Untitled

Lynx (Praha), n. s., 37: 9–12 (2006).
ISSN 0024–7774
Professor Vladimír Hanák, 75
Profesor Vladimír Hanák pětasedmdesátníkem
This volume contains the papers devoted to Vladimír Hanák, professor emeritus in Zoology
at Charles University of Prague, a leading personality in Czech vertebrate zoology and one of
the founding fathers of the modern study of mammals in the former Czechoslovakia and in
the Czech Republic, on occassion of his 75th birthdays. Of course, it should be stressed that
the senescence aspects commonly associated with that age like would not touch Vladimír in
any significant extent – even in these days he continualy works in field study of bats, finishes
a series of voluminous papers (among other, two volumes of a complete list of bat records in
the Czech Republic and detailed summaries of distributional status of all particular species),
a short time ago he coauthored immense analysis of bat banding results, a monographic survey
of Palaearctic bats, wrote dozens of popularising articles, a survey on history of zoology in
Central Europe and, at the same time, acts as a chief expert in various institutions of Nature
Conservancy and essentially contributes to supervising of various research projects including
PhD and MSc theses. Indeed, it is hard to believe that a man with so excellent mind and so
manifold spectrum of activities really achieved his 75.
Yet, it is the reality. Vladimír Hanák was born on 31 March 1931 in Znojmo, a town to which
he paid a considerable attention during last decades, among other with his essential role in designing the National Park Podyjí-Thayatal, where he organizes a board of experts and actively
promotes a large scale multidisciplinary investigations of that region. Going back to the region
of his youth and/or concentration on regional faunal summaries (like e.g. a survey of bats in
UNESCO Biospheric Reserve Třeboňsko he has finished just recently) is in no way a sign of
regression. Since beginning of his scientific career,
Hanák emphasized that a reliable primary record
is an ultimate prerequisite to any further study and
just because he ever put special attention to that
point his results and scientific achievement, which
much exceeded the regional scale of course, remain
largerly valid and invariant to extensive conceptual
and methodical shifts which appeared since time of
his beginnings.
Vladimír Hanák undoubtedly ranks among the
chief personalities who designed the framework
on which the Mid-European mammalogy of 20th
century has arisen. At the beginning of 1950s when
he started with profound critical reexamination of
the taxonomical and distributional status of mammals in Central Europe and the Balkans, the state of
knowledge was incredibly poor and to a considerable
degree patterned by numerous misinterpretations and
speculative predictions not rooted in a real evidence.
Hanák started with a critical revision of that state and despite he continously worked in more
groups of mammals and perforemed analyzes of mammalian communities and inter-regional
comparisons of mammal fauna in a general extent, he focused, already in 1950s, together with Jiří
Gaisler, his attention particularly to the group then the least known – bats. He started with large
scale field studies of that group including extensive bat banding program and multidisciplinary
gathering of various data on biology of particular species that was performed in cooperation
with his students and colleagues of his age such as Jiří Gaisler, Karel Hůrka, Ladislav Janský,
Milan Klíma, Jaroslav Figala, Leo Sigmund and others. Soon Hanák designed a large scale
project of detailed analyses of taxonomic and distributional status of bat species covering not
only the region of central Europe but the whole Palaearctic region. He succeeded to investigate
the bat collections in many prominent European institutions and came in close contacts with
almost all European specialists in that group and his approach soon became a model of a well
designed research effort widely respected in an international scale. Among the achievements
which particularly contributed to Hanák’s international reputation were first of all his pioneer
role in discovery of sibling species Plecotus austriacus and Myotis brandtii, detailed taxonomical
analyses of Myotis mystacinus group, studies on taxonomy and distribution of several least known
taxa of the Palaearctic fauna such as Rhinolophus bocharicus, Eptesicus sodalis, Myotis longipes,
1970s: Vladimír Hanák among his colleagues and students during a field excursion to southern Moravia,
the county of his childhood.
10
Hypsugo savii (and
several other mostly in coauthorship
with Jiří G aisler
or Ivan Horáček)
or the papers providing first comprehensive data on bats
of Bulgaria, Libya
and other regions.
Unfortunately, only
a smaller part of
his achievements
was properly published and the rest
remained in state of
manuscripts (such
as his voluminous
1960 PhD thesis
which summarized 2006: Vladimír Hanák with his students (and the editors of this volume) during
taxonomical and field studies in southern Bohemia.
distributional status
of all Mid-European bat species throughout whole their ranges or a prepared monograph on
the genus Myotis).
