamidae 1 sp., Ciconiidae 1 sp., Coraciidae 1
sp., Corvidae 19 spp., Cotingidae 1 sp., Cracticidae 4 spp.,
Cuculidae 3 spp., Dendrocolaptidae 1 sp., Dicruridae 2
spp., Icteridae 2 spp., Laniidae 9 spp., Laridae 8 spp.,
Malaconotidae 3 spp., Meropidae 1 sp., Momotidae 1 sp.,
Muscicapidae 1 sp., Paridae 1 sp., Phoeniculidae 1
sp., Picidae 2 spp., Ramphastidae 2 spp., Sturnidae 1 sp.,
Turdidae 1 sp., Tyrannidae 1 sp., Vireonidae 1 sp.; Appendix
S3). The three families with the highest incidence of bat
Mammal Review (2016) 2016 The Mammal Society and John Wiley & Sons Ltd

P. Mikula et al.

Predation of bats by diurnal birds

(a)

(b)

(c)

Fig. 1. Global distribution of attempted and

actual predation on bats by Accipitriformes (a),
Falconiformes (b), and other groups of diurnal
birds (non-raptors; c). Each documented case
of predation on bats is represented by a grey
dot; each black dot represents a case for each
predator group identified in (a), (b), and (c).

hunting were Cracticidae (33% of all species), Laniidae

(29%), and Bucerotidae (18%).

At least 124 species of bats from 48 genera and 11

families (Emballonuridae 4 spp., Hipposideridae 5 spp.,
Megadermatidae 1 sp., Molossidae 18 spp., Mormoopidae 3
spp., Mystacinidae 2 spp., Nycteridae 2 spp., Phyllostomidae
8 spp., Pteropodidae 22 spp., Rhinolophidae 6 spp.,
Vespertilionidae 53 spp.) were identified as victims of pre-

dation attempts by diurnal raptors. We also recorded one

capture of Noctilio sp., extending the bat family diversity for
Noctilionidae. A few other bat taxa were found to be preyed
upon by raptors (Appendix S2). Altogether, the recorded
bats represent more than 10% and 63% of global diversity
of bat species and families, respectively.
In more than 42% of the 1319 cases of bats preyed upon
by raptors, the bats were not identified even to the family
level (typically, authors were unable to identify the bats,
while in a few cases, we were unable to obtain the full text of
the original source). The highest numbers of reported pre-

Mammal Review (2016) 2016 The Mammal Society and John Wiley & Sons Ltd

Bat species caught by diurnal bird predators

Predation of bats by diurnal birds

P. Mikula et al.

Fig. 2. Kernel density maps based on source

type. Scientific literature records (a) reach
highest densities mainly in temperate and wellstudied areas, whereas the density of Internet
records (b) shows a more diffuse pattern.

dation (in descending number of cases) concerned bats

from the family Molossidae, followed by Vespertilionidae,
Pteropodidae, Phyllostomidae, Emballonuridae, Hipposideridae, and Rhinolophidae; all other families constituted
less than 1% of records (Table 1).
At least 50 species of bats from 24 genera and six families
(Emballonuridae 1 sp., Hipposideridae 1 sp., Molossidae
4 spp., Phyllostomidae 3 spp., Pteropodidae 6 spp.,
Vespertilionidae 35 spp.) were recorded as victims of

predation or predation attempts by non-raptorial diurnal

birds (Appendix S3). Identified bat taxa made up about 4%
and 32% of global diversity of bat species and families, respectively. Of the 211 records of bats being preyed upon by nonraptors, about 42% represented unidentified bats. After
unidentified bat records were removed, the bats of the family
Vespertilionidae were most frequently associated with predation, followed by Molossidae, Pteropodidae, Phyllostomidae,
Emballonuridae, and Hipposideridae (Table 1).

Fig. 3. Genera of birds known to hunt bats,

indicated in a raptor phylogeny. Visualisation
of the consensual phylogenetic tree of raptor
genera was made by the avian phylogenetic
tool available at http://birdtree.org/ (Jetz et al.
2012). Genera in which at least one bathunting species was present are highlighted in
black; others are grey. Bats as targets of
diurnal raptors predatory attempts were
recorded in the majority of extant lineages and
genera.

Mammal Review (2016) 2016 The Mammal Society and John Wiley & Sons Ltd

P. Mikula et al.

Table 1. Bat families taken by diurnal birds.

Percentages of predation records containing
identified bats are grouped by bat family and
summarised for three groups of diurnal avian
predators (Accipitriformes: n = 426;
Falconiformes: n = 335; other birds: n = 122)

Predation of bats by diurnal birds

Bat family

Taken by
Accipitriformes (%)

Taken by
Falconiformes (%)

Taken by other birds

(non-raptors; %)

Emballonuridae
Hipposideridae
Megadermatidae
Molossidae
Mormoopidae
Mystacinidae
Noctilionidae
Nycteridae
Phyllostomidae
Pteropodidae
Rhinolophidae
Vespertilionidae

1.6
1.9
0.2
43.2
0.2
0
0
0.7
3.3
29.3
1.4
22.8

1.8
0
0.6
32.2
1.2
0.3
0.3
0
2.7
3.9
0.6
58.8

0.8
0.8
0
9
0
0
0
0
4.9
8.2
0
77

In addition to the traditionally accepted view that owls are

the major avian predators of bats, here we demonstrate that
bats have regularly been observed being caught by a wide
range of diurnal birds. Our results suggest that bats are at
significant risk of predation during daylight hours and are
supportive of the avian predation hypothesis for the evolution of nocturnality in bats (Speakman 1995). It seems that
bird species capable of catching and consuming bats do so
opportunistically, although hunting success, proportion,
and species spectra of bats attacked differ among avian taxa.
Almost all bat-hunting bird taxa, including raptors, are
predominantly predators of prey types other than bats (del
Hoyo et al. 1994, Ferguson-Lees & Christie 2001). In fact,
Macheiramphus alcinus is the only species of diurnal bird
(and probably the only vertebrate predator) that consistently preys predominantly on bats (Chapin 1932, Fenton
et al. 1977, Black et al. 1979). It is also the sole diurnal
raptor with several unique morphological and behavioural
adaptations that enable it to feed primarily on bats during
crepuscular periods. Macheiramphus alcinus has shifted its
dominant activity to days with reduced visibility, when its
large eyes provide enhanced vision (Ferguson-Lees &
Christie 2001). The species enlarged gape suggests convergent evolution with swifts, swallows, and nightjars
(Caprimulgiformes), and permits the ingestion of relatively
large intact prey (Jones et al. 2012). Its prey is, moreover,
typically captured, processed, and swallowed in flight
(Eccles et al. 1969, Black et al. 1979, Ballance 1981), so that
its behaviour resembles that of aerial feeders more than that
of other raptors that usually process their prey at a perch
(Ferguson-Lees & Christie 2001). In contrast, Falco
rufigularis feeds on bats only occasionally (birds and insects
are its main diet; Beebe 1950, Seijas 1996) and has no

