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Abstract

We reviewed published and unpublished written reports on bat mortality at wind farms in northwestern Europe. The estimated number of bats killed per turbine annually was relatively low (0-3) on flat, open farmland away from the coast, higher (2-5) in more complex agricultural landscapes, and highest (5-20) at the coast and on forested hills and ridges. The species killed almost exclusively (98%) belonged to a group (Nyctalus, Pipistrellus, Vespertilio and Eptesicus spp.) adapted for open-air foraging. The bats were killed by the moving rotor blades as they hunted insects attracted to the turbines. This occurred independently of sex and age. Peak mortality varied considerably in frequency and timing among years, but the events usually (90%) occurred on nights with low wind speeds in late July to early October and to a lesser extent (10%) also in April-June. The mortality increased with turbine tower height and rotor diameter but was independent of the distance from the ground to the lowest rotor point. It was also independent of the size of the wind park (118 turbines). Bat species other than the open-air suite referred to above are usually not at risk at wind turbines, because they fly below the rotors, but are still killed occasionally (2%).
INTRODUCTION
Wind turbines cause extensive mortality among
birds and bats in certain situations, more so in some
species than in others. This has become an important
ethical problem and has resulted in serious concern
about the future of some populations (Kunz et al.,
2007a). For example, bats that encounter wind tur-
bines along the Appalachian Mountains in eastern
U.S. are killed at rates of 30–40 individuals per tur-
bine annually. The estimated total mortality rate at
wind turbines in this area alone in the year 2020,
based on projections of installed capacity, is be-
tween 33,000 and 110,000 bats per year (Kunz et al.,
2007a). Bats typically live long lives and have slow
reproductive rates (Barclay and Harder, 2003) and,
although the sizes of bat populations are generally
unknown, it is unclear if they can sustain increased
mortality of this magnitude (Kunz et al., 2007a).
Based on the North American studies reviewed
by Kunz et al. (2007a, 2007b) and Arnett et al.
(2008) and the European work available to us, it ap-
pears that death rates similar to that of the Appalach -
ians are uncommon. Nevertheless, mortality figures
nearly as high have been reported at wind turbines
on top of forested hills and ridges in southern Ger -
many (Brinkmann et al., 2006) and at the At lan tic
coast of France (Dulac, 2008). These cases will be
considered further below.
The purpose of this review is to summarize the
present knowledge about wind turbine effects on
bats, as it has developed in northwestern Europe
since 1999 (Bach et al., 1999; Ramel et al., 1999).
Our aims are: i) to synthesize current information
from bat mortality monitoring in northwestern
Europe; ii) to apply statistical analysis to data pre-
sented in the original reports, using a meta-analyt-
ical approach, to test some general hypotheses;
and iii) to compare the results of our analysis with
data from North America (reviewed by Kunz et al.,
2007a, 2007b and Arnett et al., 2008). The overall
aim is to provide an up-to-date summary of the bat
Acta Chiropterologica, 12(2): 261–274, 2010
PL ISSN 1508-1109 © Museum and Institute of Zoology PAS
doi: 10.3161/150811010X537846
Bat mortality at wind turbines in northwestern Europe
JENS RYDELL
1, 5
, LOTHAR BACH
2
, MARIE-JO DUBOURG-SAVAGE
3
, MARTIN GREEN
1
, LUISA RODRIGUES
4
,
and A
NDERS HEDENSTRÖM
1
1
Ecology Building, Lund University, SE-223 62 Lund, Sweden
2
Freilandforschung, zool. Gutachten, Hamfhofsweg 125b, D-28357 Bremen, Germany
3
Société pour l’Etude et la Protection des Mammifères, c/o Muséum d’Histoire Naturelle de Bourges, Les Rives d’Auron,
F-18000 Bourges, France
4
Instituto da Conservação da Natureza e da Biodiversidade, Rua de Santa Marta 55, P-1150-294 Lisboa, Portugal
5
Corresponding author: E-mail: jens.rydell@telia.com
We reviewed published and unpublished written reports on bat mortality at wind farms in northwestern Europe. The estimated
number of bats killed per turbine annually was relatively low (0–3) on flat, open farmland away from the coast, higher (2–5) in more
complex agricultural landscapes, and highest (5–20) at the coast and on forested hills and ridges. The species killed almost
exclusively (98%) belonged to a group (Nyctalus, Pipistrellus, Vespertilio and Eptesicus spp.) adapted for open-air foraging. The
bats were killed by the moving rotor blades as they hunted insects attracted to the turbines. This occurred independently of sex and
age. Peak mortality varied considerably in frequency and timing among years, but the events usually (90%) occurred on nights with
low wind speeds in late July to early October and to a lesser extent (10%) also in April-June. The mortality increased with turbine
tower height and rotor diameter but was independent of the distance from the ground to the lowest rotor point. It was also
independent of the size of the wind park (1–18 turbines). Bat species other than the open-air suite referred to above are usually not
at risk at wind turbines, because they fly below the rotors, but are still killed occasionally (2%).
Key words: aerial ecology, aeroecology, bat conservation, high-altitude feeding, killing factors, renewable energy, wind farming
INTRODUCTION
The Cerrado is the most extensive woodland-
savanna in South America and one of the world’s
34 biodiversity hotspots (Myers et al., 2000;
Mit ter meier et al., 2004). The rapid expansion of
agriculture and cattle ranching severely affected
the biome and reduced it to 21.6% of its original
area (Mit termeier et al., 2004). A quarter of all
grain pro duced in Brazil and nearly 40 million
heads of cattle represent important economic
activities that impact the biome, with steady growth
projected in both industries, as well as in charcoal
production. Low levels of investment in biodi-
versity research, and a small protected areas net-
work (both in number of protected areas and in
total area under protection) only make the Cerrado
conservation more urgent (Klink and Machado,
2005).
Bats compose more than 50% of the mammal
fauna in the Cerrado (Aguiar and Zortéa, 2008), and
they play key roles in ecosystem processes and serv-
ices, as pollinators of native plants (e.g., Martins and
Batalha, 2006), as seed dispersers that help to re-
store degraded areas (e.g., Bizerril and Raw, 1998),
and as predators that help to regulate and control
agriculture pests (e.g., Aguiar and Antonini, 2008).
Bats are also the mammals most commonly in-
volved in transmission of rabies to humans (Schnei -
der et al., 1996; Belotto et al., 2005). In addition to
rabies-related public health problems, disease trans-
mission by vampire bats to herbivorous species,
chiefly cattle, is also a major constraint to livestock
production in Latin America (Belotto et al., 2005).
Acta Chiropterologica, 12(2): 275–282, 2010
PL ISSN 1508-1109 © Museum and Institute of Zoology PAS
doi: 10.3161/150811010X537855
Do current vampire bat (Desmodus rotundus) population control practices pose
a threat to Dekeyser’s nectar bat’s (Lonchophylla dekeyseri) long-term persistence
in the Cerrado?
LUDMILLA M. S. AGUIAR
1
, DANIEL BRITO
2, 3, 4
, and RICARDO B. MACHADO
1
1
Universidade de Brasília, Instituto de Biologia, Departamento de Zoologia, Campus Universitário Darcy Ribeiro, Brasília,
Distrito Federal 70910-900, Brazil
2
Universidade Federal de Goiás, Instituto de Ciências Biológicas, Departamento de Ecologia, Caixa Postal 131, Goiânia,
Goiás 74001-970, Brazil
3
Universidade Estadual de Santa Cruz, Programa de Pós-Graduação em Ecologia e Conservação da Biodiversidade, Rodovia
Ilhéus-Itabuna km 16, Ilhéus, Bahia 45662-000, Brazil
4
Corresponding author: brito.dan@gmail.com
The Cerrado is rapidly losing space to agriculture, pastures and urbanization. Current management practices to control rabies
outbreaks through the eradication of vampire bat populations may put other bat species in peril. Our objective is to evaluate if the
current vampire bat population control practices could pose a threat to Lonchophylla dekeyseris persistence, an endemic bat of the
Cerrado. We used the VORTEX program to model different vampire bat management scenarios, causing low (25%), medium (50%)
or high (75%) incidental mortality to L. dekeyseri populations. Inbreeding depression has been identified as a threat to the species,
therefore we also modeled scenarios evaluating such effects. Results show that current vampire bat management practices have
serious impacts on populations of L. dekeyseri. In all cases marked declines in population sizes were observed (even when there was
no decline in survival probabilities). For medium and high incidental mortality management scenarios, we also observed decreases
in survival probability and in genetic diversity. In those scenarios evaluating vampire bat management and inbreeding depression
together, the models suggest that such interaction results in more pronounced declines. Habitat loss and fragmentation in the Cerrado
are severe threats and have already negatively impacted L. dekeyseri. Unfortunately, if currentpopulation control practices dealing
with vampire bats are not changed, inappropriate rabies management may be the coup de grace to the long-term persistence of this
species.
Key words: Cerrado, Desmodus, Lonchophylla, population viability analysis, rabies, roost
INTRODUCTION
Mammalian reproduction is energetically expen-
sive, and lactation is the most expensive component
for females (Racey and Speakman, 1987; Gittleman
and Thompson, 1988). Lactating females must meet
the energetic demands of lactation during nightly
feeding bouts (Racey and Speakman, 1987; Kurta
et al., 1989, 1990; McLean and Speakman, 1997),
while at the same time balancing time spent away
from their roost with the necessity of regularly re-
turning to feed dependent offspring. While males,
non-reproductive females, and pregnant or post-
lactating females often spend extended periods of
time in roosts between foraging bouts (Kunz, 1973;
Fenton et al., 1977; Anthony et al., 1981; Ruedi,
1993), lactating females typically spend relatively
more time foraging and less time in roosts (Bar-
clay, 1989; Aldridge and Brigham, 1991; Clark et
al., 1993; Catto et al., 1995; Entwistle et al., 1996;
Wilkinson and Barclay, 1997; Shiel et al., 1999;
O’Donnell, 2002; but see Dietz and Kalko, 2007).
Furthermore, lactating females reduce the size of
their foraging area and make repeated visits to their
day roost to nurse pups and provide additional pa -
rental care (Anthony et al., 1981; Audet, 1990; Shiel
et al., 1999; O’Donnell 2001; Henry et al., 2002;
Murray and Kurta, 2004). As lactation progresses,
lactating females typically make fewer but longer
visits to the maternity roost, presumably to spend
longer periods feeding larger offspring (Barclay,
1989). However, the total time spent in the materni-
ty roost by lactating females remains fairly constant
throughout lactation (Clark et al., 1993; Rydell,
1993; Entwistle et al., 1996). Thus, the increased
energetic demands of lactation and the needs of
dependent offspring are reflected in changes in the
nocturnal activity patterns of lactating females.
Females of most temperate bat species form ma-
ternity colonies during pregnancy and lactation, and
Acta Chiropterologica, 12(2): 283–291, 2010
PL ISSN 1508-1109 © Museum and Institute of Zoology PAS
doi: 10.3161/150811010X537864
Nocturnal activity patterns of lactating silver-haired bats (Lasionycteris noctivagans):
the influence of roost-switching behavior
MAARTEN J. VONHOF
1, 3
and BURR J. BETTS
2
1
Department of Biological Sciences and Environmental Studies Program, Western Michigan University, Kalamazoo,
MI 49008-5410, USA
2
Eastern Oregon University, One University Blvd., La Grande, OR 97850-2899, USA
3
Corresponding author: E-mail: maarten.vonhof@wmich.edu
A characteristic feature of forest-dwelling bats is that they regularly switch day roosts. The additional time associated with finding
or assessing the availability of a new roost, moving offspring to the new roost, coordinating with other group members, and meeting
any increased energetic demands that may be associated with these behaviors may influence the number and duration of feeding
bouts of reproductive females. To examine the influence of roost-switching on nocturnal activity patterns we radio-tagged lactating
female silver-haired bats (Lasionycteris noctivagans) and monitored them on nights when they moved to a new roost site and nights
when no roost shift occurred. The typical activity pattern on a non-move night consisted of an initial long bout away from the day
roost (median length: 177.4 min) followed by alternating bouts in (50.6 min) and away (57.0 min) from the roost for the rest of the
night. On move nights bats reduced the length of their first absent period (82.7 min) and increased the number of trips from the roost,
although neither the length of these subsequent absent periods nor the total time absent from the roosts differed from non-move
nights. Similarly, total time spent in roosts did not differ between move and non-move nights, but bats made more and shorter visits
on move nights. Visits to original and new roosts were not synchronous among colony members, and the length of time for
individuals from the same colony to move to a new roost varied from 1 to 4 h. Our results indicate that roost switching did not
influence the total time spent in or away from day roosts. Rather, bats moved between the original roost, the new roost, and foraging
areas with much greater frequency on move nights.
