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Mammalia 75 (2011): 97–101 2011 by Walter de Gruyter •Berlin •New York. DOI 10.1515/MAMM.2010.074
2010/063
Article in press - uncorrected proof
Short Note
Nyctalus lasiopterus Schreber, 1780 (Chiroptera:
Vespertilionidae): first record for Kornati archipelago
and first recent capture for Croatia
Dina Kovacˇ
1,2,
*, Daniela Hamidovic´
2
,
Norma Fressel
1,2
and Sanja Drakulic´
1
1
Biology Students Association – ‘‘BIUS’’, Department
of Biology, Faculty of Science, University of Zagreb,
Rooseveltov trg 6, 10000 Zagreb, Croatia,
e-mail: dinakovac@gmail.com
2
Croatian Biospeleological Society, Demetrova 1, 10000
Zagreb, Croatia
*Corresponding author
Keywords: bats; Croatia; Falco eleonorae; Kornati
archipelago; Nyctalus lasiopterus.
Chiropterofauna in Croatia consists of 34 recorded bat spe-
cies that belong to 11 genera. However, the exact distribution
of the majority of bat species is poorly known owing to
insufficient data. One more species, Mehely’s horseshoe bat
(Rhinolophus mehelyi), has not been confirmed in recent
studies and it is considered to be regionally extinct (RE)
(Antolovic´ et al. 2006, Ministry of Culture and State Institute
for Nature Protection 2010).
Nyctalus lasiopterus is considered to be the largest and
one of the rarest bat species in Europe (Popa-Lisseanu et al.
2008). Its distribution in Europe is mainly Mediterranean,
stretching from the Iberian Peninsula to the Balkans, with
records to the north being very scattered (Iba´n˜ez et al. 2004).
On the Global and Mediterranean IUCN Red List of Mam-
mals it is considered to be in the ‘‘near threatened’’ category
(Hutson et al. 2008, Temple and Cuttelod 2009), whereas on
the European and Croatian Red List of Mammals it is listed
as ‘‘data deficient’’ (Antolovic´ et al. 2006, Temple and Terry
2007).
There are only a few records of Nyctalus lasiopterus in
Croatia, most of which are historical. Kolombatovic´ (1884)
caught a female of the species near the town of Solin in
March 1883 and reported it as Vesperugo molossus.He
reported two more records in the following two years: one
specimen captured in the Neretva valley and one male near
the town of Solin (Kolombatovic´ 1885). Wettstein (1928)
gave body and skull measurements of a dead male referred
as Nyctalus maximus (synonym of N. lasiopterus) collected
in June 1926 at 600 m altitude near the town of Senj. Over
a period of 69 years, skeletal remains of only 16 specimens
were found in the Galicˇnjak cave on the island of Mljet
(Tvrtkovic´ and Baltic´ 1996). All of these records are from a
330-km stretch of the Eastern Adriatic Coast (Figure 1).
Within Croatia, the bat fauna of Kornati archipelago has
not been well-documented and no study on bats of Kornati
islands has been published. Most of the underground sites
on these islands have yet to be fully explored. The only
existing data for surrounding area are those from Telasˇc´ica
Nature Park on the island of Dugi otok, a few kilometers
away from Kornati archipelago where eight species of bats
have been recorded to date: Hypsugo savii,Myotis emargi-
natus,Myotis blythii oxygnathus,Pipistrellus kuhlii,Plecotus
austriacus,Rhinolophus ferrumequinum,Rhinolophus hip-
posideros and Tadarida teniotis (Grgurev 2002). The Bat
Research Group of the Biology Students Association –
‘‘BIUS’’ conducted the first field research of the bat fauna
in the National Park ‘‘Kornati’’, Croatia during 2009. The
aim was to determine the composition of bat fauna and to
form a basic guideline for future monitoring protocol for the
National Park. These results are the first official records on
bat fauna in the National Park ‘‘Kornati’’. Here, we report
the first record of the greater noctule, Nyctalus lasiopterus
as the first record for Kornati archipelago and the first cap-
ture of a live individual in Croatia for 83 years.
The Kornati archipelago is a special and separate group
of islands located in the Adriatic Sea that encompasses an
area of approximately 300 km
2
that includes around 140 land
units permanently or temporarily above sea level. In 1980,
part of it was designated as the Kornati National park (Fri-
ganovic´ 1996). The islands of Kornat (32.5 km
2
) and Z
ˇut
(14.8 km
2
) account for 70% of the total land area. The basic
geomorphological characteristic of the archipelago is pre-
dominantly karstic relief with a highly indented coastline
(Bognar and Grizelj 1996).
According to Koeppen’s climate classification system, the
area of Kornati National Park presents an olive climate (Csa).