special adaptations; its gape and eye size are proportionally

comparable with those of other diurnal raptors (Hall et al.
2009, Jones et al. 2012).
On a local level, however, small and medium-sized bats
most often of the families Vespertilionidae and Molossidae
occur in the diet of several falcons such as Falco rufigularis,
Falco sparverius, Falco subbuteo, Falco tinnunculus, Falco
biarmicus, and Falco columbarius (Twente 1954, Black 1956,
Baker 1962, Van Jaarsveld 1988, Yosef 1991, Seijas 1996,
Martinez & Lee 2013, Mikula et al. 2013). Exceptions are
larger falcons (e.g. Falco peregrinus and Falco biarmicus)
that are also capable of successful predation of even larger
bats of the family Pteropodidae (Clunie 1972, Worthy &
Anderson 2009). Falconids appear to be very successful bat
predators and, locally, bats can represent a substantial part
of their diet (e.g. Baker 1962, Clunie 1972, Kirven 1976,
Seijas 1996, Worthy & Anderson 2009, Foysal 2015). Large
bats (mainly Pteropodidae) are also usually caught by
Haliaeetus, Hieraaetus, Aquila, Nisaetus, Polemaetus,
Stephanoaetus, and Pithecophaga eagles (e.g. Kennedy 1985,
Stuart & Stuart 2004, Welbergen 2006, Anonymous 2010,
Fam & Nijman 2011), but predation on smaller bats has
also been recorded (Fenton et al. 1994, van Beirs 2004,
Brown 2004). Smaller accipitrids prey frequently upon
smaller bats mainly of the Molossidae and Vespertilionidae
families (e.g. Baker 1962, Lindskog 1976, Manakadan &
Natarajan 1992, Rydell 1992, Fenton et al. 1994). When
attacking bats as novel prey, raptors occasionally exhibit
initial neophobia, although they quickly overcome
this to become successful predators of bats (Mikula et al.
2013).
On the other hand, records of predation of bats were
completely absent among South American caracaras
(Falconidae) and scavenging new world and old world vultures (except genus Gyps). Moreover, predation of bats was
not recorded in raptor taxa with specialised dietary
demands (e.g. the bee-eating genus Pernis, ichtyophagous

Mammal Review (2016) 2016 The Mammal Society and John Wiley & Sons Ltd

DISCUSSION
Bats as prey of diurnal birds

Predation of bats by diurnal birds

Ichtyophaga, snake-eating Circaetus, except in one case, and

malacophagous Rostrhamus and Helicolestes). We assume
that the absence of information on predation of bats by
members of the subfamily Harpiinae, as well as in a few
other taxa, was due primarily to a scarcity of data on their
diet.
In other birds (non-raptors), bats were frequently found
to be preyed upon by gulls (Holroyd & Beaubien 1983,
Speakman 1991b), hornbills (Bucerotidae; Kemp 2001) and
especially by larger passerines (Passeriformes; Schwan &
Hikes 1979, Boxall 1982, Sarkozi & Brooks 2003, Hernndez
et al. 2007, Farina et al. 2011, Mikula 2013). We found that
out of 28 families and 94 bat-hunting species of nonraptors, 14 families and 47 species belong to the passerine
clade. However, more than one-third of passerine families
and nearly 80% of species are placed in the superfamily
Corvoidea. Members of this superfamily, and particularly
the corvids (Corvidae) that represent 40% of passerine
predators of bats, are large-brained opportunistically
feeding generalists exhibiting very flexible behaviour and
successful adaptations to local conditions (reviewed by
Jacobs et al. 2014). These adaptations enable them to utilise
bats as a food source when the opportunity arises.
Overall, a considerable proportion of the bats that were
captured by birds represented common species in their
respective geographic regions. Our results suggest that bats of
the families Molossidae, Vespertilionidae, and Pteropodidae
are among those especially vulnerable to diurnal predation by
birds. We believe that certain adaptations make these bats
particularly vulnerable to predation by mainly visually oriented predators (such as diurnal birds): their relatively frequent diurnal activity (Vespertilionidae and Pteropodidae,
mainly genus Pteropus; e.g. Speakman 1990, Thomson et al.
1998), roosting in super-abundant communities even up to
several million members (Molossidae and Pteropodidae;
Sprunt 1951, Lee & Kuo 2001, Anonymous 2010), and resting
in open roosts (Pteropodidae; Kunz & Fenton 2003).
However, high prevalence of the aforementioned bat groups
in the diet of diurnal birds may simply result from the fact that
these three families are the most species-rich bat families.

P. Mikula et al.

In natural habitats, bats are regularly taken by predatory

birds near roosting places, where bats typically live in dense
aggregations. Many raptor species are attracted to such locations to prey upon the resident bats. Around the world,
there are many examples of such behaviour, especially in
tropical and subtropical regions. For example, at
Gomantong and Niah caves at Borneo, a wide variety of
raptor species have been observed sitting or catching bats

near cave entrances. Moreover, substantial amounts of

raptors remains are found directly inside or in the vicinity
of caves, indicating long-term use of these areas by them
(Smythies 1968, Thiollay 1983, Sheldon et al. 2001, Lim &
Cranbrook 2002, Stimpson 2009). Raptor aggregations near
caves are found worldwide; observations of bat hunting in
the vicinity of bat caves came, for instance, from the USA
(Stager 1941, Constantine 1948, Sprunt 1951, Twente 1954,
Black 1956, Baker 1962, Harden 1972, Looney 1972, Lee &
Kuo 2001), Mexico (Anonymous 2014a, van Beirs 2014),
Puerto Rico (Rodrguez-Durn & Lewis 1985), Burma
(Smythies 1953), Thailand (Beckemeyer 2000, Round et al.
2006, Dreyer 2010, Anonymous 2014b), Australia (Lewis
1987), South Africa (Laycock 1982), Uganda (van Beirs
2004), and Zambia (Black et al. 1979).
Hunting near caves or other roosting places of bats is not
an unknown phenomenon among a wide variety of arthropod and vertebrate predators (Twente 1956, Molinari et al.
2005, Esbrard & Vrcibradic 2007, Nyffeler & Knrnschild
2013). Black et al. (1987) even suggested that communally
roosting bats emerging in large numbers can represent
easily accessible prey and act as a selective force in the
evolution of specialised bat-hunting raptors such as
Macheiramphus alcinus; this species is capable of hunting
bats in the open air (Lendrum 1977) but prefers to hunt
near roosting places (Finch 1978, Black et al. 1979). Other
raptors also hunt bats near caves or other roosting places. In
the new world, Accipiter cooperii, Accipiter striatus, Buteo
jamaicensis, Buteo swainsonii, Circus hudsonius, Falco
peregrinus, and Falco sparverius were observed hunting
mainly Tadarida brasiliensis bats near several caves in the
USA and Puerto Rico (Sprunt 1950, Twente 1954, Black
1956, Baker 1962, Harden 1972, Looney 1972,
Rodrguez-Durn & Lewis 1985, Lee & Kuo 2001, Martinez
& Lee 2013). Around the Mexican bat cave El Volcn de los
Murcilagos several raptor species, including Accipiter
bicolor, Accipiter cooperii, Buteo albonotatus, Buteo
brachyurus, Buteo plagiatus, Buteogallus urubitinga,
Chondrohierax uncinatus, Falco rufigularis, Micrastur
ruficollis, Micrastur semitorquatus, and Rupornis magnirostris
(Boland 2009, Binns & Beer 2011, Anonymous 2014a, c, van
Beirs 2014), were observed feeding on communally roosting
bats. Very diverse raptor assemblages can also be found near
tropical caves in Thailand and Malaysia, where mainly
Tadarida plicata bats roost in huge aggregations; among
records mainly from Internet searches we found information on numerous bat-hunting raptor species: Accipiter
badius, Accipiter gularis, Accipiter trivirgatus, Accipiter
virgatus, Aviceda jerdoni, Butastur indicus, Buteo japonicus,
Falco peregrinus, Falco severus, Falco tinnunculus, Haliastur
indus, Ictinaetus malayensis, Lophotriorchis kienerii,
Macheiramphus alcinus, Nisaetus nanus, and Spilornis cheela
(Smythies 1968, Thiollay 1983, Beckemeyer 2000, Lim &