Key words: activity patterns, lactation, Lasionycteris noctivagans, roost switching, silver-haired bat, time budget, tree-roosting
INTRODUCTION
Climatic extremes and unpredictable precipita-
tion are characteristics of desert climates and gener-
ally impose restrictions on the energy and water
intakes of desert dwelling organisms (Noy-Meir,
1973). Thus, energy-expensive activities, such as
flight, social interactions and echolocation (although
not in flight) that are a routine for bats, could make
these animals as a group, and small insectivorous
bats in particular, vulnerable to even slight vagaries
of their environment. Bats may be additionally vul-
nerable because of their high surface to volume ra-
tio that is more than twice that of similar sized, non-
flying mammals due to their naked, vascularized
wings. Foraging activity in insectivorous bats is
affected by variation in prey availability that may
change daily as a function of temperature, wind and
light (Altringham, 1999). Furthermore, insectivo-
rous bat activity may be limited in areas with low
winter temperatures, due to the paucity of available
prey. This phenomenon was described for temperate
regions in general, where, during winter, only flying
insects are active, and it most strongly affects the ac-
tivity of bats that glean arthropods from ground,
rock and plant surfaces (McNab, 1982). Indeed,
Ayal and Merkl (1994) found a significant declining
trend in terrestrial beetle activity from the beginning
to the end of winter in the Negev desert. This lack
of available prey during winter may constrain the
activity of insectivorous bats in general, and the
timing of reproduction in particular, and may force
mother bats to invest in particularly fast growth of
embryos and pups.
Female bats are exposed to additional environ-
mental stress because of the costs of maternal in-
vestment. The food and water requirements of fe-
male bats increase substantially during pregnancy
and even more so during lactation/nursing (Speak -
man and Racey, 1987; Kurta et al., 1989; Kunz et
Acta Chiropterologica, 12(2): 293–299, 2010
PL ISSN 1508-1109 © Museum and Institute of Zoology PAS
doi: 10.3161/150811010X537873
Foraging behavior of a desert dwelling arthropod-gleaning bat
(Otonycteris hemprichii) during pregnancy and nursing
SHAI DANIEL
1, 2
, CARMI KORINE
2, 3, 4
, and BERRY PINSHOW
1, 2
1
Department of Life Sciences, Ben-Gurion University of the Negev, P.O.B. 653, 84105 Beer Sheva, Israel
2
Mitrani Department of Desert Ecology, Jacob Blaustein Institutes for Desert Research, Ben-Gurion University of the Negev,
84990 Midreshet Ben-Gurion, Israel
3
Present address: Ramon Science Center, P.O.B. 194, 80600 Mitzpe Ramon, Israel
4
Corresponding author: E-mail: ckorine@bgu.ac.il
We studied the role of behavioral activities used by desert-dwelling, arthropod gleaning Hemprich’s long-eared bat (Otonycteris
hemprichii), hypothesizing that there is a trade off between their own and their offspring's food needs. Specifically, we tested the
following predictions: 1) females will bring forward their emergence time from their roosts and increase foraging bout length
progressively from the first through the second trimesters of pregnancy, and during nursing as their pups grow; but, that 2) during
the last trimester of pregnancy, namely, in the final stages of foetus development, females will emerge later and there will be
a reduction in foraging effort (time); and that 3) females will spend more time foraging during nursing than during pregnancy. We
found that the bats emerged from their roosts to forage earlier during the first trimester, but not during the last two trimesters of
pregnancy, and that they did not change their emergence time as nursing progressed. Bats that emerged later at night during
pregnancy spent more time foraging, and, except for the third trimester, the length of the first daily foraging bout increased as both
pregnancy and nursing progressed. Total daily foraging time increased as pregnancy and nursing progressed. Generally, these results
support our hypothesis; through adjustment of their foraging behavior as embryos and pups develop, breeding female Hemprich's
long-eared bats contend with the potentially conflicting food requirements of their offspring and their own needs.
Key words: desert, emergence behavior, Otonycteris hemprichii, reproduction
INTRODUCTION
The distributional range of many organisms ex-
pands and contracts over considerable time periods
(Lyons, 2003) because of natural climatic variation
due to events like glaciations over millions of years
through to regular seasonal variation and migration
(Doak et al., 2008; Olea, 2008). However, some fac-
tors may accelerate distributional shifts; particularly
for highly mobile species in spatially heterogeneous
environments.
With recent climate events and increased urbani -
zation, many species are now finding refuge in new
areas (Van Der Ree and McCarthy, 2005; Williams
et al., 2006; Kurta and Winhold, 2007; Gonzalez-
Megias et al., 2008) and their geographic distri-
bution is expanding and/or contracting at a rapid
rate as a result. Climatic modeling based on known
occur rences for species is an important new tool for
predicting geo graphic distributions (Phillips et al.,
2006a). These techniques also allow us to inves -
tigate re cords of species in contact zones and with
overlapping ranges. For example, ecological niche
based models have been used to predict the range of
overlap in European vipers in a contact zone in
northern Spain (Martinez-Freiria et al., 2008) and
spiny pocket mice in South America (Anderson et
al., 2002).
Large flying-foxes (Pteropus spp.) are capable of
making long-distance flights over a sustained period
of time, such as during southerly summer migra-
tions (Ratcliffe, 1932; Nelson, 1965; Tidemann and
Nelson, 2004). Studies of radio-tagged P. polioce -
pha lus indicate movements of 610 km in 32 days,
210 km in 10 days (Eby, 1991), and up to 750 km
over 6–18 months (Spencer et al., 1991). Two satel-
lite-tagged P. poliocephalus achieved roundtrips
ex ceed ing 2,000 km and > 4° latitude over 7–10
months (Tidemann and Nelson, 2004). Similar sea-
sonal patterns and range of movements have been
recorded for three other Pteropus spp. in Australia,
the black flying-fox, P. alecto (Tidemann et al.,
1999), the spectacled flying-fox, P. conspicillatus
(L. A. Shilton, D. A. Westcott, and P. J. Latch, un-
published data) and the little red flying-fox, P. sca -
pulatus (Sinclair et al., 1996). However, we do not
Acta Chiropterologica, 12(2): 301–309, 2010
PL ISSN 1508-1109 © Museum and Institute of Zoology PAS
doi: 10.3161/150811010X537882
The implications of sympatry in the spectacled and grey headed flying-fox,
Pteropus conspicillatus and P. poliocephalus (Chiroptera: Pteropodidae)
JENNIFER G. PARSONS
1, 4
, JEREMY VAN DER WAL
1, 2
, SIMON K. A. ROBSON
1
, and LOUISE A. SHILTON
1, 3
1
School of Marine and Tropical Biology, James Cook University, Qld, Australia, 4811
2
Centre for Tropical Biodiversity and Climate Change, James Cook University, Qld, Australia, 4811
3
Spatial Conservation, P.O. Box 511, Edge Hill, Qld, Australia, 4870
4
Corresponding author: jennifer.parsons@jcu.edu.au
Sympatry in flying-foxes is common and it occurs throughout coastal Australia, however Pteropus conspicillatus and P. poliocephalus
are thought to have allopatric populations. We examine the distribution of these two flying-fox species and report on the first flying-
fox camp (day roost) with all four species of Australian mainland Pteropus co-occurring. Climatic models were developed from
previous records to determine if the location of this new camp is climatically suitable for these species. We found that this location
is climatically suitable to some degree for both P. conspicillatus and P. poliocephalus but that the latter had a higher climatic
suitability at this site. Historical records exist for P. poliocephalus close to this location but not for P. conspicillatus. The location of
this mixed-species flying-fox camp is the most southerly distribution for P. conspicillatus, being 500 km further south than previous
records. This area of overlap creates potential opportunities for interbreeding between P. conspicillatus and P. poliocephalus.
Therefore, monitoring of this region for the location of further mixed-species camps and the degree of panmixia at those locations
is highly desirable.
Key words: distribution, flying-fox, Pteropus, spatial modeling, sympatry
Echolocation call analysis and presence-only modelling as conservation monitoring
tools for rhinolophoid bats in Thailand
ALICE C. HUGHES
1, 4
, CHUTAMAS SATASOOK
2
, PAUL J. J. BATES
3
, PIPAT SOISOOK
2
, TUANJIT SRITONGCHUAY
2
,
G
ARETH JONES
1
, and SARA BUMRUNGSRI
2
1
School of Biological Sciences, University of Bristol, Woodland Road, Bristol, BS8 1UG, United Kingdom
2
Princess Maha Chakri Sirindhorn Natural History Museum, Faculty of Science, Prince of Songkla University, 15 Karnjanavanit
Road, Hat Yai, Songkhla 90112, Thailand
3
Harrison Institute, Centre for Systematics and Biodiversity Research, Bowerwood House, 15 St Botolph’s Road, Sevenoaks, Kent,
TN13 3AQ, United Kingdom
4
Corresponding author: E-mail: AH3881@bristol.ac.uk
Bats are an important component of biodiversity in Southeast Asia, and are key indicators of habitat quality. Acoustic analysis of
echolocation calls not only allows the identification of bat species that are difficult to capture, but also allows for rapid and
standardised ways to survey and monitor bats over large areas. However keys based on call parameters must also account for
geographic variation in call parameters, and consider any effects of morphology and sex on call frequency. Presence-only modelling
can predict likely geographic locations of specific taxa, and can used to refine decision making so that species unlikely to occur in
a specific region can be omitted from more localised acoustic libraries. Here we develop an acoustic library for the echolocation
calls of rhinolophoid bats in Thailand, and use presence-only modelling (Maxent) to explore how species with similar calls in
a library developed at the national level can sometimes be largely allopatric, and hence identifiable, once geographic range is
predicted. Presence-only modelling can also be used to explore whether species with similar calls adjust call frequency in likely
areas of sympatry. We analysed calls from fourteen species of rhinolophid and twelve hipposiderid species from Thailand. Calls from
a further three rhinolophid and one hipposiderid species are also described but not analysed statistically because of small sample
sizes. Even without considering geographic variation, 69% of rhinolophid (14 species with a minimum of five individuals/ species)
and 91% of hipposiderid calls (12 species) could be classified successfully to species using two call parameters (frequency of most
energy (FMAXE) and duration) in a discriminant function analysis. Most of the discrimination was achieved because species often
utilised different frequency bands. Morphology can also affect call frequency both across and within species. In both rhinolophids
and hipposiderid families there was a negative relationship between FMAXE and forearm length. Within species, FMAXE was
negatively related to forearm length in Rhinolophus microglobosus, R. pusillus and R. thomasi, and positively related to forearm
length in R. affinis and R. pearsonii. Male R. pusillus called at higher frequencies than females, although there was no sexual size
dimorphism in forearm length. Call frequency was negatively related to relative humidity in R. pusillus, suggesting that bats called
at lower frequencies in humid environments to counter increases in atmospheric attenuation. Presence-only modelling was
used to show that some species with similar call frequencies (e.g., R. lepidus and R. microglobosus; R. yunanensis and
R. trifoliatus are predicted to occur largely in allopatry, and so could be identified reliably in particular parts of the country. Presence-
only modeling can assist in predicting areas of overlap between species with similar call frequencies. Other species (e.g.,
R. malayanus, R. coelophyllus) overlap in frequency when data from all of Thailand are combined, but seem to avoid call overlap
when syntopic. Hence acoustic identification can be improved by taking into account local distribution patterns and patterns of
species coexistence. The creation of call libraries at a local scale would have extensive potential as a resource to monitor changes
in species distributions through time.
Key words: constant frequency, acoustic identification, horseshoe bats, Old World leaf-nosed bats, Southeast Asia
INTRODUCTION
Bats are important indicators of ecosystem health
(Jones et al., 2009), and Southeast Asia (Indochina,
Sumatra, Borneo) is one of the three major areas
of bat diversity globally (Proches, 2005), with over
330 species confirmed, including 197 endemics
(King ston, 2010). Southeast Asia also encompasses
four of the world’s biodiversity hotspots (Myers et
al., 2000), and 30–50% of the mammal species in
this region are bats (Kingston et al., 2006). However
many issues threaten the biodiversity of Southeast
Acta Chiropterologica, 12(2): 311–327, 2010
PL ISSN 1508-1109 © Museum and Institute of Zoology PAS
doi: 10.3161/150811010X537891
INTRODUCTION
Although there has been a great deal of research
on the structure and function of echolocation calls
used for foraging and commuting, relatively little
research has been conducted on the communication
functions of bat vocalizations (Pfalzer and Kusch,
2003). In addition to echolocation calls, bats pro-
duce social calls which differ from echolocation
calls in structure and function (Fenton, 2003). Social
calls are usually longer in duration and lower in fre-
quency than echolocation calls and thus, are more
effective at longer distances (Pfalzer and Kusch,
2003). Social calls include mating calls, alarm calls,
territorial calls, and food calls (Wilkinson, 1995)
and can aid in finding patchily distributed food
resources (Wilkinson, 1992; Safi and Kerth, 2007)
and roosts (Ruczyński et al., 2007, 2009). In con-
trast, echolocation calls are used to gain information
about the environment (Fenton, 2003). Because
echo location calls can be intercepted by other bats,
they also can serve a communication function, as in
the case of eavesdropping (e.g., Barclay, 1982; Bal -
combe and Fenton, 1988; Gillam, 2007). Further,
differences in echolocation call structure may play
an important role in species recognition (Russo et
al., 2007).