Nevertheless, it was not only his research effort and publications for which Hanák became
a leading personality of the post-war generation of Czech zoologists and a man who actually
imprinted the development of the discipline in this country for next decades. His university
teaching and the design of the vertebrate zoology program at the Charles University to which
he essentially contributed present undoubtedly particularly significant part of Hanák’s achievements.
Almost all Czech zoologists of today generations are direct or indirect students of Vladimír
Hanák and perhaps all, similarly as many foreign zoologists who met Hanák personally were
deeply impressed of his charming personality. It is not easy to identify which is source of the
charm similarly as it is not easy to specify what all atributed the fact which all Hanák’s students
like to stress – namely, that it was just the Hanák’s role as a university teacher what became
an essential imprinting factor that stimulated their further career in zoology, their view of the
specific qualities of that discipline and the meaning of the scientific study in that branch. The
Hanák’s lectures, the practical courses or field excursions were, of course, traditionally performed as a top of professional standard and despite of Hanák’s sense of humor and tolerance
with quite strong requirements on students’ comprehension. Yet, neither that was perhaps the
most essential. What may have played a more significant role was probably the context in
which Hanák the specificities of the branch and the particular scientific problems he uses to
expose. Quite characteristic is his vivid, unostentatious but highly qualified interest in a broad
spectrum of phenomena and processes which make together the phenomen of Nature, his deep
respect to their aesthetic dimensions as well as to historical and cultural heritages which form
our comprehension to them. Hanák perceives the Nature and animals like instant source area
11
of aesthetic and intelectual pleasures and the science of life grows thus in his representation in
an extremelly attractive domain which opens a real chance to integrate an intelectual creativity
with a meaningfull way of life rich in aesthetic and cultural issues. Vladimír has been quite
consequent in respecting such a view in his own life and particularly with that, with his way to
treat the pleasures and troubles of life and with his richly human interest on life if his students
he has instantly demonstrated actual relevance and qualities of that view and of the way of life
indexed by it.
Moreover, Hanák never applied techniques that nowdays are nearly obligatory prerequisite to
a scientific career: conceptual opportunism, affiliation of own view of a topic to the mainstream interpretations, overexposing or overestimation of own achievements. In consequence, his
publications are not excessive either in number or in impact factors of the periodical in which
they appeared but without exception they bring the concise information which reliability stays
without any doubt. The ultimate index of Hanák’s scientific efforts was the actual meaning
of the facts and their relevance within a broad contextual background in which his studies did
exposed. The respect to his own sense of meaning of particular facts and the deep meaning of
the world was valued for him much more than aspects of his academic career, secular honours
or public fame.
His straightforward way of presenting his opinions and deeply rooted sense of justice brought
him many troubles, particularly in the period after occupation of Czechoslovakia by foreign
armies in 1968. His career as well as contacts with the colleagues abroad were stopped for long
years. The situation dramatically changed after the “velvet revolution” in 1989 when Hanák
was immediately appointed associate professor and elected to various academic positions in
which he considerably help in transformation of the university study of biology into a vivid
modern institution.
We do not wish, however, to repeat here the biographic data of Vladimír Hanák, neither
enumerate his particular research achievements nor will provide list of his publications, by the
way extensivelly enlarged during the recent decades. The reader can find these information in
the volume Prague Studies in Mammalogy edited on occassion of Hanák’s 60th birthdays.
Here we would like only to express, on behalf of numerous students and friends, warm thanks
to Vladimír for all the gifts which he provided us and to wish him to the decades to come a good
health, optimistic mind and lot of pleasure which he uses so generously to diseminate around
throughout his life.
The Editors
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Lynx (Praha), n. s., 37: 13–17 (2006).