Mammal Review (2016) 2016 The Mammal Society and John Wiley & Sons Ltd

Where are bats caught?

NATURAL HABITATS

P. Mikula et al.

Cranbrook 2002, Stimpson 2009, Dreyer 2010, Anonymous

2014b). Additionally, numerous bat predation records came
from the vicinity of the Murchison Falls in Uganda, where
bats emerging from the cave are pursued by Aquila rapax,
Buteo buteo, Falco ardosiaceus, Falco cuvierii, Falco
peregrinus, Gypohierax angolensis, Haliaeetus vocifer,
Hieraaetus wahlbergi, Macheiramphus alcinus, and Milvus
migrans (Brown 2004, Dobbs & Twino-beku 2004, van Beirs
2004, Woolley & Casson 2008). Many of these records,
mainly from tropical areas, are derived not from the scientific literature but from Internet searches. Supported by our
analyses, Internet sources can provide important information on birdbat relationships in the tropics.
Another example of the capacity of raptors to exploit rare
but aggregated food sources is provided by the Fibwe bat
forest in Zambia, where pteropodids (mainly Eidolon
helvum) aggregate seasonally in vast numbers (up to 10
million individuals; Stuart & Stuart 2004, Anonymous 2010,
Willems pers. comm.). These aggregations are accompanied
by many species of medium-sized to large raptors (Aquila
nipalensis, Aquila spilogaster, Buteo buteo, Clanga clanga,
Clanga pomarina, Haliaeetus vocifer, Hieraaetus ayresii,
Milvus migrans, Polemaetus bellicosus, and Stephanoaetus
coronatus) that visit this site to prey upon roosting bats. Furthermore, highly unusual raptors such as vultures
(Gypohierax angolensis and Gyps africanus) also take advantage of the opportunity to prey on these bats (Stuart &
Stuart 2004, Anonymous 2010, Willems pers. comm.).
As well as hunting near caves and dense bat aggregations,
many raptor species were observed lurking near bat shelters
in cliffs, man-made structures, and vegetation, and attacking migrating or released bats, as described for several
accipitrids (Mumford 1980, Kemp & Rautenbach 1987, Byre
1990, Rautenbach et al. 1990, Fenton et al. 1994, Pikacha
et al. 2012) as well as falconids (Twente 1954, Meinesz et al.
1982, Thomsett 1987, Fenton et al. 1994, Yancey et al. 1996).
Aerial attackers of bats, such as Falco rufigularis, prefer to
hunt bats in or over forest canopy and above lakes and
rivers (Beebe 1950, Pierson & Donahue 1983, Robinson
1994, Parker 1997). In fact, attacks on bats away from bat
roosts are not exceptional events for open-air hunting or
ambush raptors preying upon bats (Borowski 1968,
Lindskog 1976, Van Jaarsveld 1988, Roworth & Wright
1989, Byre 1990, Yosef 1991, Michalak 1997, Rollie &
Christie 2006).

Predation of bats by diurnal birds

by Russo & Ancillotto 2015). Attacks on bats by falconids

(Fry 1964, Garber 1977, Hanmer & Blackwood 1982, Negro
et al. 1992, 2000, Hunter 2009, Martinez & Lee 2013, Mikula
et al. 2013) and accipitrids (Eccles et al. 1969, Mumford
1980, Manakadan & Natarajan 1992) were often reported
near the bats roosting places in houses in parts of old
buildings such as church towers, or when bats were released
from captivity. In some cases, raptors were even observed
systematically waiting for emerging bats in the vicinity of
their roosting places and catching them directly at roost
openings (Martinez & Lee 2013, Mikula et al. 2013). Successful attacks on bats further from their roosting places
and in open air were also observed in urban areas (Metcalf
1983, Craves 1994).
Several urban avian predators of bats take advantage of
artificial light sources to extend their activity into nocturnal
times, and sometimes successfully hunt bats attracted to the
insects found near lights (Tryjanowski & Lorek 1998, Negro
et al. 2000, Rejt 2004, DeCandido & Allen 2006). These
findings support the hypothesis that anthropogenic roosting places can act as ecological traps for synurbic species of
bats via their exposure to increased predation pressure
(reviewed by Russo & Ancillotto 2015); however, the overall
impact of predation on bat populations needs to be studied
in greater detail.

Non-raptors as predators

Bats now frequently use man-made buildings and structures

in urban habitats as roosting places (Lausen & Barclay 2006,
Lesinski et al. 2009a). However, roosting in buildings may
expose them to increased predation by opportunistic predators that can reach high numbers in urban areas (reviewed

We found considerably less information on predation of

bats by birds other than raptors. Most predation records
represented hunting attempts by opportunistic omnivores,
e.g. corvids, gulls, hornbills, roadrunners Geococcyx spp., tits
Parus major, and carnivorous shrikes (Laniidae). In most
cases, this behaviour was spontaneous and opportunistic;
attacks usually occurred in the open air and, therefore,
hunting success was rather incidental. Furthermore, the lack
of morphological and behavioural adaptations the predators have for this prey type probably leads to low hunting
success. For instance, non-raptorial bird species usually use
their beaks to catch prey (Lefevre 2005, Collin 2011) instead
of using the claws employed by raptors (Sprunt 1951,
Garber 1977, Byre 1990, Rollie & Christie 2006). In these
occasional bat-hunting species, analysis of their diet and
systematic observations may reveal important insights into
their feeding behaviour. For instance, although information
on predation of bats by ravens Corvus corax from direct
observations is very fragmentary, detailed studies of their
pellets from long-term accumulations confirm the occurrence of eight bat species in their diet (Obuch 2007).
In general, bats comprise only very small proportions of
the diet of diurnal birds, and hunting attempts are only
occasionally successful. An exception to this rule can occur
in local bird populations hunting near roosting places of