The monthly air temperature ranges from an average 88Cin
February to an average 258C in July and August and the
maximum rainfall recorded in October (Penzar and Penzar
1996). The entire area belongs to the EU-Mediterranean veg-
etation zone where the larger islands are covered by karstic
pasture vegetation as a dominant habitat type, with small
areas of olive groves (Olea europaea) and woodland on larg-
er islands. The basic climate-zonal woodland vegetation is
represented by the Myrto-Quercetum ilicis, whereas the rock
vegetation is represented by the Phagnalo-Centaureetum
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98 D. Kovacˇ et al.: First record and recent capture of Nyctalus lasiopterus
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Figure 1 Circled on the map of Croatia is the location of the Kornati archipelago. The icon of a black bat represents new data on Nyctalus
lasiopterus from the Kornati archipelago and recent data from the island of Mljet (Tvrtkovic´ and Baltic´ 1996). The icon of a white bat
represents historical data (Kolombatovic´ 1884, Kolombatovic´ 1885, Wettstein 1928). The Falco eleonorae icon represents nesting areas
(Piasevoli and S
ˇc´ etaric´ 2001, Radovic´ et al. 2003, S
ˇc´ etaric´-Legan and Piasevoli 2005, G. Piasevoli personal communication). The circled
F. eleonorae icon represents potential nesting sites. Nature Parks are marked with dark grey and National Parks with light grey color. The
map was constructed in the SAGA free geographic information system (GIS).
ragusinae, and the coastal cliff vegetation by the Plantagini-
Limonietum cancellati community (Trinajstic´ 1996).
During the hot spring and summer months, bats can visit
a few available freshwater sources for foraging and drinking
of water. There is only one pond that contains water through-
out the year and it is located on the island of Kornat, in the
Tarac field (43849.7639,15816.0509) surrounded with olive
groves. Other water bodies are extremely rare and dependant
on weather conditions. There are also some manmade water-
ing sites for sheep but most of these are smaller than 0.5 m
2
.
We conducted sampling sessions by mist netting around
the pond in Tarac field and watering site near Lovric´ev stan
on the island of Kornat in May, August and September 2009,
for two nights each month. We used six mist nets (two
1.0=2.5 m, one 9.0=2.5 m, two 6.0=2.5 m and one
3.0=2.5 m) surrounding the water bodies to capture bats.
Mist nets were set up at sunset and remained opened for a
minimum of 6 h, often until sunrise and were checked every
15 min. Captured individuals were weighted with the Pesola
spring balance (60 g) and forearm length was measured with
calliper Meba ("0.1 mm). Bats were aged, sexed, identified
following Dietz and von Helversen (2004) and released. One
individual was photographed and the photographs are avail-
able on request.
During our research of Kornati archipelago, we recorded
10 bat species: Hypsugo savii, Miniopterus schreibersii,
Myotis mystacinus, Myotis emarginatus, Myotis blythii oxy-
gnathus, Nyctalus lasiopterus, Pipistrellus kuhlii, Plecotus
kolombatovici, Rhinolophus ferrumequinum and Tadarida
teniotis (Kovacˇ and Fressel in press).
We captured three adult males of greater noctule bat
(Nyctalus lasiopterus) in the Tarac field. The first one was
captured at 02:15 h on August 22, 2009 (forearm length:
63.6 mm, weight: )60.0 g), the second at 22:10 h on Sep-
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D. Kovacˇ et al.: First record and recent capture of Nyctalus lasiopterus 99
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tember 20, 2009 (forearm length: 63.3 mm, weight: )60.0 g)
and the third one at 22:30 h on September 21, 2009 (forearm
length: 64.0 mm, weight: )60.0 g). All three males had
enlarged and conspicuous testes, as well as buccal glands.
For a tree-dwelling bat such as Nyctalus lasiopterus, the
availability of old trees with appropriate cavities for roosting
is an important factor (Esto´k et al. 2007). The largest com-
plex of holm oak (Quercus ilex) forests is in its degradation
stages with trees around 6 m high only being found sporad-
ically (Matic´ et al. 2001). The same situation exists in the
olive groves with trees not higher than 6 m. Studies of this
species in Corsica have shown they mostly roost (96%) in
Corsican pines (Pinus nigra subsp. laricio) and some Euro-
pean silver fir (Abies alba) (Beuneux et al. 2010). It is pos-
sible that the roost trees on Kornati are aleppo pines (Pinus
halepensis) because trees higher than 11 m can be found on
the archipelago. There is also a large forest complex of P.
halepensis in Telasˇc´ica Nature Park where the trees are
approximately 8 m high (Matic´ et al. 2001). Another tree
species that can be found on the archipelago is the stone pine
(Pinus pinea), but there are no data about older trees and
their height. Because there are few roosting opportunities on
Kornati archipelago, the presence of N. lasiopterus suggests
that some other resource might be important enough to
attract the species.