ISSN 0024–7774
Holocene remains of the Pine marten (Martes martes) from
the Tatra Mts. (Poland) – skull morphology and population structure
(Carnivora: Mus­te­li­dae)
Zmienność morfologiczna czaszek holoceńskiej populacji kuny leśnej (Martes
martes) z jaskiń tatrzańskich i populacji współczesnej (Carnivora: Mustelidae)
Justyna Bachanek & Bronisław W. Wołoszyn
Polish Academy of Sciences, Institute of Systematics and Evolution of Animals,
ul. Sławkowska 17, PL–31-016 Kraków, Poland
received on 6 June 2006
Abstract. A morphological analysis of 36 skulls of the pine marten Martes martes (Linnaeus, 1758) col­
lec­ted in caves in the Tatra Mts. (southern Poland) was carried out. Special attention was paid to age and
sex structure of this Holocene population. The results of skull measurements of the subfossil population
were compared with those of recent pine marten populations from Poland.
INTRODUCTION
The paper is aimed at the analysis of skull morphology of subfossil remains of the pine mar­ten
Martes martes (Linnaeus, 1758) from Poland regarding age and sex structure of the po­pu­la­ti­on.
So far, no studies have been focused particularly on this aspect. Most publications are devoted
to morphological comparison of skulls between Martes martes and Martes foina (Erxleben,
1777). The differences between the two species were presented by Anderson (1970), Altuna
(1973), Steiner & Steiner (1986) using non-metrical parameters, and Ge­ra­si­mov (1985) using
a multivariate analysis of variance. In Europe, the skull morphology of martens was studied in
various aspects and according to various criteria: age classes – Röt­tcher (1965), Habermehl
& Röttcher (1967), sex dimorphism Bree et al. (1970), Ros­so­li­mo & Pavlinov (1974), as well
as variability and asymmetry of dentition of pine and beech martens in Poland (Wolsan 1985,
1986, 1989) and in Europe (Reig 1989). The above men­ti­o­ned publications were aimed only
at com­pa­ri­sons of recent populations of the two marten species, whereas there are no articles
comparing recent and subfossil populations.
MATERIAL AND METHODS
Several hundreds of caves are known in the Tatra Mts., mostly in their western part. These caves are
characterized by specific ecoclimate and fauna. The material of 36 pine marten skulls was collected in
the following Tatra caves: Czarna (7 specimens), Zimna (4), Mała Świstówka (1), Miętusia (1), Cho­
chołowska (2), Bielska (1), Ptasia Studnia (6), Niedźwiedzia (1), Za siedmioma Progami (12), Wysoka
(1). The studied material was collected between 1961 and 1981, mostly by K. Kowalski, A. Woźnica,
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M. Zagórny, M. Gębołys, B. W. Wołoszyn and has been deposited in the Institute of Systematics and
Evolution of Animals, PAS, in Kraków.
The subfossil skulls of the pine marten were found on the surface of cave floor and most probably are
of the Holocene age.
The following measurements were taken: condylobasal length (CBL), alveoli condylobasal length
(CBLa); zygomatic width (ZYW); postorbital width (PorW); mastoid width (MstW). Males and females
were di­stin­guis­hed on the basis of differences in alveoli condylobasal length (CBLa) and zygomatic
width (ZYW).
We used the simplified age classification presented by Buchalczyk & Ruprecht (1989), based on the
following characters: (1) state of preservation of sutura internasalis, s. nasomaxillaris, s. nasofrontalis,
and s. maxillofrontalis; (2) degree of tooth wear; and (3) crista sagittalis development.
Three age classes were distinguished: (I) young individuals, contours of s. nasomaxillaris, s. na­so­fron­ta­
lis, and s. maxillofrontalis visible; beginning of crista sagittalis formation; rough surface of the toothrow;
pre­su­med age: 5–8 months; (II) mature specimens, crista sagittalis situated below the line, rough surface
of the toothrow; presumed age: between 9 months and 2 years; (III) old individuals, long crista sagittalis,
smooth surface of the toothrow, some teeth may be missing; age: 2 years or more.
The data on skull morphology of the studied Holocene population were compared with the results of
measurements of the recent marten population taken from the publication by Reig & Ruprecht (1989).
These authors studied Martes martes from four regions of Poland: Masurian and Pomeranian Lakelands,
Wielkopolska-Kujawy Lowland, Białowieża Forest, and Silesia.