Mammal Review (2016) 2016 The Mammal Society and John Wiley & Sons Ltd

URBAN AREAS

Predation of bats by diurnal birds

bats. For instance, aerial attacks on bats by true shrikes

Lanius spp. are very common, but infrequently result in
success (Bent 1950, Young 1980, Gorman 1998). However, it
seems that shrikes may increase their hunting success by
remaining near crevices used by bats as roosts (Herrmann
2004). This hunting technique was observed in several
species of bird that were able to catch bats very successfully
at roost entrances, directly in roosting places or when bats
were unable to escape and their mobility was reduced
(when they fell on the ground, or were hibernating or
injured). For instance, such predatory attempts were
recorded in tits (Estk et al. 2010), corvids (Hochachka &
Scharf 1986, Pacenovsk 2006, Mikula 2013, Michaelsen
et al. 2014), gulls (Holroyd & Beaubien 1983, Collin 2011,
Michaelsen et al. 2014), butcherbirds (Cracticidae; Young
1980), and roadrunners (Herreid 1960, Martnez-Coronel
et al. 2009). In this way, some urban populations of corvids
and gulls can be very effective bat predators (e.g. Forestman
2006, Collin 2011). This suggests that bat populations may
be effectively regulated by bird predators at the local level
and, moreover, points to the vulnerability of urban bat
populations to opportunistic predators. In addition, some
birds, especially corvids and gulls, are able to utilise communal hunting to their advantage to increase their success
at hunting bats (Clay 1959, Cleeves 1969, Lefevre 2005,
Hernndez et al. 2007).

P. Mikula et al.

the presence in tropical ecosystems of avian bat predators,

such as Macheiramphus alcinus and Falco rufigularis, and of
a higher diversity of carnivorous birds such as owls, which
can represent substantial risk of predation for bats during
moonlit nights (del Hoyo et al. 1994, 1999, Kissling et al.
2011). Presence of light avoidance behaviour was, moreover,
experimentally supported in insectivorous and frugivorous
bats (Stone et al. 2009, Lewanzik & Voigt 2014). The nocturnal activity pattern of bats could therefore be permanently
maintained by the constantly present high risk of diurnal
predation by avian predators (Speakman 1995).

CONCLUSIONS
Our results show that attacks on bats by diurnal birds are
globally distributed and reach the geographical limits of the
coexistence of both animal groups. Such predatory attempts
are present in the majority of extant raptor lineages as well
as in numerous other groups of diurnal birds. Moreover,
bats in the majority of extant bat families fall victim to predation by diurnal birds. Our results are thus supportive of
the avian predation hypothesis: predation by diurnal birds
could act as one of major factors affecting the scarcity of
daytime activity in bats, and as a driver of the evolution of
bat nocturnality.

ACKNOWLEDGEMENTS
Support for the avian predation hypothesis
Additional support for the avian predation hypothesis for
the evolution of nocturnality in bats came from observations on bats living on raptor-free islands. Perhaps the bestknown case is that of Nyctalus azoreum, a bat species
endemic to the Azorean archipelago. This species exhibits an
unusually high degree of diurnal activity that appears to
reflect its release from predation risk by diurnal raptors
(Moore 1975, Speakman & Webb 1993). Such an activity
pattern has also been found in some other island bat populations where large avian predators are absent or rare, such
as in the insectivorous bat Hipposideros ruber on So Tom
Island (Russo et al. 2011) and several pteropodids such as
Pteropus melanotus on Christmas Island (Hall et al. 2014)
and Pteropus samoensis on American Samoa (Thomson
et al. 1998). On Fiji, however, where Falco peregrinus is
present (Clunie 1972), the subspecies of Pteropus samoensis
is also diurnal but appears to behave more cryptically
(Wilson & Engbring 1992).
Bats have been observed to shift their emergence time
when diurnal predators are present (e.g. Fenton et al. 1994,
Welbergen 2006). In addition, lunar phobia exists mainly
among tropical bats of all trophic guilds but is almost
absent among temperate insectivorous bats (reviewed by
Lima & OKeefe 2013). Such a pattern could be explained by

We would like to thank Renate Alton, Omar Al-Sheikhly,

Mark D. Anderson, Peter Backor, Colin Beale, Keith Betton,
Rob Bijlsma, Massimo Biondi, BirdWatch Zambia, Brian
Blaylock, Maurice Bot, Mareike Bollen, Mark Brown,
Herve Brule, Center for Wildlife Conservation Cambodia,
Nicoletta Chiozzi, Dominique Conrad, Julie A. Craves,
Stephen Debus, Christiane Denys, Joe DiCostanzo, Pieter
Van Dorsselaer, Pavla Doubkov, Christian Dronneau,
Muriel Dubray, Petra Fahle, Andy Featherstone, Karl
Frafjord, Lenka Glosov, Oscar Gordo, Marco A. M.
Granzinolli, Margaret A. Griffiths, Troy W. Grovenburg,
Ferenc Halsz, Peter Hewitt, Kuan-Chieh Hung, Tom
Jenner, Rudy Jocqu, Ron Johnstone, Holger Kolberg, Margaret Koopman, Alexander Krikellis, Sergei Kruskop, Robinson Lance, Lauraine Leigh, Grzegorz Lesinski, Wen-Horn
Lin, Rebecca Mabuza, Ian Macalpine-Leny, Bojan Marceta,
Brian Marshall, Bruno Massa, tefan Matis, Jean Meguin,
Tore C. Michaelsen, Edita Mikov, Campbell Murn, Renaud
Nadal, Barure Nirmala, Hans Oelke, Han Olff, Jerry Olsen,
Karin Persson, Gary Presland, Hans-Ulrich Raake, Wolfgang
Rackow, Roger Riddington, Chris Risley, Travis Rosenberry,
Martin Sanford, Norbert Schffer, Anja Schuhn, Ron Searle
and his family, Donald Seriff, Dione Seripierri, Clark
Sherman, Christoph Siems-Wedhorn, John R. Speakman,
Gero Speer, Ian M. Spence, Tadeusz Stawarczyk, Toshitaka

10

Mammal Review (2016) 2016 The Mammal Society and John Wiley & Sons Ltd

P. Mikula et al.

N. Suzuki, Craig Symes, Wendy Tatar, Russell Thorstrom,

Stephanie Tyler, Marcel Uhrin, Sam Veansa, Ben Verboom,
Anne Weiserbs, Chip Weseloh, Frank Willems, and Joseph
Wunderle, who kindly sent us copies of some articles or
offered personal observations in this topic. We are also very
thankful to Ivan Horcek, who helped us with the identification of bat taxa from Internet photos.