Bats exhibit a wide range of responses to echolo-
cation and social calls made by other bats. Calls may
attract bats to a common foraging area (e.g., Wil kin -
son, 1992) or act as a spacing mechanism (Leonard
and Fenton, 1984; Barlow and Jones, 1997). Some
bats respond only to the calls of conspecifics where-
as others respond to both conspecifics and het-
erospecifics. Myotis yumanensis and Myotis lucifu-
gus in British Columbia only respond to the echolo-
cation calls of conspecifics (Balcombe and Fenton,
1988), but M. lucifugus in Ontario respond to echo -
location calls of conspecifics as well as to calls of
Eptesicus fuscus (Barclay, 1982). Further more, re-
sponses may vary with echolocation call type.
Individuals of Tadarida bra si liensis are attracted to
Acta Chiropterologica, 12(2): 329–336, 2010
PL ISSN 1508-1109 © Museum and Institute of Zoology PAS
doi: 10.3161/150811010X537909
Intra- and interspecific responses to Rafinesque’s big-eared bat
(Corynorhinus rafinesquii) social calls
SUSAN C. LOEB
1, 3
and ERIC R. BRITZKE
2
1
USDA Forest Service, Southern Research Station, Department of Forestry and Natural Resources, Clemson University,
Clemson, SC 29634, USA
2
US Army Engineer Research and Development Center, 3909 Halls Ferry Road, Vicksburg, MS 39180, USA
3
Corresponding author: E-mail: sloeb@fs.fed.us
Bats respond to the calls of conspecifics as well as to calls of other species; however, few studies have attempted to quantify these
responses or understand the functions of these calls. We tested the response of Rafinesque’s big-eared bats (Corynorhinus
rafinesquii) to social calls as a possible method to increase capture success and to understand the function of social calls. We also
tested if calls of bats within the range of the previously designated subspecies differed, if the responses of Rafinesque’s big-eared
bats varied with geographic origin of the calls, and if other species responded to the calls of C. rafinesquii. We recorded calls of
Rafinesque’s big-eared bats at two colony roost sites in South Carolina, USA. Calls were recorded while bats were in the roosts and
as they exited. Playback sequences for each site were created by copying typical pulses into the playback file. Two mist nets were
placed approximately 50–500 m from known roost sites; the net with the playback equipment served as the Experimental net and
the one without the equipment served as the Control net. Call structures differed significantly between the Mountain and Coastal
Plains populations with calls from the Mountains being of higher frequency and longer duration. Ten of 11 Rafinesque’s big-eared
bats were caught in the Control nets and, 13 of 19 bats of other species were captured at Experimental nets even though overall bat
activity did not differ significantly between Control and Experimental nets. Our results suggest that Rafinesque’s big-eared bats are
not attracted to conspecifics’ calls and that these calls may act as an intraspecific spacing mechanism during foraging.
Key words: call playback, Corynorhinus rafinesquii, eavesdropping, echolocation, social calls
Acta Chiropterologica, 12(2): 337–340, 2010
PL ISSN 1508-1109 © Museum and Institute of Zoology PAS
doi: 10.3161/150811010X537918
Why the bat did not cross the road?
ARTHUR A. ZURCHER
1, 4
, DALE W. SPARKS
2
, and VICTORIA J. BENNETT
3
1
Center for North American Bat Research and Conservation, Department of Biology, Indiana State University, Terre Haute,
IN 47809, USA
2
Environmental Solutions and Innovations, 781 Neeb Road, Cinncinati, OH 45233, USA
3
Department of Forestry and Natural Resources, Purdue University, West Lafayette, IN 47907, USA
4
Corresponding author: E-mail: azurcher1@indstate.edu
Roadways are nearly ubiquitous parts of the modern landscape, but their impact on bats remain relatively unknown. We studied the
influence of vehicular traffic on the behaviour of commuting bats near the Indianapolis International Airport. A previous study at
this site documented that Indiana bats (Myotis sodalis) were much more likely to cross roads with low traffic volumes. One potential
interpretation of this result is that bats perceive motor vehicles as a threat and exhibit avoidance behaviour whether or not the bats
are in immediate danger. To test this hypothesis, we observed 211 cases of bats approaching roads that bisected their commuting
routes. Information recorded at the time included the presence or absence of vehicles, the height the bat was flying, whether a bat
reversed course prior to crossing the road and if so the distance from the road or vehicle (if present) when it altered its direction, and
finally the speed, type and relative level of noise emitted by vehicles. Results revealed that bats were more than twice as likely to
reverse course when vehicles were present as opposed to their absence. When automobiles were present 60% of bats exhibited
avoidance behaviour, reversing course at an average of 10 m from a vehicle. Conversely, when no automobiles were present, only
32% of bats reversed their course and 68% crossed the road. The height a bat flew, speed of the vehicle, type of vehicle or level of
noise emitted by vehicles had no effect on the likelihood of bats reversing course. These data support the hypothesis that bats
perceive vehicles as a threat and display anti-predator avoidance behaviour in response to their presence.
Key words: anti-predator behaviour, automobiles, bats, behavioural modification, commuting, roadways
INTRODUCTION
Despite their nearly ubiquitous nature, the direct
and indirect effects of roadways remain poorly stud-
ied for many taxa. This is especially true for bats,
a taxon for which the first studies are just now being
published (Lesiński, 2007, 2008; Gaisler et al.,
2009; Russell et al., 2009). These papers as well
as scattered anecdotal (and often unpublished) ac-
counts of bats hit by vehicles (Sparks and Choate,
2000; Butchkowski and Hassinger, 2002; Dodd et
al., 2004; Capo et al., 2006; Choquene, 2006; Glista
and DeVault, 2008; Russell et al., 2008; Hein et al.,
2009; Whitaker and Mumford, 2009) suggest that
roadways may be a significant source of mortality
for bats. However, direct mortality is not the only
impact of road networks. Roadways may also serve
as a geographic barrier reducing the ability of bats
to access suitable foraging or roosting sites (Kerth
and Melber, 2009; M. A. McGuire, D. W. Sparks,
J. O. Whitaker, Jr., and V. Brack, Jr., unpublished
data). Currently, there is very little information on
how bats respond to the roads they encounter while
commuting to foraging habitats. Comparative stud-
ies have been conducted to explore foraging behav-
iour (Waters et al., 1999; Loeb and O’Keefe, 2006;
Hein et al., 2009). For example, some bats have
been found to avoid foraging near roadways as the
noise of passing traffic disrupts their ability to
echolocate effectively (Schaub et al., 2008). How -
ever, bats do not necessarily depend on echolocation
to commute (Russell et al., 2009). While commuting
bats have two primary objectives: 1) To reach
foraging habi tat in smallest amount of time (i.e., rate
of mov ement is faster and flight paths are straight
(Schaub and Schnitzler, 2007; Grodziński et al.,
2009) and 2) to avoid predators (i.e., by flying par-
allel to linear features and/or under cloud cover
(Limpens and Kap teyn, 1991; Verboom and Spoel -
stra, 1999; Cie cha now ski et al., 2007). Thus the
presence of a road along a commuting route can, as
an open habitat feature, expose the bats predation
INTRODUCTION
The Old World fruit bats (Chiroptera: Pteropodi -
dae) are important for the survival of more than
114 plant species of the world (Mickleburgh et al.,
1992). Their positive roles as pollinators, seed dis-
persers, and as agents for maintaining plant commu-
nity diversity has been acknowledged universally
(van der Pijl, 1982; Marshall, 1983; Cox et al.,
1991; Fujita and Tuttle, 1991; Mickleburgh et al.,
1992; Rainey et al., 1995; Eby, 1996; Banack,
1998). Pteropodids are represented by three genera
and four species in Pakistan (Roberts, 1997; Mah -
mood-ul-Hassan et al., 2009) and includes the short-
nosed fruit bat (Cynopterus sphinx), the Indian fly-
ing fox (Pteropus giganteus), the Egyptian fruit bat
(Rousettus aegyptiacus) and the fulvous fruit bat
(Rousettus leschenaultii).
Distributed in Maldive Isles, India (including
Andaman Isles), Sri Lanka, Pakistan, Bangladesh,
Nepal and Burma, P. giganteus is one of the largest
bats in the world (Simmons, 2005). It is often la-
beled as ‘vermin’ on the assumption that it poaches
ripe fruits from orchards and defecates in public
places (Chakraverthy et al., 2009; Mahmood-ul-
Hassan et al., 2009). This bat is thought to cause
heavy economic losses to arecanut (Areca catechu),
sapota (Achrus zapota), guava (Psidium guajava),
mango (Mangifera indica), and jaman (Syzigium
jam bolanum) (Roberts, 1997; Chakraverthy and Gi -
rish, 2003). Pteropus giganteus is given no protec-
tion by Pakistani law and is hunted for its body fat
to be used as potions and as putative cures for rheu-
matic pains, by local medical practitioners known as
‘hakeems’ (Roberts, 1997). Pteropus giganteus is
included in the fourth schedule of the Punjab
Wildlife (Protection, Preservation, Conservation and
Management) Act 1974 section 2 (v), which lists the
species among those that are given no legal protec-
tion and can be hunted.
Although extensive literature exists on the food
habits of some Pteropus species (e.g., Dobat and
Peikert-Holle, 1985; Marshall, 1985; Mickleburgh
et al., 1992; Wiles and Fujita, 1992), few studies
offer a detailed investigation of their diet in a single
area (Banack, 1998). Information is particularly
scanty regarding the diet of P. g. giganteus in
urban landscapes and no attempt has been made
to quantify the ecological role of this species. The
present study was designed to determine whether
Acta Chiropterologica, 12(2): 341–347, 2010
PL ISSN 1508-1109 © Museum and Institute of Zoology PAS
doi: 10.3161/150811010X537927
The diet of Indian flying-foxes (Pteropus giganteus) in urban habitats of Pakistan
MUHAMMAD MAHMOOD-UL-HASSAN
1, 3
, TAYIBA L. GULRAIZ
1
, SHAHNAZ A. RANA
2
, and ARSHAD JAVID
1
1
Department of Wildlife and Ecology, University of Veterinary and Animal Sciences, Syed Abdul Qadir Jilani Raod, Lahore, Pakistan
2
Department of Zoology, University of Agriculture, Faisalabad, Pakistan
3
Corresponding author: E-mail: drmmhassan@gmail.com
We studied the diet of the Indian flying fox (Pteropus giganteus) in Pakistan from March 2008 to February 2009 and found that
the bats fed on 20 species belonging to 11 plant families. Of these, four families (Anacardiaceae, Bignonaceae, Malvaceae, and
Sapotaceae) were identified from remnants of flower petals in food boluses while the remaining families (Annonaceae, Arecaceae,
Ebenaceae, Meliaceae, Moraceae, Myrtaceae, and Sapindaceae) were identified from the seeds in the boluses and from guano
samples. Plants in the family Moraceae (50.7%) comprised most of the bat’s diet. Fruit of Ficus retusa (27.5%) and F. carica (23.0%)
during winter, F. glomerata (30.9%) and F. religiosa (28.1%) during spring, Psidium guajava (19.6%), F. bengalensis (18.7%) and
Diospyros peregrina (17.8%) during summer, and D. peregrina (71.9%) during autumn, were the most frequently identified items.
The four seasonal diets varied significantly (χ
2
= 435, d.f. = 18, P < 0.01). Results confirm that the ecological services rendered by
P. giganteus, such as pollination and seed dispersal, outweigh its losses, such as damage to the ripe fruit. Hence, the species should
not be regarded as a pest; rather efforts should be made to ensure its conservation.
Key words: conservation, Ficus, fruit bat, Pakistan, pest, seed dispersal, urban habitat
INTRODUCTION
Foraging habitats of British bats are largely de-
termined by the abundance of their insect prey (Ra -
cey and Swift, 1985). In recent decades changing
agricultural practices have negatively influenced
farm land bat populations (Duvergé and Jones,
2003). These have ranged from agricultural intensi-
fication and the use of chemicals on farmland
(McAney and Fairley, 1989; Wickramasinghe et al.,
2003) to the production of silage rather than hay
(Entwistle et al., 2001).