Since the subfossil material was damaged, it was impossible to take some of the measurements usually
taken on recent material. The CBL length of the skull of recent specimens is measured from the anterior
margin of incissivi to the posterior margin of condylus occipitalis. However, incisors are missing in most
subfossil skulls, therefore the CBLa was taken. Results of both measurements taken on the same recent
Table 1. Range and mean values of skull measurements of subfossil and recent martens (males and females
separately). Explanations: CBLa – alveolar condylobasal length; CBL – condylobasal length; ZYW – zy­
go­ma­tic width; PorW – postorbital width
Tab. 1. Zróżnicowanie holoceńskiej populacji kun ze względu na różnorodność badanych parametrów.
measurement
mean±SD
Holocene
min–max
females
CBLa
74.97±1.83
72.15–78.33
CBL
ZYW
42.68±2.75
36.47–48.69
PorW
18.25±1.38
16.35–21.50
mean±SD
Recent
min–max
78.5±4.46
75.0±4.27
45.3±2.40
17.3±6.30
72.15–88.15
75.50–81.60
43.00–48.00
15.30–20.00
males
CBLa
83.18±2.22
79.80–88.15
CBL
85.6±1.70
ZYW
47.29±2.62
43.23–50.70
51.1± 3.20
PorW
19.24±0.83
17.77–21.01
19.1±6.50
82.0–87.90
47.6–55.20
15.8–21.30
all individuals
CBLa
78.50±4.46
72.15–88.15
CBL
82.2±1.43
ZYw
44.20± 3.46
36.50–50.70
48.2±1.35
PorW
18.50±1.55
15.25–22.55
18.2±1.16
78.75–84.75
45.30–51.60
15.50–20.65
14
specimens were compared. An average difference between the two measurements was 0.6 mm, i.e. 0.01%,
being within statistical error limits.
An analysis regarding sex and age structure of the population was carried out. Statistical significance
level of all differences was below 0.005. Values were transformed before the statistical analysis. All P-values reported are two tailed, the T-test of the differences between two means was performed.
RESULTS
Sexual dimorphism and population structure
Both the Holocene and recent marten po­pu­la­ti­ons exhibit a distinctive sex dimorphism: ma­les
are larger than females. The di­stin­guis­hing characters are: condylobasal length (CBL) or alveoli
condylobasal length (CBLa), and zy­go­ma­tic width (ZYW). In the subfossil population, differences in CBL and ZYW between the two sexes are statistically significant (p<0.001). Based
on these characters, we classified 15 spe­ci­mens as females, and 11 as males (Fig. 1), while skull
dimensions of the remaining 10 specimens did not allow their classification.
Comparison of subfossil and recent material
Within the sex groups, the Holocene marten skulls are shorter than the recent ones by 3.85 mm
on average (sta­tis­ti­cal­ly significant difference, p<0.005; Table 1).
Fig. 1. Skull dimensions of Holocene martens – sex groups.
Rys. 1. Zróżnicowanie holoceńskiej populacji kun ze względu na płeć.
15
Fig. 2. Sex and age structure of the studied Holocene marten population.
Rys. 2. Struktura populacji holoceńskiej pod względem płci i wieku.
Age of individuals
The Holocene marten sample comprises 36 specimens. Most of them are adults from the II
age class (n=24). Older specimens (of III age class) are less numerous (n=10), and only two
individuals were classified as young.
DISCUSSION
Trogloxenes – animals inhabiting caves incidentally or using them as seasonal shelters – are
the most numerous group in the fauna of caves. The pine marten Martes martes is one of such
species: it occurs in caves sporadically but gets through cave corridors far from the entrance.
The authors analysed the structure of a subfossil marten population on the basis of skull
morphology. Our results show that the Holocene martens had on average by 3.85 mm smaller
skulls than the recent ones. This difference may be related to thermoregulation requirements in
different climatic conditions: according to Bergmann’s rule animals living in warmer cli­ma­te
tend to have a relatively higher body surface to body mass ratio (e.g. smaller body size) than
animals living in colder climate where reduction of relative body surface (e.g. larger body size)
is favored. It is likely that the remains of martens from the Tatra caves originate from the Holocene optimal climatic period, Atlantic or Subatlantic, when smaller individuals could easily
radiate excess of heat. The results of radiocarbon dating (C14) of the subfossil material could
bring a definitive answer to this question.
The age structure analysis was carried out using simplified age categories proposed by Bu­
chal­c­zyk & Ruprecht (1989). These authors assessed the age of 596 marten specimens, grou­
ping them into five age groups. For the small Holocene sample (n=36) we reduced age groups
to three. Mature specimens prevailed in our material, while old animals (of 2 or more years of
age) were by half less numerous. Young individuals (of less than 8 months) were rare – only
two specimens out of the 36. The low proportion of young animals indicates that caves were
used by martens as temporary shelters or hunting grounds, not for raising offspring (Fig. 2).