Predation of bats by diurnal birds

Anonymous (2010) Kasanka Trust Ltd. Kasanka Trust Newsletter

2: 15.
Anonymous (2012) ArcGIS Desktop: release 10.1. Environmental
Systems Research Institute, Redlands, California, USA.
Anonymous (2014a) 6th11th March 2014 The Yucatan
Peninsula, Mexico part 2. Chichen Itza, Felipe Carillo Puerto
and Calakmul. https://gryllosblog.wordpress.com/2014/
05/10/6th-11th-march-2014-the-yucatan-peninsula-mexico
-part-2-chichen-itza-felipe-carillo-puerto-and-calakmul/
(accessed 9 July 2015).
Anonymous (2014b) Back in cyber-space again!!! Report from
Khao Yai! http://orientbirding.com/blog/back-in-cyber
-space-again-report-from-khao-yai/ (accessed 9 July 2015).
Anonymous (2014c) The Yucatan, Mexico. http://www.
birdholidays.co.uk/Yucatan.birdwatching.holiday.htm
(accessed 9 July 2015).
Baker JK (1962) The manner and efficiency of raptor
depredations on bats. The Condor 64: 500504.
Ballance TC (1981) Observations on bat hawk hunting.
Honeyguide 106: 2930.
Beckemeyer R (2000) Phop gan mai iik khrap, Thailand! Argia
13: 912.
Beebe W (1950) Home life of the bat falcon, Falco albigularis
albigularis Daudin. Zoologica 35: 6986.
Begon M, Townsend CR, Harper JL (2005) Ecology: From
Individuals to Ecosystems, 4th ed. Blackwell Publishing,
Oxford, UK.
Bekker JP, Mostert K (1991) Predation on bats in the
Netherlands. Lutra 34: 126.
Bent AC (1950) Life Histories of North American Wagtails,
Shrikes, Vireos, and Their Allies. Dover Publications, New York,
USA.
Binns A, Beer D (2011) Yucatan: Balamku raptors and bat
spectacle. http://notesfromthewildside.com/2011/
03/yucatan-balamku-raptors-and-bat-spectacle/ (accessed 9
July 2015).
Black HL, Howard G, Stjernstedt R (1979) Observations on the
feeding behavior of the bat kite (Macheiramphus alcinus).
Biotropica 11: 1821.
Black HL, Thurow TL, White JD (1987) (Abstract) Bats: an
ecological opportunity and a force in the evolution of the bat
kite. In: Annual Meeting of the Raptor Research Foundation,
Inc., Boise, Idaho, 41. Raptor Research Foundation, Boise,
USA.

Black TH (1956) Bat predators at Carlsbad Caverns. Carlsbad

Caverns Natural History Files, No. N1427.
Boland M (2009) Barred forest falcon at bat cave collared?
https://www.youtube.com/watch?v=wsBHyc-0oVM (accessed
9 July 2015).
Borowski S (1968) Sparrow hawk hunting on the bats. Notatki
Ornitologiczne 9: 4041.
Boxall PC (1982) Further observations of predation by
black-billed magpies on small mammals. Journal of Field
Ornithology 53: 172173.
Brown R (2004) Uganda, 13th26th January 2004. http://www.
surfbirds.com/trip_report.php?id=93 (accessed 10 July 2015).
Byre VJ (1990) A group of young peregrine falcons prey on
migrating bats. The Wilson Bulletin 102: 728730.
Caro T (2005) Antipredator Defenses in Birds and Mammals.
University of Chicago Press, Chicago, USA.
Chapin JP (1932) The birds of the Belgian Congo, Vol. 1.
Bulletin of the American Museum of Natural History 65: 1756.
Clay WM (1959) Blue jays attack a red bat. The Auk 76: 532.
Cleeves TR (1969) Herring gull catching and eating bat. British
Birds 62: 333.
Clunie F (1972) A contribution to the natural history of the Fiji
peregrine. Notornis 19: 302321.
Collin PN (2011) Herring gull catching and eating live bats.
Scottish birds 31: 318319.
Constantine DG (1948) Great bat colonies attract predators.
Bulletin of the National Speleological Society 10: 100.
Craves JA (1994) American kestrel feeds on red bat. Michigan
Birds and Natural History 1: 1415.
de Noronha JD, Battirola LD, Chagas A, de Miranda RM,
Carpanedo RD, Rodrigues DD (2015) Predation of bat
(Molossus molossus: Molossidae) by the centipede Scolopendra
viridicornis (Scolopendridae) in Southern Amazonia. Acta
Amazonica 45: 333336.
del Hoyo J, Elliot A, Sargatal J (1994) Handbook of the Birds of
the World. Vol. 2, New World Vultures to Guineafowl. Lynx
Editions, Barcelona, Spain.
del Hoyo J, Elliot A, Sargatal J (1999) Handbook of the Birds of
the World. Vol. 5, Barn-owls to Hummingbirds. Lynx Edicions,
Barcelona, Spain.
del Hoyo J, Elliott A, Sargatal J, Christie DA, de Juana E (2015)
Handbook of the Birds of the World Alive. Lynx Editions,
Barcelona, Spain. (accessed 19 July 2015;
http://www.hbw.com/).
DeCandido R, Allen D (2006) Nocturnal hunting by peregrine
falcons at the Empire State Building, New York City. The
Wilson Journal of Ornithology 118: 5358.
Dobbs G, Twino-beku A (2004) Uganda, July 28th to August 17th
2004. http://www.birdtours.co.uk/tripreports/uganda/
uganda7/ugan-04.htm (accessed 10 July 2015).
Dreyer NP (2010) Thailand, January 2010. http://dreyerfoto.
dk/index/travel (accessed 19 June 2014).
Drummond AJ, Rambaut A (2007) BEAST: Bayesian
evolutionary analysis by sampling trees. BMC Evolutionary
Biology 7: 214.