Whilst much conservation effort has concentrat-
ed on conservation of bat roosts, future management
should also aim to conserve and correctly manage
key foraging habitats surrounding roosts (Mitchell-
Jones et al., 1993; Jones et al., 1995; Entwistle et
al., 1996; Racey, 1998; Arlettaz et al., 2000; Bon ta -
dina et al., 2001). Most microbat species forage
within a few kilometres of their day roosts (e.g.,
Catto et al., 1996; Siemers et al., 1999; Bon ta dina et
al., 2002; Dense and Rahmel, 2002; Steinhauser,
2002; Cordes, 2004; Encar nacão et al., 2005; Nich -
olls and Racey, 2006; Mackie and Racey, 2007;
Smith and Racey, 2008), hence habitats within this
area should be critical for habitat conservation.
Riparian areas, broad-leaved woodland and lin-
ear landscape features such as hedges are recognised
as important habitat features for foraging bats
(Boon man et al., 1995; Walsh and Harris, 1996;
Verboom and Huitema, 1997; Entwistle et al., 2001;
Downs and Racey, 2006). Grazed pasture, particu-
larly that grazed by domestic cattle (Bos taurus), is
also increasingly being considered as an important
foraging habitat. Maintenance of unimproved pas-
ture through cattle grazing ensures a supply of dung
associated insects (Entwistle et al., 2001). This has
been well documented with regard to the greater
Acta Chiropterologica, 12(2): 349–358, 2010
PL ISSN 1508-1109 © Museum and Institute of Zoology PAS
doi: 10.3161/150811010X537936
Do bats forage over cattle dung or over cattle?
NICK C. DOWNS
1, 3
and LAURA J. SANDERSON
1,2
1
Hyder Consulting Limited, Cresswell Associates, The Mill, Brimscombe Port, Stroud, Gloucestershire, GL5 2QG,
United Kingdom
2
School of Animal and Microbial Sciences, University of Reading, Whiteknights, P.O. Box 228, Reading, Berkshire,
RG6 6AJ, United Kingdom
3
Corresponding author: E-mail: nhi768@yahoo.co.uk
Although the importance of cattle grazed pasture to foraging bats has already been well documented, the relative importance of cattle
dung compared with the cattle themselves is largely unknown. This was investigated by comparing two adjacent fields: a current
and a recent area of organic cattle-grazed pasture in England. The cattle were exchanged between fields during each of two separate
observation periods in June and July 2005, one of eight evenings and the other of ten. Hand-held detectors were used to record
time-expanded bat passes simultaneously from comparable positions within each field for 150 minutes after sunset. In addition, two
separate observation periods (one of three evenings and the other of four) was used to investigate bat activity over and away from
cattle within the same field. In general, bat activity was statistically greater within the field containing cattle than in the field
without cattle (paired t-test, t
21
= 3.97, P < 0.001). Within the field containing cattle, bat activity was greater over cows than away
from them (paired t-test, t
6
= 2.48, P < 0.05). It is therefore suggested that cattle, rather than dung are the primary attractant
for foraging bats. Specifically, the difference in bat activity between the fields with and without cattle was statistically significant
for Eptesicus serotinus (paired t-test, t
21
= 4.14, P < 0.001), Pipistrellus pipistrellus (paired t-test, t
21
= 3.23, P < 0.01) and
Myotis (Dunn’s test, Z = 2.99, P < 0.01) bats. Management recommendations include keeping cattle within bat foraging areas,
minimizing the use of anthelminthic avermectin-based drugs for cattle, and maintaining woodland/trees within and around cattle-
grazed pasture.
Key words: Chiroptera, farming, foraging, management, England
INTRODUCTION
Domestic cats, Felis catus, are opportunistic for-
agers especially adapted to prey on small burrowing
rodents and hares. However, in their feral or semi-
feral state, their ability to use alternate prey, or even
scavenge, enables them to maintain their abundance
when their main prey is not available (Fitzgerald and
Turner, 2000; Harper, 2004; Calver et al., 2007).
Their long association with humans has lead the way
for a large body of literature on foraging behavior of
cats, both feral and otherwise (e.g., Biben, 1979;
Liberg, 1984; Harper, 2005), mainly in the temper-
ate zones. This association with humans has led to
the establishment of large populations of cats, re-
sulting in a variety of problems in urban, rural, and
natural areas.
Free-roaming cats include owned cats allowed
outdoors, abandoned or lost cats, as well as feral and
semi-feral cats. Semi-feral cats are defined as cats
that are directly fed by humans on a regular basis,
but not owned (McGee Hill, 2006), while feral cats
are never purposely fed by humans. In this study we
examine a little studied aspect of cat’s foraging
behavior: bat predation on a tropical island where
bats comprise the dominant component of the mam-
malian fauna.
M
ATERIALS AND METHODS
Observations were made at Culebrones Cave in northern
Puerto Rico, West Indies (18°24’51.59”N, 66°43’43.58”W).
The cave is located within the northern karst region, where bat
densities are higher than anywhere else on the Island. Cule -
brones cave is classified as a ‘hot cave’ (Rodríguez-Durán, 2009),
with a multispecies assemblage of bats estimated at 300,000 in-
dividuals of six species (Rodríguez-Durán, 1996): Brachyphylla
ca ver narum, Erophylla bombifrons, Monophyllus redmani,
Mor moops blainvillei, Pteronotus quadridens and P. parnellii.
The cave consists of a main tunnel over 600 m deep with mul-
tiple chambers where bats distribute according to species, and
constrictions that function as heat traps. The entrance, at the
base of a hill, slants downwards about 45°.
We visited the site two to four times a month from April to
August, 2006 and from March to July, 2007, and at least once
monthly at other times of the year. The cave had been visited on
a regular basis since 1995. Wing and leg remains left by the cats
after consuming the bats were collected and compared to study
collections at Universidad Interamericana to identify the bat
species to which they belonged. On each visit, all wing remains
were removed from the area surrounding the cave opening
Acta Chiropterologica, 12(2): 359–362, 2010
PL ISSN 1508-1109 © Museum and Institute of Zoology PAS
doi: 10.3161/150811010X537945
Predation by free-roaming cats on an insular population of bats
ARMANDO RODRÍGUEZ-DURÁN
1, 2
, JANSEL PÉREZ
1
, MARI ANA MONTALBÁN
1
, and JEAN M. SANDOVAL
1
1
Universidad Interamericana, 500 John W. Harris Ave., Bayamón, PR 00957, USA
2
Corresponding author: E-mail: arodriguez@bc.inter.edu
Free-roaming cats are known to adversely impact native faunas in the areas where they have been introduced, an impact that is even
greater on islands. We examine the predation of bats by cats at Culebrones cave, Puerto Rico, West Indies. Culebrones cave is a hot
cave located in the karst region of northern Puerto Rico. The temperature gradient inside the cave sustains a multi-species
assemblage of bats consisting of approximately 300,000 individuals of six species, namely: Brachyphylla cavernarum, Erophylla
bombifrons, Monophyllus redmani, Mormoops blainvillei, Pteronotus quadridens and Pteronotus parnellii. Even though rats are
often their primary prey, cats will use alternative prey, which enables them to maintain their abundance when one prey is not
available. In Puerto Rico, birds and reptiles are known to be preyed upon by cats. Although cats are commonly observed in or
around bat caves in Puerto Rico, this is the first systematic attempt to evaluate their role as bat predators. We made observations
of the hunting strategy of cats using an infrared camera and recorded the number of wings left as remains of these hunting
bouts. Wings were identified to species. Cat scats were also recovered and examined to identify prey species. Our results
suggest that captures of different species of bats is not a function of their abundance in the cave. While M. blainvillei (11 g) and
P. quadridens (5 g) are the most abundant species in the cave, B. cavernarum (50 g) and M. redmani (11 g) are captured in greater
numbers by the cats.
Key words: islands, tropical bats, predation, cats, foraging behavior
INTRODUCTION
Numerous methodologies exist for the study of
mammals including live traps, kill traps, acoustic
monitoring, and visual observation (Gannon et al.,
2007). However, commonly used techniques some-
times become the standard for studying particular
taxa, despite a lack of critical assessment of proce-
dures available. Significant declines in populations
of bats over the past several decades have been
reported for certain species in Europe, the United
States, and Mexico (Hutson et al., 2001). More
recently, a fungal-condition known as white-nose
syndrome (WNS) has been linked to massive die-
offs of bats in the northeastern United States
(Blehert et al., 2009; Reichard and Kunz, 2009).
Therefore, without conclusive data on the status of
most populations, there is an urgency to explore the
best methods for surveying these mammals.
Methods commonly used to study bats include
acoustic surveys using ultrasonic bat detectors or
capture devices such as bat traps and mist nets
(Kunz, 1988). While ultrasonic surveys are less in-
vasive and have been shown to improve estimates of
species richness and habitat use over conventional
mist-netting (Murray, et al., 1999), studies seeking
demographic data on populations must rely on actu-
al captures. Less attention has been given to com-
parisons of capture techniques and to our knowl-
edge, no studies have been done on Nearctic bats.
Bat traps were found to be more efficient than mist
nets in Australia (Tidemann and Woodside, 1978)
and Malaysia (Francis, 1989). Because of the small
size of bat traps, they are more commonly used for
trapping small corridors and cave entrances. Mist
nets are the most frequently used method to capture
bats in flight, most likely due to cost and the ability
to use multiple nets to cover a greater area. Many
different ways of deploying mist nets have been
proposed, including canopy nets, low nets, and dif-
ferent combinations of nets (Kunz and Kurta, 1988).
In Malaysia, studies comparing low nets to canopy
nets have found a significantly higher capture rate
and species diversity in canopy nets (Francis, 1994;
Zubaid, 1994). Others have suggested it is impor-
tant to use multiple net configurations (Helman and
Churchill, 1986; Kunz and Kurta, 1988; Larsen et
al., 2007).
Acta Chiropterologica, 12(2): 363–369, 2010
PL ISSN 1508-1109 © Museum and Institute of Zoology PAS
doi: 10.3161/150811010X537954
A test of mist-net configurations in capturing bats over stream corridors
L. MICHELLE GILLEY
1, 3
and MICHAEL L. KENNEDY
2
1
Department of Biological Sciences, 331 Funchess Hall, Auburn University, Alabama 36849, USA
2
Department of Biology, Ellington Hall 307, The University of Memphis, Tennessee 38152, USA
3
Corresponding author: E-mail: gillelm@auburn.edu
Most surveys for bats are conducted using mist nets in riparian areas along stream corridors. Various methods exist for deploying
mist nets, but success of using different configurations has not been assessed. We tested efficiency of three configurations of mist
nets during summers of 2000 and 2001 at the Milan Army Ammunition Plant (Carroll and Gibson counties) in western Tennessee.
Configurations of mist nets included: I (one net placed transverse to stream), T (one net placed transverse to stream, and one net
positioned perpendicular to first net in midstream), and Z (two nets positioned parallel to stream, and a center net positioned
diagonally between the two nets). The study consisted of 347 net nights and 220 captures of bats (85 I, 62 T, 73 Z). Four species
were captured including: 133 Lasiurus borealis, 63 Perimyotis subflavus, 15 Nycticeius humeralis, and nine Myotis austroriparius.
Sex-ratios for adults were female biased, while juvenile sex-ratios were near equal. Netting results suggest that traditional
I-configurations were statistically equal to T- and Z-configurations for all analyses of total captures and for the two dominant species
captured: L. borealis and P. subflavus. Because the I-configuration requires less equipment and time for set-up, capturing bats in
linear corridors could be optimized by using more I-nets rather than multiple net configurations.
Key words: bats, capture techniques, Lasiurus borealis, mist-net configuration, Perimyotis subflavus, Tennessee
INTRODUCTION
The distribution of Southern African bat species
is poorly known compared with other small mam-
mal taxa such as rodents (Skinner and Chimimba,
2005; Monadjem et al., 2010), or compared with
other regions such as Europe (Mitchell-Jones et al.,
1999). Our knowledge of the bat fauna of Mo -
zambique is particularly inadequate, given that the
most recent synopsis is 35 years old (Smithers and
Lobão Tello, 1976). At that time, only a single site
(Ilha de Mozambique) had been surveyed in the
northern provinces (north of the Zambezi River), an
area covering almost 50% of the country. The re-
maining ‘southern’ half of the country was only
patchily covered, with 19 species known from single
localities and a further nine from just two localities.