Nyctalus lasiopterus is known as a predator which regu-
larly preys on migrating passerines, as well as on insects
(Dondini and Vergari 2000, Iba´n˜ez et al. 2001). Its presence
on the island of Kornat could possibly be explained by an
abundant food source. As proposed by Kralj et al. (2007),
the Croatian coast represents a crossroad for migratory song-
birds (warblers) using southwestern, southeastern and central
Mediterranean flyways. There are few bigger wetlands in the
eastern Adriatic coast such as Lake Vransko and Kolansko
Blato, which represent important stopover sites for many
migratory birds (Kralj et al. 2007). Some of the migratory
passerines (Saxicola rubetra,Sylvia atricapilla,Sylvia borin
and Phylloscopus trochilus) were recorded in the study area
at the same period as our research (S
ˇupraha in press). This
can be further linked with the suggestion that distribution of
greater noctule closely follows the migration paths and high
abundance of passerine birds. As a nocturnal ‘‘ecological
equivalent’’ of N. lasiopterus, Eleonora’s Falcon, Falco eleo-
norae, catches prey in the air during the autumn migration
from Europe to Africa, and switches its diet from mostly
large insects to catching migrating birds (Ristow et al. 1986,
Popa-Lisseanu et al. 2007). A few sightings of F. eleonorae
were recorded in the Kornati archipelago (Lukacˇ et al. 1997),
and nests were confirmed on cliffs for islands distant from
the coastline (Figure 1, Piasevoli and S
ˇc´etaric´ 2001, S
ˇc´etaric´-
Legan and Piasevoli 2005, G. Piasevoli personal communi-
cation). Because both species occur on the Kornati
archipelago, there is a possibility that N. lasiopterus uses a
similar hunting strategy and catches prey above the open sea
similar to F. eleonorae (Rose´n et al. 1999). The distance
between the island of Kornati and the mainland where more
suitable roosting sites are available is 20 km and therefore
is not an obstacle for species such as N. lasiopterus which
can fly more than 130 km in a single night (Popa-Lisseanu
et al. 2009). This could explain its presence near the pond
in Tarac field at the Kornat island. To test this hypothesis
further insight into species dietary preferences would be
needed from the Kornati archipelago, as well as in areas
around F. eleonorae nesting sites and surrounding islands
(Figure 1).
Another issue concerns the fact that all captured individ-
uals were adult reproductively active males, similar to studies
of Nyctalus lasiopterus in France where throughout the year
only males are recorded on the island of Corsica (Beuneux
et al. 2010). In Loze`re, France, no females were caught dur-
ing the breeding period in the summer but were recorded
along with males in the autumn migration and mating period
(Destre 2008, Sane´ 2008). Sexual segregation of the popu-
lation most probably results from sex-biased migration (Iba´-
n˜ez et al. 2009). The presence of females in Hungary (Esto´k
2007) and their absence in Greece during summer (Helversen
and Weid 1990) suggests that N. lasiopterus is a migratory
species in Eastern Europe, which females possibly migrate
south, along the Adriatic coast, mating with the males along
the way. Therefore, further research should encompass a
year-round study to investigate the sexual composition of the
species. Echolocation analysis should be based on recent
studies (Haquart and Disca 2007, Esto´k and Siemers 2009,
Haquart et al. 2010) to differentiate N. lasiopterus from other
bat species such as T. teniotis which is present in the same
area and has similar echolocation characteristics. Radio-
tracking should also be conducted to study habitat use and
roost selection for the species (Esto´k et al. 2007, Sane´ 2008,
Popa-Lisseanu et al. 2009). In this way, more productive
monitoring guidelines and environmental study assessments
can be made, especially considering the vulnerability of Nyc-
talus species as one of the most affected European species
by wind turbines (Rodrigues et al. 2010).
Acknowledgements
Thanks are due to Martina Markov Podvinski as a project leader of
the ‘‘Biodiversity Protection in Croatia’s Kornati Archipelago:
Inventorying and monitoring of terrestrial species’’ (Ecological
association Argonauta in cooperation with BIUS) and Vanja C
ˇavrak
as a project leader of the Biological camp ‘‘Kornati ’09’’ (BIUS in
cooperation with Argonauta) through which the research has been
conducted. We also thank Zlatko Ruzˇanovic´ and the National Park
Kornati for all their help, as well as our colleagues Valerija Aptree-
va, Z
ˇeljka Drdar, Darija Josic´, Jure Krasic´, Ana Pusˇic´, Dania Randi
and Martina Ratko who participated in the fieldwork. We are grate-
ful to Goran Rnjak from Speleological Section of the Croatian
Mountaineering Society ‘‘Mosor’’ who helped us in more ways than
one and Marina Kipson for all her advice. We thank Ana Popa-
Lisseanu, Pe´ ter Esto´ k and the Associate Editor for improving the
manuscript and for their valuable suggestions, as well as Henry
Schofield for English proof-reading. Sandra Dobric´ kindly helped
by translating the French literature and Boris Krstinic´ helped by
preparing the map and illustrations. This research was partially
funded by the Royal Norwegian, Netherlands and the Embassy of
Finland in Zagreb. All capture and handling of bats were approved
by the Ministry of Culture Nature Protection Directorate of the
Republic of Croatia.
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100 D. Kovacˇ et al.: First record and recent capture of Nyctalus lasiopterus
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