Mammal Review (2016) 2016 The Mammal Society and John Wiley & Sons Ltd

11

REFERENCES

Predation of bats by diurnal birds

P. Mikula et al.

Duquet M, Nadal R (2012) La capture de chauves-souris par des

rapaces diurnes en France: essai de synthse. Ornithos 19:
184195.
Eccles DH, Jensen RAC, Jensen MK (1969) Feeding behaviour of
the bat hawk. Ostrich 40: 2627.
Esbrard CE, Vrcibradic D (2007) Snakes preying on bats: new
records from Brazil and a review of recorded cases in the
Neotropical Region. Revista Brasileira de Zoologia 24:
848853.
Estk P, Zsebok S, Siemers BM (2010) Great tits search for,
capture, kill and eat hibernating bats. Biology Letters 6: 5962.
Fam SD, Nijman V (2011) Spizaetus hawk-eagles as predators of
arboreal colobines. Primates 52: 105110.
Farina O, de Carvalho C, Pedro WA (2011) Predation of
Platyrrhinus lineatus (E. Geoffroy, 1810) (Chiroptera:
Phyllostomidae) by Cyanocorax chrysops (Vieillot, 1818)
(Passeriformes: Corvidae). Chiroptera Neotropical 17:
993996.
Fenton MB, Cumming DHM, Oxley DJ (1977) Prey of bat
hawks and availability of bats. The Condor 79: 495497.
Fenton MB, Rautenbach IL, Smith SE, Swanepoel CM, Grosell J,
Van Jaarsveld J (1994) Raptors and bats: threats and
opportunities. Animal Behaviour 48: 918.
Ferguson-Lees J, Christie DA (2001) Raptors of the World. Helm
Identification Guides, London, UK.
Finch BW (1978) Observations and notes on the bat hawk
Machaeramphus alcinus in Papua. Papua New Guinea Bird
Society Newsletter 146: 812.
Forestman (2006) Urraca cazando murcilagos. https://www.
youtube.com/watch?v=E881fLF-EJ0 (accessed 18 August
2015).
Foysal M (2015) Observations of red-headed falcon Falco
chicquera (Aves: Falconiformes: Falconidae) nest at
Keraniganj, Dhaka, Bangladesh, with a focus on post-fledging
behavior. Journal of Threatened Taxa 7: 71387145.
Frafjord K (2012) Observations of a merlin (Falco columbarius)
hunting northern bats (Eptesicus nilssonii) in midnight sun
(Northern Norway). Nyctalus 17: 107111.
Fry CH (1964) Red-necked falcon Falco chicquera hunting bats.
Bulletin of the Nigerian Ornithologists Society 1: 19.
Garber SD (1977) Bat predation by the American kestrel, Falco
sparverius (Aves: Falconiformes). Bat Research News 18:
3738.
Gorman G (1998) Great grey shrike Lanius excubitor hunting
noctule bats. Aquila 103104: 153.
Gunnell GF, Simmons NB (2012) Evolutionary History of Bats
Fossils, Molecules and Morphology. Cambridge University
Press, Cambridge, UK.
Haensel J, Smmer P (2002) Birds of prey catch bats and flying
foxes an attempt of a general survey most recent
knowledge about bat hunting of the fastest falcons in
Germany. Ornithologische Jahresberichte des Museums
Heineanum 20: 99141.
Hall J, Rose K, Smith C, De Jong C, Phalen D, Austen J, Field H
(2014) Health assessment of the Christmas Island flying fox

(Pteropus melanotus natalis). Journal of Wildlife Diseases 50:

447458.
Hall MI, Iwaniuk AN, Gutirrez-Ibez C (2009) Optic foramen
morphology and activity pattern in birds. The Anatomical
Record 292: 18271845.
Hanmer DB, Blackwood JGV (1982) A bat-eating kestrel, Falco
dickinsoni. Ostrich 53: 188189.
Harden WD (1972) Predation by hawks on bats at Vickery
Bat Cave. Bulletin of the Oklahoma Ornithological Society 5:
45.
Hernndez DL, Mell JJ, Eaton MD (2007) Aerial predation of a
bat by an American crow. The Wilson Journal of Ornithology
119: 763764.
Herreid CF (1960) Roadrunner a predator of bats. The Condor
62: 67.
Herrmann E (2004) Common fiscal Lanius collaris preying on a
cape serotine bat Eptesicus capensis. Ostrich 75: 333.
Hochachka WM, Scharf CS (1986) Black-billed magpie, Pica
pica, predation on bats. The Canadian Field-naturalist 100:
121122.
Holroyd GL, Beaubien EG (1983) Ring-billed gull, Larus
delawarensis, predation on bat, Myotis, injured by an
American kestrel, Falco sparverius. The Canadian
Field-naturalist 97: 452.
Hunter S (2009) Rock kestrel hunting bats at roost. Gabar 20:
40.
Jacobs IF, Osvath M, Osvath H, Mioduszewska B, von Bayern
AM, Kacelnik A (2014) Object caching in corvids:
incidence and significance. Behavioural Processes 102:
2532.
Jetz W, Thomas GH, Joy JB, Hartmann K, Mooers AO (2012)
The global diversity of birds in space and time. Nature 491:
444448.
Jedrzejewska B, Jedrzejewski W (1998) Predation in Vertebrate
Communities: the Biaowieza Primeval Forest as a Case Study.
Springer-Verlag, Berlin, Germany.
Jones LR, Black HL, White CM (2012) Evidence for convergent
evolution in gape morphology of the bat hawk
(Macheiramphus alcinus) with swifts, swallows, and
goatsuckers. Biotropica 44: 386393.
Kemp AC (2001) Family Bucerotidae (hornbills). In: del Hoyo J,
Elliott A, Sargatal J (eds) Handbook of the Birds of the World,
Vol. 6, Mousebirds to Hornbills, 436523. Lynx Editions,
Barcelona, Spain.
Kemp AC, Rautenbach IL (1987) Bat hawks or bat-eating hawks?
Gabar 2: 46.
Kennedy RS (1985) Conservation research of the Philippine
Eagle. National Geographic Society Research Report 18:
401414.
Kervyn T (1998) Attacks of serotine bats (Eptesicus serotinus) by
breeding starlings (Sturnus vulgaris) and common swifts
(Apus apus). Aves 35: 225229.
Kirven MN (1976) The Ecology and Behavior of the Bat Falcon,
Falco rufigularis. PhD thesis, University of Colorado, Boulder,
Colorado, USA.

12

Mammal Review (2016) 2016 The Mammal Society and John Wiley & Sons Ltd

P. Mikula et al.

Predation of bats by diurnal birds

Kissling WD, Sekercioglu CH, Jetz W (2011) Bird dietary guild

richness across latitudes, environments and biogeographic
regions. Global Ecology and Biogeography 21: 328340.
Kunz TH, Fenton MB (2003) Bat Ecology. University of Chicago
Press, Chicago, USA.
Lausen CL, Barclay RMR (2006) Benefits of living in a building:
big brown bats (Eptesicus fuscus) in rocks versus buildings.
Journal of Mammalogy 87: 362370.
Laycock P (1982) Avian predation on cave dwelling
insectivorous bats. Bokmakierie 34: 1718.
Lee YF, Kuo YM (2001) Predation on Mexican free-tailed bats
by peregrine falcons and red-tailed hawks. Journal of Raptor
Research 35: 115123.
Lefevre KL (2005) Predation of a bat by American crows, Corvus
brachyrhynchos. The Canadian Field-naturalist 119: 443444.
Lendrum AL (1977) Observations on a bat hawk. Honeyguide
90: 45.
Lesinski G, Gryz J, Kowalski M (2009a) Bat predation by tawny
owls Strix aluco in differently human-transformed habitats.
Italian Journal of Zoology 76: 415421.
Lesinski G, Ignaczak M, Manias J (2009b) Opportunistic
predation on bats by the tawny owl Strix aluco. Animal
Biology 59: 283288.
Lewanzik D, Voigt CC (2014) Artificial light puts ecosystem
services of frugivorous bats at risk. Journal of Applied Ecology
51: 388394.
Lewis MJ (1987) Australian kestrels Falco cenchroides feeding on
bats. Australian Bird Watcher 12: 126127.
Lim CK, Cranbrook E (2002) Swiftlets of Borneo: Builders of
Edible Nests. Natural History Publications (Borneo), Kota
Kinabalu, Malaysia.
Lima SL, Dill LM (1990) Behavioral decisions made under the
risk of predation: a review and prospectus. Canadian Journal
of Zoology 68: 619640.
Lima SL, OKeefe JM (2013) Do predators influence the
behaviour of bats? Biological Reviews 88: 626644.
Lindskog SR (1976) Sparrow hawk Accipiter nisus catching a bat.
Anser 14: 281.
Looney MW (1972) Predation of bats by hawks and owls.
Bulletin of the Oklahoma Ornithological Society 5: 14.
Manakadan R, Natarajan V (1992) Brahminy kite Haliastur
indus (Boddaert) preying on bats. Journal of the Bombay
Natural History Society 89: 367.
Martnez-Coronel M, Morales-Medina X, Mdespacher-Ziehl C
(2009) Depredadores de murcilagos en la cueva de Los
Laguitos, Chiapas, Mxico. Revista Mexicana De Mastozoologa
(Nueva poca) 13: 8291.
Martinez SG, Lee TE (2013) Predation on Mexican free-tailed
bats (Tadarida brasiliensis) by merlin (Falco columbarius). The
Southwestern Naturalist 58: 508512.
Mason CF, Stebbings RE, Winn GP (1972) Noctules (Nyctalus
noctula) and starlings (Sturnus vulgaris) competing for
roosting holes. Journal of Zoology 166: 467.
Meinesz A, Meinesz S, Bennett G (1982) Rock kestrel takes a bat.
Bokmakierie 34: 4041.