Hence, prior to 2000, a total of 56 bat species were
known to occur in Mozambique, and 28 (50%) of
these were known from two or fewer sites (Smithers
and Lobão Tello, 1976) in a country covering over
801,000 km
2
. (At least three of these 56 species are
based on misidentifications — see Results below).
A further indication of the under-sampled state of
bats in Mozambique is that smaller neighbouring
countries have higher species richness: for example,
Acta Chiropterologica, 12(2): 371–391, 2010
PL ISSN 1508-1109 © Museum and Institute of Zoology PAS
doi: 10.3161/150811010X537963
A recent inventory of the bats of Mozambique with documentation of seven
new species for the country
ARA MONADJEM
1, 13
, M. CORRIE SCHOEMAN
2
, APRIL RESIDE
3
, DOROTHEA V. P IO
4
, SAMANTHA STOFFBERG
5
,
J
ULIAN BAYLISS
6, 7
, F. P. D. (WOODY) COTTERILL
8
, MICHAEL CURRAN
9, 10
, MIRJAM KOPP
9
, and
P
ETER J. TAYLOR
11, 12
1
All Out Africa Research Unit, Department of Biological Sciences, University of Swaziland, Private Bag 4, Kwaluseni, Swaziland
2
School of Biological and Conservation Sciences, University of KwaZulu-Natal, Durban, Republic of South Africa
3
All Out Africa, P.O. Box 153, Lobamba, Swaziland
4
Département d’Ecologie et Evolution, Université de Lausanne, Biophore 1015, Lausanne, Switzerland
5
Evolutionary Genomics Group, Department of Botany and Zoology, University of Stellenbosch, Private Bag X1, Matieland,
Stellenbosch, Republic of South Africa
6
Mulanje Mountain Conservation Trust (MMCT), P.O. Box 139, Mulanje, Malawi
7
Conservation Science Group, Department of Zoology, University of Cambridge, Cambridge, UK
8
AEON – Africa Earth Observatory Network, Departments of Geological Sciences, and Molecular and Cell Biology,
University of Cape Town, Rondebosch 7701, Republic of South Africa
9
Institute of Biogeography, University of Basel, St. Johanns-Vorstadt 10, CH-4056, Switzerland
10
Institute of Environmental Engineering, ETH Zurich, HIF C 13, Wolfgang-Pauli-Str. 15, CH-8093 Zurich, Switzerland
11
Durban Natural Science Museum, P. O. Box 4085, Durban, Republic of South Africa
12
Department of Ecology and Resource Management, School of Environmental Sciences, University of Venda,
Private Bag X5050, Thohoyandou, 0950, Republic of South Africa
13
Corresponding author: E-mail: ara@uniswacc.uniswa.sz
The bat fauna of Mozambique is poorly documented. We conducted a series of inventories across the country between 2005 and
2009, resulting in the identification of 50 species from 41 sites. Of these, seven species represent new national records that increase
the country total to 67 species. These data include results from the first detailed surveys across northern Mozambique, over an area
representing almost 50% of the country. We detail information on new distribution records and measurements of these specimens.
Special attention is paid to the Rhinolophidae, because these include several taxa that are currently in a state of taxonomic confusion.
Furthermore, we also present some notes on taxonomy, ecology and echolocation calls. Finally, we combine modelled distributions
to present predicted species richness across the country. Species richness was lowest across the coastal plain, to the east and far north,
and is predicted to increase in association with rising altitude and higher topographic unevenness of the landscape.
Key words: Mozambique, Chiroptera, distribution, check-list, conservation
INTRODUCTION
Bats of the families Rhinolophidae and Hippo -
sideridae have been recently documented as sister
taxa within the superfamily Rhinolophoidea and are
considered sister groups of Megachiroptera within
the clade of Pteropodiformes (Teeling et al., 2005;
Hutcheon and Kirsch, 2006; Jones and Teeling,
2006). The rhinolophid bats are widely distributed
in the Old World and their range extends from
Western Europe and Africa to Japan, Philippines
and Australia. Within the family Rhino lophidae,
only a single genus is recognized consisting of 77
extant species (Corbet and Hill, 1992; Sim mons,
2005). Chromosomal data are available for 38 spe -
cies from Europe, Africa, Asia Minor, Middle East,
India, Japan, China, Thailand, and Ma lay sia (see
Zima et al., 1992 for review; Rickart et al., 1999;
Volleth et al., 2002; Gu et al., 2003; Wu et al., 2004;
Wu and Harada, 2005; Karataş et al., 2006; Mao et
al., 2007; Puerma et al., 2008). Most species have
a karyotype with a 2n of 58 or 62.
Hipposiderid bats are distributed in the tropics
and subtropics of the Old World and the family
includes nine genera and 81 species. The genus
Hipposideros is the largest within the family with
67 species (Bogdanowicz and Owen, 1998; Sim -
mons, 2005). The karyotypes of 22 species of the
family Hipposideridae have been reported so far
(Ray-Chaudhuri and Pathak, 1966; Ray-Chaudhuri
et al., 1971; Ando et al., 1980; Handa and Kaur,
1980; Zhang and Wan, 1992; Choudhury and Patro,
1993; Sreepada et al., 1993; Gu, 2002a, 2002b,
2006; Wu et al., 2006) and a limited extent of varia-
tion is evident. In most of the species studied,
a karyotype with 32 biarmed chromosomes is pres-
ent (with the fundamental number of autosomal
arms FNa = 60). Within the genus Hipposide ros,
exceptions from this uniform pattern were found
in H. com mersoni from South Africa (Rauten bach
et al., 1993) and in H. obscurus from Philip pines
(Rickart et al., 1999). The karyotype of H. com mer -
soni had a 2n of 52 whereas that of H. obscurus a 2n
of 24 and FNa = 44.
In this paper, non-differentially stained karyo -
types of ten species of the genera Rhinolophus
and Hipposideros from western Africa and south-
ern India are examined and these karyotypes
are compared to those in the literature. The aim
is to demonstrate that considerable chromosomal
Acta Chiropterologica, 12(2): 393–400, 2010
PL ISSN 1508-1109 © Museum and Institute of Zoology PAS
doi: 10.3161/150811010X537972
Karyotypic variation in rhinolophid and hipposiderid bats (Chiroptera:
Rhinolophidae, Hipposideridae)
DARINA KOUBÍNOVÁ
1, 4
, K. S. SREEPADA
2
, PETR KOUBEK
3
, and JAN ZIMA
3
1
Department of Zoology, Faculty of Science, Charles University, Viničná 7, CZ-12844 Praha 2, Czech Republic
2
Department of Applied Zoology, Mangalore University, Mangalagangotri - 574 199, Karnataka, India
3
Institute of Vertebrate Biology, Academy of Sciences, Květná 8, CZ-60365 Brno, Czech Republic
4
Corresponding author: E-mail: darina.koubinova@gmail.com
According to current phylogenetic hypotheses, the bats of the families Rhinolophidae and Hipposideridae are sister groups nested
within the clade of Pteropodiformes. A conservative nature of karyotypic evolution was previously reported within the two families.
Karyotypes with diploid number (2n) varying between 58 and 62 chromosomes were assumed to prevail among the rhinolophid
species, whereas, karyotypes with 32 chromosomes were found in most of the hipposiderid bats. However, divergent lower or higher
2n numbers have been recorded in some species in both families. Variation is documented in the present paper by examination of
non-differentially stained karyotypes in 10 species belonging to genera Rhinolophus and Hipposideros from western Africa and
southern India. Among the species studied, the karyotypes with a 2n of 32, 36, 52, 56, 58, and 62, and with relatively stable number
of autosomal arms (FNa = 60, 62, 64) were recorded.
Key words: chromosomal evolution, karyosystematics, Robertsonian translocations
INTRODUCTION
Due to increased potential for long distance dis-
persal, bats commonly exhibit less pronounced pop-
ulation structure than nonvolant mammals (Ditch -
field, 2000). Comparative phylogeographic studies
have confirmed this prediction for many chiropteran
species (Patterson et al., 1992; McCracken et al.,
1994; Sinclair et al., 1996; Webb and Tidemann,
1996; Petit and Mayer, 1999, 2000; Ditchfield,
2000; Russell et al., 2005; Bryja et al., 2009). While
increased dispersal ability is a major reason for
reduced population structure in volant organisms,
the migratory behavior of many bat species (Mc -
Cracken et al., 1994; Webb and Tidemann, 1996;
Petit and Mayer, 1999, 2000; Russell et al., 2005;
Bryja et al., 2009), as well as birds (Buerkle, 1999;
Davis et al., 2006; Hellgren et al., 2008) also con-
tributes. Seasonal migrations allow what are tem-
porarily isolated populations to exchange genes,
often homogenizing populations over large geo-
graphic areas.
In spite of the potential for reduced popula-
tion structure, divergent genetic lineages have been
detected in some bat species (Worthington Wilmer
et al., 1994; Wilkinson and Fleming, 1996; Lloyd,
2003; Miller-Butterworth et al., 2003; Roberts,
2006; Weyandt and Van Den Bussche, 2007). Most
commonly, these are nonmigratory species and/or
species that exhibit strong female philopatry evident
using mitochondrial DNA (mtDNA) markers. When
biparentally inherited nuclear markers are used,
strong population structure often disappears (Cas -
tella et al., 2001; Kerth et al., 2002), although
there are exceptions (Miller-Butterworth et al.,
2003). The pattern of male-biased dispersal is
extremely common in mammals (Greenwood,
1980), yet mammalian phylogeographic studies are
dominated by maternally inherited mtDNA markers.
While these markers can provide insight into his-
torical evolution and population demographics,
their utility in identifying and quantifying patterns
of contemporary gene flow in mammals is limit-
ed, and the reliability of a mtDNA-only approach
has been questioned (Galtier et al., 2009). A more
comprehensive approach includes biparentally
inherited nuclear markers in conjunction with
mtDNA.
Acta Chiropterologica, 12(2): 401–413, 2010
PL ISSN 1508-1109 © Museum and Institute of Zoology PAS
doi: 10.3161/150811010X537981
Range-wide population genetic structure of the pallid bat (Antrozous pallidus)
— incongruent results from nuclear and mitochondrial DNA
JUSTIN B. LACK
1, 2
, JEREMY E. WILKINSON
1
, and RONALD A. VAN DEN BUSSCHE
1
1
Department of Zoology, 501 Life Sciences West, Oklahoma State University, Stillwater, OK 74078, USA
2
Corresponding author: E-mail: justin.lack@okstate.edu
Male-biased dispersal with female philopatry is a common pattern for many mammals. Because mtDNA is inherited maternally,
biparentally inherited nuclear markers are often utilized to estimate population structure and gene flow. The pallid bat (Antrozous
pallidus) exhibits a relatively continuous distribution across arid western North America. A previous mtDNA study suggested little
gene flow among populations and identified three distinct phylogroups. We examined population structure of A. pallidus using
amplified fragment length polymorphisms (AFLP) in 187 individuals from 29 localities across the distribution of the species. Eight
primer pairs identified 797 polymorphic loci. All analyses indicated that populations in California, British Columbia, and the Baja
Peninsula were relatively distinct. Cluster analysis indicated gene flow has been occurring between the Baja Peninsula population
and populations to the north and east. The pallid bat appears to be characterized by male-mediated dispersal and gene flow, while
females are largely philopatric. While several populations appear to be somewhat distinct, the overall pattern of divergence for
A. pallidus is indicative of isolation by distance throughout the majority its range, suggesting significant gene flow has been
occurring since populations diverged during Pliocene desert formation and mountain building.
Key words: Antrozous pallidus, philopatry, phylogeography, AFLP, sex-biased dispersal, Pliocene
INTRODUCTION
The taxonomy of Hipposideros bicolor Tem-
m inck, 1834 and Hipposideros pomona Andersen,
1918 is complex and confusing. H. bicolor was de-
scribed without either a holotype or type locality be-
ing designated. Later Temminck (1835) provided
a more detailed description which was based on 14
specimens from Java, Amboina and Timor. How -
ever, according to Hill (1963), this series was a com-
posite, with the smaller specimens from Amboina
referable to a form subsequently described as Phyl -
lo rhina amboinensis Peters, 1871, which is now con -
sidered to be a synonym of Hipposideros ater Temp -
leton, 1848. Subsequently, Tate (1941) designated
a lectotype of H. bicolor based on a specimen in
the Rijks museum van Natuurlijke Historie, Lei den
and restricted the locality to Côte d’Anjer in north -
western Java. This lectotype and others in the series
were reviewed by Bergmans and van Bree (1986).
Meanwhile, Andersen (1918) briefly reviewed
the H. bicolor group. Unfortunately, he was con-
fused about the taxon bicolor and according to Hill
(1963) his short description appears more referable
to H. ater. In this paper, he also named H. pomona
from southern In dia and H. gentilis from Myanmar,
both in the H. bicolor group. He then named and
referred three new taxa, sinensis, atrox, and major to
H. gentilis as subspecies. Tate (1941) followed
Ander sen (1918) and also included these latter taxa
in H. gentilis. However, this view was not accepted
by Hill (1963) who included all five taxa, pomona,
gentilis, sinensis, atrox and major as subspecies of
H. bicolor. Subsequently, Hill et al. (1986) revised
this opinion and considered that H. bicolor, sensu
Hill (1963), actually comprised two species H. bico -
lor and H. pomona. They referred the taxa major
and atrox to H. bicolor and gentilis and sinensis
to H. pomona. This view was followed by Yen bu-
tra and Felten (1986), Zubaid and Davison (1987),
Corbet and Hill (1992), and Simmons (2005).
Kingston et al. (2001) noted that there were
two clearly defined phonic types of H. bicolor,
based on acoustic data collected in peninsular
Malaysia. One had a peak frequency of maximum
energy at ca. 131 kHz and the other at ca. 142 kHz.
Acta Chiropterologica, 12(2): 415–438, 2010
PL ISSN 1508-1109 © Museum and Institute of Zoology PAS
doi: 10.3161/150811010X537990
A taxonomic review of the Hipposideros bicolor species complex and H. pomona
(Chiroptera: Hipposideridae) in Thailand
BOUNSAVANE DOUANGBOUBPHA
1, 2, 5
, SARA BUMRUNGSRI
2
, PIPAT SOISOOK
3
, CHUTAMAS SATASOOK
3
,
N
IKKY M. THOMAS
4
, and PAUL J. J. BATES
4
1
Faculty of Environmental Sciences, National University of Laos, Dong Dok Campus, Xaythany District, Vientiane Capital, Lao PDR
2
Department of Biology, Faculty of Science, Prince of Songkla University, Hat Yai 90112, Thailand
3
Princess Maha Chakri Sirindhorn Natural History Museum, Faculty of Science, Prince of Songkla University, Hat Yai 90112, Thailand
4
Harrison Institute, Centre for Systematics and Biodiversity Research, Bowerwood House, St. Botolph’s Road, Sevenoaks,
Kent, TN13 3AQ, Great Britain
5
Corresponding author: E-mail: bounsavanhd@yahoo.com
Following extensive field work in Thailand (2006–2008), this paper reviews the taxonomic status of the three species of the
Hipposideros bicolor group. Based on morphometric characters and acoustic data, the two phonic types, H. bicolor (131 kHz) and
H. bicolor (142 kHz) are treated as distinct species. Hipposideros bicolor (131 kHz) is designated as H. bicolor; H. bicolor (142
kHz) is provisionally designated as H. atrox. The morphometric characters, acoustic data, and geographical distribution of
H. pomona are also reviewed. The diagnostic characters of these frequently confused taxa are discussed, with a detailed study of the
external, cranio-dental, and bacular morphology, and acoustic features. New data on the conservation status, distribution and ecology
of these three species are included.
Key words: Hipposideros bicolor group, H. atrox, H. pomona, taxonomy, distribution, Thailand
Acta Chiropterologica, 12(2): 439–448, 2010
PL ISSN 1508-1109 © Museum and Institute of Zoology PAS
doi: 10.3161/150811010X538007
Morphological evidence for hybridization in the sister species Myotis myotis and
Myotis oxygnathus (Chiroptera: Vespertilionidae) in the Carpathian Basin
JUSTYNA BACHANEK
1, 2
and TOMASZ POSTAWA
1
1
Institute of Systematics and Evolution of Animals, Polish Academy of Sciences, Sławkowska 17, 31-016 Kraków, Poland
2
Corresponding author: E-mail: jbachan@gmail.com
A recent molecular-based study suggested that two sister species of mouse-eared bats (Myotis myotis and M. oxygnathus) share some
mtDNA haplotypes when they occur in sympatry. We used traditional morphometric methods in order to find potential hybrid
specimens. Multivariate morphometric analysis of 22 cranial and dental characters was applied to a sample of 298 adult specimens
of mouse-eared bats (Myotis myotis and M. oxygnathus) from their sympatric range in the Carpathian Basin. Additionally, we
included several juvenile exemplars of both species in order to exclude the treatment of juvenile specimens as individuals with
intermediate characteristics. Principal Component Analyses (PCA) and Discriminant Function Analysis (DFA) revealed a distinct
separation between specimens of M. myotis and M. oxygnathus, with 6 specimens in an intermediate position. All intermediate
specimens come from the contact area of M. myotis and M. oxygnathus. A subsequent detailed analysis showed that intermediate
specimens are similar to M. myotis in skull and mandible size, but with tooth-row size similar to M. oxygnathus. An analogous
situation has been described in several species of small mammals. Bats designated as intermediate are probable hybrids, i.e. their
phenotypes lay between the parental forms.
Key words: Mouse-eared bat, cranial measurements, intermediate individuals, morphological analysis, Carpathian Basin
INTRODUCTION
Hybridization has been reported in several
groups of mammals, including bats, although exam-
ples from this group are very uncommon (Baker and
Bradley, 2006). The vast majority of cases were de-
tected using molecular methods in species with mor-
phological differences (Patton and Smith, 1990;
Bradley et al., 2003; Berthier et al., 2006) as well as
in cryptic species (Hoffmann et al., 2003; Baker and
Bradley, 2006). Some potential hybrids distin-
guished on the basis of morphometry (Herd and
Fenton, 1983) have not been confirmed by molecu-
lar studies (Weller et al., 2007). Morphological
study of individuals of two mouse subspecies and
two moles identified as hybrids suggested genetic
differentiation between them and the parental
species, according to both size (Auffray et al., 1996;
Loy et al., 2001) and shape (Renaud et al., 2009).
The mouse-eared myotis Myotis myotis (Bork hau -
sen, 1797) and the Monticelli’s myotis Myotis oxy -
gna thus (Monticelli, 1885) are a pair of closely re-
lated species occurring in Europe and Asia Minor.
Both have similar ranges in southern and central
Europe (Simmons, 2005; Dietz et al., 2007). The
genetic distance between them in mtDNA is small,
reaching 2.5–2.8% (Ruedi and Mayer, 2001), more-
over, Berthier et al. (2006) have shown that the
two species share some mtDNA haplotypes in
sympatric populations from the Alps. The results
of studies using nuclear (RAG2) and mitochon-
drial markers also indicate hybridization in the
Car pathian Basin (Bogdanowicz et al., 2009). This
sug gests incomplete reproductive isolation. In con-
trast to genetic distance measures and difficult to
recognize external features (Arlettaz, 1995), cranial
measurements provide enough precision to deter-
mine the correct species (Horáček, 1985; Ben-
da, 1994; Benda and Horáček, 1995). If M. myotis
and M. oxygnathus are relatively easily identifiable
on the basis of the skull, it should be possible to
identify hybrid specimens with intermediate cha-
racteristics as compared to the parental species. The
aim of this study was to conduct a morphological
analysis on Central European populations of M. my-
otis and M. oxygnathus in order to test hypotheses
on the occurrence of intermediate individuals, indi-
cating hybridization.
INTRODUCTION
The genus Myotis (Chiroptera: Vespertilionidae)
is widely distributed in America, Africa, Eurasia,
and Australia, and comprising between about 84
spe cies (Koopman, 1994) and up to 103 species
(Sim mons, 2005). The large-footed myotis, Myotis
ad versus (Horsfield, 1824), is considered to have
a wide distribution, ranging from Taiwan island,
Malay Peninsula, Sulawesi, Java and many islands
(Indonesia) to Western New Guinea and to the
Solomon Is lands, and Vanuatu (Simmons, 2005;
Wilson, 2008). Simmons (2005) recognized the fol -
lowing five subspecies as associated with adversus:
carimatae Miller 1906, orientis Hill 1983, taiwa-
nensis Ärnbäck-Christie-Linde 1908, tanimba ren-
sis Kitche ner 1995, and wetarensis Kitchener 1995.
Simmons (2005) also follows Kitchener et al.
(1995) in their treatment of two former subspecies
of M. adversus, namely macropus Gould 1854 and
mo luccarum Thomas 1915. Kitchener et al. (1995)
split M. adversus into three species based on cranial,
external and bacular morphology: M. adversus from
New South Wales and Indonesia, M. moluccarum
from northern Australia and New Guinea which
tentatively included solo monis, and M. macropus
from Victoria and South Australia. In addition, it
was also suggested that a specimen of M. adversus
from New South Wales may represent a distinct sub-
species, but it was not formally described (Kitchener
et al., 1995). Sub sequently, the molecular phylogeny
of three taxa (ad ver sus, macropus, moluccarum) in
Australia and Indo nesia was tested by mitochondri-
al genes, Cyt b and partial nicotinamide adenine din-
ucleotide dehydrogenase subunit 2 (ND2) (Cooper
et al., 2001). That study found evidence that only
one species (M. macropus) occurs in Australia and
most authors consider that adversus is not present in
Australia (Churchill, 2008; Richards et al., 2008) or
Papua New Guinea (Bonaccorso, 1998).
Taiwan’s taxon (Myotis taiwanensis) was first
described by Ärnbäck-Christie-Linde (1908) with
the type locality given as Takao, Anping, Tainan,
For mosa (= southern Taiwan). The species-level
clas sification was accepted by Tate (1941), but
Eller man and Morrison-Scott (1951) considered
Acta Chiropterologica, 12(2): 449–456, 2010
PL ISSN 1508-1109 © Museum and Institute of Zoology PAS
doi: 10.3161/150811010X538016
Revalidation of Myotis taiwanensis Ärnbäck-Christie-Linde 1908 and
its molecular relationship with M. adversus (Horsfield 1824)
(Vespertilionidae, Chiroptera)
NAI-JIAN HAN
1
, JIN-SHUO ZHANG
1
, TERRY REARDON
2
, LIANG-KONG LIN
3
, JUN-PENG ZHANG
4
, and SHU-YI ZHANG
4, 5
1
Institute of Zoology, and Graduate University of Chinese Academy of Sciences, 100101, Beijing, China
2
Evolutionary Biology Unit/Vertebrates, South Australian Museum, North Terrace, Adelaide SA 5000, Australia
3
Department of Life Science, Tunghai University, Taichung, 40704, Taiwan, China
4
School of Life Science, East China Normal University, 200062, Shanghai, China
5
Corresponding author: E-mail: syzhang@bio.ecnu.edu.cn
A combined approach using molecular and morphological data has revealed that the taxon Myotis taiwanensis, which until now has
been usually considered as a subspecies of Myotis adversus, should be reinstated as a full species, as originally described by
Ärnbäck-Christie-Linde (1908) from Takao, Anping, Tainan, Formosa (= southern Taiwan). In our genetic analysis using two nuclear
DNA segments of protein kinase C iota (PRKCI) and ubiquitin specific peptidase 9 (Usp9x), X-linked genes, together with
two mitochondrial genes, i.e., nicotinamide adenine dinucleotide dehydrogenase subunit 1 (ND1) and cytochrome b (Cyt b), we
demonstrate that M. taiwanensis is closely related to M. pilosus, and largely divergent from two subspecies of M. adversus. Our
analysis further shows that M. taiwanensis differs considerably from M. adversus in external and dental features. New records of
M. taiwanensis from Shandong and Anhui provinces in eastern China are presented.
Key words: Myotis adversus, M. taiwanensis, molecular phylogeny, morphology
INTRODUCTION
Compared to continental Europe and Britain,
Ireland has a relatively impoverished fauna, most
likely resulting from its recent glacial history, small
land area, island status and position at the north
western edge of the European land mass. Current-
ly, there are 34 terrestrial species of mammals in Ire -
land of which 10 are bats (Marnell et al., 2009).
Over the last 12 years Ireland’s species list has seen
the addition of three new bat species. In 1997, two
phonic types of Pipistrellus pipistrellus, were reclas-
sified to Pipistrellus pipistrellus (common pipi strel -
le) and P. pygmaeus (soprano pipi strel le) (Barratt et
al., 1997). Pipistrellus nathusii (Nathu sius’s pipi -
strel le) was confirmed to be breeding in Northern
Ireland in 1998 (Russ et al., 1998) and most recent-
ly a few suspected Myotis brandtii (Brandt’s bat)
have been found (Kelleher, 2005; Har ris 2006;
Mullen, 2006).
Myotis brandtii is distributed throughout central
and northern Europe, across Russia to the Urals,
being sparsely distributed in Western Europe, the
Mediter ranean and the Balkans (Dietz et al., 2009).