Menchetti M, Scalera R, Mori E (2014) First record of a possibly

overlooked impact by alien parrots on a bat (Nyctalus leisleri).
Hystrix, the Italian Journal of Mammalogy 25: 6162.
Metcalf EC (1983) Australian hobbies hunting bats. Australian
Raptors Association News 4: 78.
Michaelsen TC, Jensen KH, Hgstedt G (2014) Roost site
selection in pregnant and lactating soprano pipistrelles
(Pipistrellus pygmaeus Leach, 1825) at the species northern
extreme: the importance of warm and safe roosts. Acta
Chiropterologica 16: 349357.
Michalak L (1997) Sharp-shinned hawk preys on bat. Ontario
Birds 15: 2728.
Mikula P (2013) Western jackdaw (Corvus monedula) attacking
bats (Chiroptera): observations from Bardejov, northeastern
Slovakia. Sylvia 49: 157159.
Mikula P, Hromada M, Tryjanowski P (2013) Bats and swifts as
food of the European kestrel (Falco tinnunculus) in a small
town in Slovakia. Ornis Fennica 90: 178185.
Molinari J, Gutirrez EE, Asceno AA, Nassar JM, Arends A,
Mrquez RJ (2005) Predation by giant centipedes, Scolopendra
gigantea, on three species of bats in a Venezuelan cave.
Caribbean Journal of Science 41: 340346.
Moore NW (1975) The diurnal flight of the Azorean bat and
the avifauna of the Azores. Journal of Zoology 117:
583586.
Mumford RE (1980) Two cases of hawk predation on bats. Bat
Research News 21: 11.
Negro JJ, Ibez C, Prezjord JL, Delariva MJ (1992) Winter
predation by common kestrel Falco tinnunculus on pipistrelle
bats Pipistrellus pipistrellus in southern Spain. Bird Study 39:
195199.
Negro JJ, Bustamante J, Melguizo C, Ruiz JL, Grande JM (2000)
Nocturnal activity of lesser kestrels under artificial lighting
conditions in Seville, Spain. Journal of Raptor Research 34:
327329.
Nyffeler M, Knrnschild M (2013) Bat predation by spiders.
PLoS ONE 8: e58120.
Obuch J (1998) The representation of bats (Chiroptera) in the
diet of owls (Strigiformes) in Slovakia. Vespertilio 3:
6574.
Obuch J (2007) Food of the raven (Corvus corax) in Slovakia.
Tichodroma 19: 110.
Ojeda VS, Chazarreta ML (2006) Provisioning of Magellanic
woodpecker (Campephilus magellanicus) nestlings with
vertebrate prey. The Wilson Journal of Ornithology 118:
251254.
Olsen PD, Debus SJS, Czechura GV, Mooney NJ (1990)
Comparative feeding ecology of the grey goshawk Accipiter
novaehollandiae and brown goshawk Accipiter fasciatus.
Australian Bird Watcher 13: 178192.
Pacenovsk S (2006) Attack of the black-billed magpie (Pica
pica) on the noctule bat (Nyctalus noctula). Vespertilio 910:
231232.
Pagnoni G (2013) Halconcito colorado (Falco sparverius)
cazando murcilagos. Nuestras Aves 58: 5960.

Mammal Review (2016) 2016 The Mammal Society and John Wiley & Sons Ltd

13

Predation of bats by diurnal birds

P. Mikula et al.

Parker M (1997) Ecology of Nesting Laughing Falcons and Bat

Falcons in Tikal National Park, Guatemala: Foraging and Niche
Breadth. MSc thesis, Boise State University, Boise, USA.
Pierson JE, Donahue P (1983) Peregrine falcon feeding on bats
in Suriname, South America. American Birds 37: 257259.
Pikacha P, Sirikolo M, Boseto D, Filardi C (2012) Ecological
observations on Sanfords sea-eagle Haliaeetus (leucogaster)
sanfordi. Australian Field Ornithology 29: 143148.
Rautenbach IL, Fenton MB, Kemp AC, Van Jaarsveld SJ (1990)
Home range and activity of African goshawks Accipiter tachiro
in relation to their predation on bats. Koedoe 33: 1721.
Rejt L (2004) Nocturnal feeding of young by urban peregrine
falcons Falco peregrinus in Warsaw (Poland). Polish Journal of
Ecology 52: 6368.
Robinson SK (1994) Habitat selection and foraging ecology of
raptors in Amazonian Peru. Biotropica 26: 443458.
Rodrguez-Durn A, Lewis AR (1985) Seasonal predation by
merlins on sooty mustached bats in western Puerto Rico.
Biotropica 17: 7174.
Rollie C, Christie G (2006) Peregrine falcon catching and killing
a bat in daylight. Scottish Birds 26: 4546.
Round PD, Jukmongkol R, Kasorndorkbua C (2006) Recent
reports MarchJune 2006. http://www.worldtwitch.com/2006
_thailand_bird_reports.htm (accessed 9 July 2015).
Roworth P, Wright E (1989) Sparrowhawk attacking noctule
bats. British Birds 82: 564565.
Russo D, Ancillotto L (2015) Sensitivity of bats to urbanization:
a review. Mammalian Biology 80: 205212.
Russo D, Maglio G, Rainho A, Meyer CF, Palmeirim JM (2011)
Out of the dark: diurnal activity in the bat Hipposideros ruber
on So Tom Island (West Africa). Mammalian Biology 76:
701708.
Rydell J (1992) Occurrence of bats in northernmost Sweden
(65N) and their feeding ecology in summer. Journal of
Zoology 227: 517529.
Rydell J, Speakman JR (1995) Evolution of nocturnality in bats:
potential competitors and predators during their early history.
Biological Journal of the Linnean Society 54: 183191.
Rydell J, Entwistle A, Racey PA (1996) Timing of foraging flights
of three species of bats in relation to insect activity and
predation risk. Oikos 76: 243252.
Sarkozi DL, Brooks DM (2003) Eastern red bat (Lasiurus
borealis) impaled by a loggerhead shrike (Lanius
ludovicianus). Southwestern Naturalist 48: 301303.
Schwan TG, Hikes N (1979) Fiscal shrike predation on the bat
Pipistrellus kuhli in Kenya. Biotropica 11: 21.
Seijas AE (1996) Feeding of the bat falcon (Falco rufigularis) in
an urban environment. Journal of Raptor Research 30: 3335.
Sheldon FH, Moyle RG, Kennard J (2001) Ornithology of Sabah:
history, gazetteer, annotated checklist, and bibliography.
Ornithological Monographs 52: 1285.
Smythies BE (1953) The Birds of Burma. Oliver & Boyd,
Edinburgh, UK.
Smythies BE (1968) The Birds of Borneo. 2nd ed. Oliver & Boyd,
Edinburgh, UK.