In Brit ain and mainland Europe, M. brandtii are
associated with woodland and habitats near water
(Harris and Yalden, 2008; Dietz et al., 2009), and
roost in crevices in buildings or trees (Dietz et
al., 2009). Myotis brandtii is a cryptic species, diffi-
cult to separate morphologically from Myotis
mystacinus (whisker ed bats), which are found across
Europe including Ireland; Myotis alcathoe (Alca -
thoe’s bat) and Myotis aurascens (steppe whiskered
bat) which are found in continental Europe (Dietz et
al., 2009). Separating M. mystacinus (whiskered
bat) and M. brandtii based on morphology alone
requires the use of unreliable characters such as
the premolars (Berge, 2007) and penis shape (Harris
and Yalden, 2008; Dietz et al., 2009). Genet-
ically, these two species are quite distinct, with
M. brandtii grouping with the New World Myotis,
rather than the Old World Myotis which is typical
of all other European myotid species (Ruedi and
Mayer, 2001).
Acta Chiropterologica, 12(2): 457–461, 2010
PL ISSN 1508-1109 © Museum and Institute of Zoology PAS
doi: 10.3161/150811010X538025
The status of the cryptic bat species, Myotis mystacinus and Myotis brandtii
in Ireland
EMMA S. M. BOSTON
1
, DANIEL J. BUCKLEY
1
, MICHAËL BEKAERT
1
, YANN GAGER
1
, MATHIEU G. LUNDY
2
,
D
AVID D. SCOTT
2
, PAULO A. PRODÖHL
2
, W. IAN MONTGOMERY
2
, FERDIA MARNELL
3
,
and E
MMA C. TEELING
1, 4
1
Centre for Irish Bat Research, University College Dublin, School of Biology and Environmental Science,
Belfield, Dublin 4, Ireland
2
Centre for Irish Bat Research, Queen’s University Belfast, Medical and Biological Centre, 97 Lisburn Road,
Belfast, Northern Ireland, United Kingdom
3
National Parks and Wildlife Service, 7 Ely Place, Dublin 2, Ireland
4
Corresponding author: E-mail: emma.teeling@ucd.ie
The recent identification of Myotis brandtii in Ireland raised the possibility that many roosts previously identified as M. mystacinus
had the potential of being misidentified M. brandtii. Thus, the distribution and population estimates for M. mystacinus may have
been over-estimated, while M. brandtii may have been under-estimated. Results from an all Ireland genetic survey of known
M. mystacinus maternity roosts confirm that no long term misidentification has taken place. All specimens caught and sampled were
M. mystacinus. Additonally, no further records of M. brandtii were found during six nights of woodland trapping using the acoustic
lure. While the status of M. mystacinus in Ireland is now listed as ‘least concern’ in the Irish Red List, M. brandtii is listed as ‘data
deficient’ and cannot currently be considered a resident species
Key words: Vespertilionidae, Myotis, cryptic species, Irish mammals
INTRODUCTION
The species Vespertilio murinus Linnaeus, 1758
is widespread in the Palaearctic region, ranging
from eastern France and Switzerland northward to
Scandinavia north to 61ºN, south of 55–60ºN in
Russia, eastward to the Ussuri region of eastern
Siberia, Mongolia, north-east China and Korea (e.g.,
Baagøe, 1999, 2001; Simmons, 2005). This species
is considered polytypic, with two recognized sub -
species, V. m. murinus (Europe to Upper Amur) and
V. m ussuriensis Wallin, 1969 (eastern Siberia, Us -
suri region, possibly north east China and Korea)
(Koopman, 1994; Rydell and Baagøe, 1994; Tiunov,
1997; Simmons, 2005). Banding studies have shown
that individuals of V. murinus migrate long distances
between their summer and winter roosts (Strelkov,
1969). The longest recorded flight is 1,440 km in
a young male which was banded at the end of July
1988 in Estonia, and discovered the same year in
November in Austria (Masing, 1989). Numerous va-
grants are found far outside their normal range, as
far as the Faroe islands (Baagøe and Bloch, 1994;
Baagøe, 1999, 2001). There are several records from
Far East Russia (Wallin, 1969). However, prior to
the first record on Rebun Island near Hokkaido
(Satô and Maeda, 2003), V. murinus was not consid-
ered to be present in Japan.
Within the Japanese archipelago, there have been
several reports of movements in bats, in most cases
involving Vespertilio sinensis (= Vespertilio super-
ans Thomas, 1899 — see Horáček, 1997). Banding
studies in this species indicate that it is capable of
long distance movements, ranging up to 780 km
(Mukohyama, 2002; see also Kimura et al., 2003).
Miniopterus fuliginosus, another Japanese species,
is known to fly distances up to 200 km (Xu et al.,
2005). However, neither of these taxa have been
recorded crossing the ocean. In contrast, one indi-
vidual of Nyctalus aviator was recaptured 140 km
from the banding site in Hokkaido across the Tsu -
garu Strait to Aomori, Honshu (M. Mukohyama,
personal communication). Nevertheless, there are
no records of movement of bats between Japan and
neighboring countries.
Between 17 and 27 December 2005, three bats of
indeterminate identity were captured in northern
Japan. Their forearm lengths were slightly smaller
than the average for V. sinensis, raising the pos -
sibility that they represented V. murinus. If these
Acta Chiropterologica, 12(2): 463–470, 2010
PL ISSN 1508-1109 © Museum and Institute of Zoology PAS
doi: 10.3161/150811010X538034
Vespertilio murinus Linnaeus, 1758 confirmed in Japan from morphology
and mitochondrial DNA
KUNIKO KAWAI
1, 2, 7
, DAI FUKUI
3
, MASAHIKO SAT Ô
4
, MASASHI HARADA
5
, and KISHIO MAEDA
6
1
Field Science Center for Northern Biosphere, Hokkaido University, Sapporo, Hokkaido, Japan
2
Division of Mammals, National Museum of Natural History, Smithsonian Institution, Washington, D.C., USA
3
Hokkaido Research Center, Forestry and Forest Products Research Institute, Sapporo, Hokkaido, Japan
4
Rishiri Town Museum, Senhoshi, Rishiri Island, Hokkaido, Japan
5
Laboratory Animal Center, Osaka City University, Graduate School of Medicine, Abeno, Osaka, Japan
6
Center for Natural Environment Education, Nara University of Education, Takabatake, Nara-City, Japan
7
Corresponding author: E-mail: kkawai@fsc.hokudai.ac.jp
In December 2005, three bats of indeterminate identity were captured in northern Japan. Their forearm lengths were slightly smaller
than the average for Vespertilio sinensis. We identified these three individuals as V. murinus using both morphological and molecular
approaches, and discussed their possible migration or vagrancy routes. Prior to the first record of V. murinus on Rebun Island near
Hokkaido in 2002, there was no record of this species in Japan. These new occurrences illustrate the importance of determining
possible migration or vagrancy routes of bats for understanding and preventing the spread of zoonotic diseases.
Key words: Vespertilio murinus, mtDNA, cytochrome-b, ND1, migration, occurrence, Japan
INTRODUCTION
Our understanding of the distribution and diver-
sity of bats has advanced significantly in recent
years through a combination of more intensive study
and the application of molecular markers and DNA
sequencing to enable the identification of morpho-
logically cryptic species. New species have been de-
scribed and ranges have been redefined, not only in
the world’s biologically rich, but less well studied
tropical regions (e.g., Esselstyn et al., 2008; Baker
et al., 2009), but also in intensively studied temper-
ate regions. In Europe, a number of new bat spe-
cies have been described recently: e.g., Pipistrel-
lus pygma eus (Barratt et al., 1997) and several new
Ple cotus species (e.g., Spitzenberger et al., 2006).
The biology, distribution and conservation status of
these new species are still being investigated, but
despite their morphological similarity, some are
ecologically distinct from their near relatives and
are likely to have different conservation require-
ments. One of those new species is Myotis alcathoe,
Al ca thoe bat (Helversen et al., 2001), a small mem-
ber of the genus described from specimens caught in
northern Greece and Hungary. Since its discovery it
has been reported from Poland, Slovakia and Ro -
mania in the east to Spain, France and Switzerland
in the west (Ruedi et al., 2002; Niermann et al.,
2007). The known distribution remains patchy, but
it is likely to be widely distributed across Europe
and new records are reported regularly. Numerous
records in western France suggest that M. alcathoe
could also be present in the UK and perhaps even
Ireland. We report the discovery of M. alcathoe at
two autumn swarming sites in southern England and
a third site in northern England, based on micro -
satellite genotyping and mitochondrial sequencing
of genomic DNA obtained from wing biopsy sam-
ples collected between 2003 and 2009.
M
ATERIALS AND METHODS
Study Sites and Sampling
As part of ongoing studies of the ecology and population bi-
ology of swarming species, we caught bats at sites across
Europe (Fig. 1a) primarily between 2007 and 2009 and have
been catching at sites in the UK for over 10 years. Morphol -
ogical measurements were taken together with 3 mm biopsies of
Acta Chiropterologica, 12(2): 471–483, 2010
PL ISSN 1508-1109 © Museum and Institute of Zoology PAS
doi: 10.3161/150811010X538043
Myotis alcathoe confirmed in the UK from mitochondrial and microsatellite DNA
CAMILLE M. I. JAN
1, 3
, KATIE FRITH
2, 3
, ANITA M. GLOVER
1
, ROGER K. BUTLIN
2
, CHRIS D. SCOTT
1
,
F
RANK GREENAWAY
1
, MANUEL RUEDI
4
, ALAIN C. FRANTZ
3
, DEBORAH A. DAWSON
3
, and JOHN D. ALTRINGHAM
1, 5
1
IICB, Faculty of Biological Sciences, University of Leeds, Leeds, LS2 9JT, United Kingdom
2
Animal and Plant Sciences, University of Sheffield, Sheffield, S10 2TN, United Kingdom
3
NERC Biomolecular Analysis Facility, Department of Animal and Plant Sciences, University of Sheffield, Sheffield,
S10 2TN, United Kingdom
4
Muséum d’Histoire Naturelle, 1 route de Malagnou, CH-1208 Genève, Suisse
5
Corresponding author: E-mail: j.d.altringham@leeds.ac.uk
We describe the recent discovery of Alcathoe bat, Myotis alcathoe, at locations in the south and north of England. First described in
2001 from individuals caught in Greece, the species’ known range has steadily been extended across much of Europe. Its presence
in the UK was confirmed by mtDNA (ND1) and microsatellite analysis of wing biopsy samples collected between 2003–2009,
primarily at swarming sites. Morphological and echolocation call data are consistent with the genetic interpretation. The presence
of M. alcathoe at sites 350 km apart, and the probability that it makes up a significant proportion of the small Myotis bats at these
sites, suggest it is a resident species. Preliminary assessment of its distribution in the UK will be facilitated by its distinctive
echolocation call, an FM sweep that typically terminates at 43–46 kHz, above those of other Myotis species.
Key words: Myotis alcathoe, UK, mtDNA, microsatellite genotyping, swarming
... Our results demonstrate an association between wind turbine height and fatality counts of some species of bats and swallows, based on a large dataset of turbines extending 119 to 186 m into the aerosphere. Our findings are consistent with previous studies showing that turbine height explains a portion of bat fatalities at wind energy facilities (Johnson et al. 2003;Fiedler et al. 2007;Barclay et al. 2007;Rydell et al. 2010;Georgiakakis et al. 2012;Huso et al. 2021), although other factors, such as the proximity of wind energy facilities to key habitats, migration corridors, and roosts, also play a role (Barclay et al. 2007;Cryan and Brown 2007;Ferreira et al. 2015;Thompson et al. 2017;Davy et al. 2020). Greater uncertainty in the relationship between turbine height and fatalities that we observed in spring could be the result of fewer carcass observations in Table 2. Back-transformed model predicted estimates (and 95% confidence intervals) of the number of bats killed per turbine (M) at the shortest and tallest turbines in our study. ...
... Broadly, our results and those of other studies (Johnson et al. 2003;Fiedler et al. 2007;Barclay et al. 2007;Rydell et al. 2010;Georgiakakis et al. 2012) indicate a general, ongoing pattern of more bat fatalities at taller turbines. While more bat fatalities at taller turbines is a concern with the increasing size of turbines (Wiser et al. 2011(Wiser et al. , 2018, fatality rates of bats and birds appear to be relatively consistent per unit of energy produced (GWh) by a turbine (Huso et al. 2021). ...