Sparks DW, Roberts KJ, Jones C (2000) Vertebrate predators on

bats in North America North of Mexico. Reflections of a
Naturalist. In: Choate JR (ed) Papers Honoring Professor
Eugene D. Fleharty, 229241. Fort Hays Studies, Hays, Kansas,
USA.
Speakman JR (1990) The function of daylight flying in British
bats. Journal of Zoology London 220: 101113.
Speakman JR (1991a) The impact of predation by birds on bat
populations in the British Isles. Mammal Review 21: 123142.
Speakman JR (1991b) Why do insectivorous bats in Britain not
fly in daylight more frequently? Functional Ecology 5:
518524.
Speakman JR (1995) Chiropteran nocturnality. Symposia of the
Zoological Society of London 67: 187201.
Speakman JR, Webb PI (1993) Taxonomy, status and
distribution of the Azorean bat (Nyctalus azoreum). Journal of
Zoology 231: 2738.
Speakman JR, Lumsden LF, Hays GC (1994) Predation rates on
bats released to fly during daylight in south-eastern Australia.
Journal of Zoology 233: 318321.
Sprunt A (1950) Hawk predation at the bat caves of Texas. Texas
Journal of Science 2: 463470.
Sprunt A (1951) Aerial feeding of duck hawk, Falco p. anatum.
The Auk 68: 372373.
Stager KE (1941) A group of bat-eating duck hawks. The Condor
43: 137139.
Stimpson CM (2009) Raptor and owl bone from Niah Caves,
Sarawak: identifications and morphological variation in the
humerus and tarsometatarsus of selected raptors.
International Journal of Osteoarchaeology 19: 476490.
Stone EL, Jones G, Harris S (2009) Street lighting disturbs
commuting bats. Current Biology 19: 11231127.
Stuart C, Stuart T (2004) African white-backed vultures Gyps
africanus feeding on straw-coloured fruit bats Eidolon helvum.
Vulture News 50: 3435.
Suzuki TN (2012) Mobbing to death of a Japanese long-eared
bat Plecotus sacrimontis by two species of tit. Forktail 28:
171172.
Thiollay JM (1983) Evolution actuelle du peuplement de rapaces
diurnes dans le nord de Borno. Alauda 51: 109123.
Thomsett S (1987) Bat hunting by lanner falcons in Kenya.
Gabar 2: 78.
Thomson SC, Brooke AP, Speakman JR (1998) Diurnal activity
in the Samoan flying fox, Pteropus samoensis. Philosophical
Transactions of the Royal Society B: Biological Sciences 353:
15951606.
Tryjanowski P, Lorek G (1998) Kestrel and great grey shrike
catch insects after sunset in the dark, lightened by an artificial
light. British Birds 91: 327.
Tugendhat M (1966) Swallows mobbing pipistrelle bat. British
Birds 59: 435.
Twente JW (1954) Predation on bats by hawks and owls. The
Wilson Bulletin 66: 135136.
Twente JW (1956) Ecological observations on a colony of
Tadarida mexicana. Journal of Mammalogy 37: 4247.

14

Mammal Review (2016) 2016 The Mammal Society and John Wiley & Sons Ltd

P. Mikula et al.

van Beirs M (2004) The best of Uganda, 28 November13

December 2004. http://www.birdquest-tours.com/
pdfs/report/UGANDA%20(BEST)%20REP%2004.pdf
(accessed 10 July 2015).
van Beirs M (2014) The Yucatan Peninsula, 28 February11
March 2014. http://www.birdquest-tours.com/pdfs/report/
MEXICO%20-YUCATAN-%20REP%2014-ebook.pdf
(accessed 9 July 2015).
Van Jaarsveld J (1988) African goshawks and European hobbies
bat-hawking. Gabar 3: 2931.
Welbergen JA (2006) Timing of the evening emergence from day
roosts of the grey-headed flying fox, Pteropus poliocephalus:
the effects of predation risk, foraging needs, and social
context. Behavioral Ecology and Sociobiology 60: 311322.
Wilson DE, Engbring J (1992) The flying foxes Pteropus
samoensis and Pteropus tonganus: status in Fiji and Samoa. In:
Wilson DE, Graham GL (eds) Pacific Island Flying Foxes:
Proceedings of an International Conference, 74101. US Fish
and Wildlife Service Biological Report, Washington, USA.
Woolley S, Casson J (2008) Murchison Falls.
http://jjcskw.com/trip%20reports/uganda/murchison.htm
(accessed at 10 July 2015).
Worthy TH, Anderson A (2009) Results of palaeofaunal
research. In: Clark G, Anderson A (eds) The Prehistory of
Early Fiji, Terra Australis 31, 4162. Australian National
University, Canberra, Australia.
Worthy TH, Holdaway RN (1995) Quaternary fossil faunas from
caves on Mt Cookson, North Canterbury, South Island, New
Zealand. Journal of the Royal Society of New Zealand 25:
333370.

Mammal Review (2016) 2016 The Mammal Society and John Wiley & Sons Ltd

Predation of bats by diurnal birds

Yancey FD, Roberts KJ, Jones C (1996) Prairie falcon predation

on Brazilian free-tailed bats. Prairie Naturalist 28: 146.
Yosef R (1991) Foraging habits, hunting and breeding success of
lanner falcons in Israel. Journal of Raptor Research 25: 7781.
Young RA (1980) Observations on the vulnerability of two
species of wattled bats Chalinolobus to diurnal avian
predators. Victorian Naturalist 97: 258263.

SUPPORTING INFORMATION
Additional supporting information may be found in the
online version of this article at the publishers web-site.
Appendix S1. Index of all languages used in literature
searching.
Appendix S2. Summary table of diurnal raptors
(Accipitriformes and Falconiformes) reported to capture
bats with list of localities at country level and bat taxa
(arranged in alphabetical order).
Appendix S3. Summary table of non-raptorial diurnal
birds reported as bat hunters with list of localities at
country level and bat taxa (arranged in alphabetical order).
Appendix S4. References used in the Appendices.

15