Article
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Understanding the relationship between the height of wind turbines and wildlife fatalities is important for informing and mitigating wildlife collisions as ever taller and denser arrays of wind turbines are erected across the landscape. We examined relationships between turbine height and fatalities of bats and swallows at 811 turbines in Ontario, Canada, ranging from 119 to 186 m tall. We accounted for cut-in speeds, operational mitigation, and taller turbines projecting carcasses farther from the turbine base than shorter turbines. Fatalities of hoary bats (Lasiurus cinereus Palisot de Beauvois, 1796), silver-haired bats (Lasionycteris noctivagans Le Conte, 1831), and big brown bats (Eptesicus fuscus Palisot de Beauvois, 1796) increased with increased maximum blade height of turbines. In contrast, fatalities of little brown bat (Myotis lucifugus Le Conte, 1831) and eastern red bat (Lasiurus borealis Müller, 1776) decreased with increased turbine height. Fatalities of purple martins (Progne subis Linnaeus, 1758) and tree swallows (Tachycineta bicolor Vieillot, 1808) were higher at taller turbines than shorter turbines. However, fatalities of cliff swallow (Petrochelidon pyrrhonota Vieillot, 1817) and barn swallow (Hirundo rustica Linnaeus, 1758) were not associated with turbine height. Our results suggest that varying flight height among species may be one factor affecting collision risk.
... El impacto de la energía eólica en murciélagos está ampliamente constatado a escala mundial (Kunz et al. 2007, Arnett et al. 2015, O'Shea et al. 2016, habiéndose invertido cantidades desiguales en su investigación, sobre todo en la aplicación de medidas de mitigación, con grandes diferencias entre países y promotores, especialmente Estados Unidos frente al resto del mundo (Baerwald et al. 2009, Arnett et al. 2013, Department of Energy 2019. En Europa, destacan un mayor número de estudios en la Región Eurosiberiana, especialmente Alemania (Rydell et al. 2010, Korner-Nievergelt et al. 2013, Lehnert et al. 2014, mientras que en la Mediterránea, y salvo Portugal (ICNB 2010, Amorim et al. 2012, Santos et al. 2013, Ferreira et al. 2015, Peste et al. 2015, Silva et al. 2017, no se ha pasado de un análisis más puntual. En Italia (Ferri et al. 2011, Roscioni et al. 2013, Croacia (Zagmajster et al. 2007), Grecia (Georgiakakis et al. 2012) e incluso España (Alcalde & Sáenz 2004, de la cruz Sánchez et al. 2020, Muñoz & Farfán 2020, mayoritariamente se recogen listados de número de individuos y especies afectadas sin entrar en cuestiones metodológicas (a pesar del seguimiento post-construcción más continuado que se desarrolla en España). ...
... En Canadá los valores más elevados son de 23-24 murciélagos/ae/año (Zimmerling & Francis 2016). En Europa, en Alemania, Black Forest) 41 murciélagos/ae/año (Rydell et al. 2010), y en Estados Unidos hasta 69,6 murciélagos/ae/año (Arnett et al. 2008) El principal estimador utilizado por la consultora a lo largo del año (Winkelman 1989) está obsoleto y presenta grandes sesgos (Abad 2012), pero es que la consultora sólo estimó la mortalidad para las aves. Estudios como Smallwood (2020) y referencias en dicho trabajo, sugieren una mayor siniestralidad de quirópteros que aves. ...
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Este trabajo analiza la siniestralidad de murciélagos (27,09 ae/año) y aves (10,09 ae/ año) en dos parques eólicos de cinco y seis aerogeneradores en Aragón a lo largo de un ciclo anual en el Valle del Ebro. Comparamos los resultados de este estudio con los del promotor, profundizando en los errores de detectabilidad y tasa de permanencia asociados que afectan a la estima de mortalidad. Se identificaron cinco especies Pipistrellus pipistrellus, Pipistrellus pygmaeus y Pipistrellus kuhlii, Eptesicus serotinus e Hypsugo savii que colisionaron principalmente entre junio y octubre. La mortalidad anual de quirópteros se estimó entre 388-5.460 ejemplares al año, una de las mayores mortalidades detectadas a nivel mundial. A pesar de ser especies comunes de distribución amplia el impacto cuantitativo no es asumible. Con las conclusiones obtenidas del Plan de Vigilancia Ambiental (PVA) no se evaluaron correctamente los impactos que pasaron desapercibidos por la utilización errónea de la metodología de seguimiento referida a la frecuencia de visitas o área de búsqueda, y su análisis posterior de las tasas de predación, eficiencia del observador y estima de la mortalidad. Proponemos un programa de mitigación basado en el retraso de la velocidad de arranque de los aerogeneradores hasta los 6 m/s en ese período que reduciría la siniestralidad de quirópteros observada en torno al 46-54%. Además, es urgente por el Gobierno de Aragón revisar el protocolo de seguimiento de parques eólicos en lo referente a su duración, frecuencia de visitas y errores mencionados anteriormente, así como las Declaraciones de Impacto Ambiental. Los promotores y sus consultoras deben demostrar con análisis robustos las conclusiones de sus trabajos en vez de limitarse a trabajos descriptivos. Los parques también afectaron a las aves, especialmente rapaces como el buitre leonado Gyps fulvus o el cernícalo primilla Falco naumanni, los aláudidos e incluso grullas (Grus grus). En estas especies, las colisiones se asociaron a aspectos concretos de la ecología de estas especies como la presencia de carroña, el hábitat de cultivo o la presencia en época de migración e invernada.
... There are multiple scenarios envisaged to address the gap in energy production, and the global climate crisis will limit the contribution of fossil fuels. The growing concern over the impact of wind turbines and solar panels on the local environment and public acceptance provides further motivation for the search for other sources of clean energy [5]. ...
Conference Paper
The growing energy demand and the global warming crisis require increasing the share of sustainable energy sources. Fusion energy has the potential to provide a zero-carbon, low-waste energy source that revolutionizes the energy industry. A gram of hydrogen isotope as fusion fuel can give equal energy to burning 16 m3 of coal. In a novel approach to producing fusion energy, the Orbitron concept has been introduced in which ions are electrostatically confined around a high voltage (H.V.) cathode. In this method, the cathode voltage should be as high as hundreds of kVs. One of the major engineering challenges lies in the transmission of such voltage levels from the atmospheric pressure to a high vacuum environment (p< 1e-8 Torr). One should be concerned not only about the breakdown of the vacuum or dielectric but also about the surface flashover across the insulator's surface. This study proposes an innovative design for a compact, 300-kV bushing to achieve the aforementioned goal.
... Although there is still no evaluation of the effect of modifying wind turbine distance to woody edges on the reduction of bat fatality risk (Berthinussen et al., 2021), the EUROBATS guideline was evidently based on the best available science and formulated as a precautionary principle. Moreover, three studies show that the increase of tree cover around wind turbines and the distance to woody habitats are ones of the main factors of collision risks in Europe (Roemer et al., 2019;Rydell et al., 2010;Santos et al., 2013). In addition, a recent study highlighted that wind turbines no longer attract bats beyond a distance of at least about a 100 m from woody edges (Leroux et al., 2022). ...
Article
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Wind power generation has grown exponentially over the past 20 years to meet international goals of increasing the share of renewables in energy production. Yet, this process has too often been conducted at the cost of airborne biodiversity such as birds and bats. The latter are severely threatened due to deaths by collision at wind turbine. The UNEP/EUROBATS agreement that came into force in 1994 is now ratified by 37 countries; since 2008, it recommends to site wind turbines at least 200 m away from woody edges to decrease bat fatality risks. However, 14 years later we still do not know to what extent this international recommendation has been applied in Europe. We assessed siting distances between woody edges and wind turbines for the largest wind energy producers among the UNEP/EUROBATS parties: the UK, Germany, and France. We show that 61%, 78%, and 56%, respectively, of the installed wind turbines did not comply with UNEP/EUROBATS guidelines, without improvement over time. We identified probable causes of these findings and provided key policy recommendations to achieve compliance to UNEP/EUROBATS guidelines such as better: (i) inclusion in regulatory texts, (ii) notification of the environmental authorities, and (iii) strategic, well-informed, forward planning of areas suitable for wind turbine development.
... For raptors, such as golden eagles (Aquila chrysaetos) and white-tailed eagles (Haliaeetus albicilla), there are relatively few collisions compared to other bird species, but this small impact compounded with other stressors, such as electrocution with power lines, can affect population sizes (May, 2015;Allison et al., 2017). Migratory bats, including hoary bats (Lasiurus cinereus) and noctule bats (Nyctalus noctula), often comprise the highest proportion of fatalities at wind farms (Arnett and Baerwald, 2013;Rydell et al., 2010). Although population data for these species are limited, it is possible that the current fatality rate is unsustainable (Friedenberg and Frick, 2021). ...
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Non-technical summary A substantial increase in wind energy deployment worldwide is required to help achieve international targets for decreasing global carbon emissions and limiting the impacts of climate change. In response to global concerns regarding the environmental effects of wind energy, the International Energy Agency Wind Technical Collaborative Program initiated Task 34 – Working Together to Resolve Environmental Effects of Wind Energy or WREN. As part of WREN, this study performed an international assessment with the global wind energy and environmental community to determine priority environmental issues over the next 5‒10 years and help support collaborative interactions among researchers, developers, regulators, and stakeholders. Technical summary A systematic assessment was performed using feedback from the international community to identify priority environmental issues for land-based and offshore wind energy development. Given the global nature of wind energy development, feedback was of interest from all countries where such development is underway or planned to help meet United Nations Intergovernmental Panel on Climate Change targets. The assessment prioritized environmental issues over the next 5–10 years associated with wind energy development and received a total of 294 responses from 28 countries. For land-based wind, the highest-ranked issues included turbine collision risk for volant species (birds and bats), cumulative effects on species and ecosystems, and indirect effects such as avoidance and displacement. For offshore wind, the highest-ranked issues included cumulative effects, turbine collision risk, underwater noise (e.g. marine mammals and fish), and displacement. Emerging considerations for these priorities include potential application to future technologies (e.g. larger turbines and floating turbines), new stressors and species in frontier regions, and cumulative effects for multiple projects at a regional scale. For both land-based and offshore wind, effectiveness of minimization measures (e.g. detection and deterrence technologies) and costs for monitoring, minimization, and mitigation were identified as overarching challenges. Social media summary Turbine collisions and cumulative effects among the international environmental priorities for wind energy development.
... Offshore wind energy production is increasing worldwide, specifically in north-western Europe (Global Wind Energy Council, 2021;Kaldellis & Kapsali, 2013). For the onshore sector, it is known that large numbers of bats die at wind turbines, with migratory species being most vulnerable (O'Shea et al., 2016;Rydell et al., 2010;Voigt et al., 2015). For the offshore sector, it was suggested that wind energy production may pose a threat to bats as well (Ahlén et al., 2009;Gaultier et al., 2020;Hüppop et al., 2019;Rydell et al., 2014), since past acoustic surveys documented the presence of bats above marine waterbodies (Ahlén et al., 2009;Brabant et al., 2019;Cryan & Brown, 2007;Gaultier et al., 2020;Hüppop & Hill, 2016;Ijäs et al., 2017;Lagerveld et al., 2021;Peterson et al., 2016;Rydell & Wickman, 2015;Rydell et al., 2014, Seebens-Hoyer et al., 2021. ...
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The wind energy-bat conflict is well documented for the onshore sector, with high numbers of casualties, specifically for migratory bat species. Offshore wind turbines might be a threat to bats as well, yet offshore bat migration is poorly documented. Accordingly, potential conflicts between bat conservation and offshore wind energy production are difficult to evaluate. Here, we used automated radio-telemetry to track 50 km continuous offshore movements of two Nathusius' pipistrelles (Pipistrellus nathusii) within the Motus network. After crossing the marine waterbody, tagged bats traveled over several hundred kilometers along the coastline from Germany towards the Netherlands and Belgium. Our study highlights the possibility for migratory bats to collide with offshore and coastal wind turbines. Therefore, we plead for implementing pre- and post-construction surveys and adequate mitigation schemes at offshore wind turbines in sensitive areas of the North and Baltic Sea if not already practised.
... Wind turbine collisions do not appear to be random incidents. Bird and bat species are likely drawn to turbines either directly because turbines resemble roosting locations (Cryan et al., 2014) or indirectly because turbines attract insects that birds and bats feed on (Rydell et al., 2010a(Rydell et al., , 2010b. Nine bird individual species were collision victims due to the KFW wind farm in our study. ...
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... Las cuevas que utilizan muchas especies de murciélagos son comúnmente usadas con fines turísticos, o son quemadas con el único objetivo de eliminar a los murciélagos (Mickleburgh et al., 2002). Recientemente los campos de aerogeneradores en diferentes partes del mundo matan diferentes especies de murciélagos que prefieren volar en campos abiertos (Rydell et al., 2010). ...
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