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Podsumowanie
Zbadano holoceńską populację kun leśnych (Martes martes) z jaskiń tatrzańskich. Wybrane parametry
po­mi­a­rów czaszki porównano z pomiarami wykonanymi na materiale współczesnym. Mniejsze roz­mi­a­ry
po­pu­la­c­ji holoceńskiej (Tab.1) wskazują zgodnie z Regułą Bergmanna, że badana populacja zasiedlała ten
teren w cieplejszym klimacie, prawdopodobnie w optimum klimatycznym Holocenu.
REFERENCES
Altuna J., 1973: Distincion craneal entre la Marta (Martes martes) y la Foina (Martes foina) (Mammalia).
Munibe, 25: 33–38.
Anderson E., 1970: Quaternary evolution of the genus Martes (Carnivora, Mustelidae). Acta Zool. Fenn.,
130: 1–132.
Buchalczyk T. & Ruprecht A. L., 1977: Skull variability of Mustela putorius Linnaeus, 1758. Acta The­
ri­ol., 22: 87–120.
van Bree P. J. H., van Mensh P. J. A. & van Utrecht W. L., 1970: Sur le dimorphisme sexuel des canines
chez la Foine, Martes foina (Erxleben, 1777). Mammalia, 34: 676–682.
Gerasimov S., 1985: Species and sex determination of Martes martes and Martes foina by use of systems
of craniometrical indices developed by stepwise discriminant analysis. Mammalia, 49: 235–248.
Habermehl K. H. & Röttcher D., 1967: Die Möglichkeiten der Altersbestimmung beim Marder und Iltis.
Ztschr. Jagdwiss., 13: 89–102.
Reig S., 1989: Morphological variability of Martes martes and Martes foina in Europe. Ph.D. Thesis,
Jagiellonian University, Cracow, 145 pp.
Reig S. & Ruprecht A. L., 1989: Skull variability of Martes martes and Martes foina from Poland. Acta
Theriol., 34: 595–624.
Rossolimo O. & Pavlinov I., 1974: Sexual dimorphism in the development, size and proportions of the
skull in the pine marten (Martes martes Linn.: Mammalia, Mustelidae). Pp.: 180–191. In: King C. M.
(ed.): Biology of the Mustelids: some Soviet Research. Volume 1. British Library, Boston Spa, Yorkshire,
266 pp.
Röttcher D., 1965: Beträge zur Altersbestimmung bei Nerz, Steinmarder und Iltis. Inaugural-Dis­ser­ta­ti­
on zur Erlangung des Doktorgrades. Veterinärmedizinischen Fakultät der Justus Liebieg-Universität,
Gies­sen, 83 pp.
Steiner H. M. & Steiner F. M., 1986: Die nicht-metrische Unterscheidung von Schädeln mitteleuropäischer Baum- und Steinmarder (Martes martes und Martes foina, Mammalia). Ann. Naturhist. Mus.
Wien, 88/89 B: 267–280.
Wolsan M., Ruprecht A. L. & Buchalczyk T., 1985: Variation and asymmetry in the dentition of the pine
and stone martens (Martes martes and M. foina) from Poland. Acta Theriol., 30: 79–114.
Wolsan M., 1986: Morphological variation of the first upper molar in the genus Martes (Carnivora, Mus­
te­li­dae). Mem. Mus. Natn. Hist. Natur. Paris (S. C), 53: 241–254.
Wolsan M., 1989: Dental polymorphism in the genus Martes (Carnivora: Mustelidae) and its evo­lu­ti­o­na­ry
Significance. Acta Theriol., 34: 545–593.
Wołoszyn W., 1996: Fauna jaskiń tatrzańskich [Fauna of the Tatra caves]. Pp.: 525–533. In: Mirek Z.,
Głowaciński Z., Klimek K. & Pięko-Mirkowa H. (eds.): Przyroda Tatrzańskiego Parku Narodowego
[Nature of the Tatra National Park]. Tatrzański Park Narodowy, Kraków-Zakopane.
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Lynx (Praha), n. s., 37: 19–21 (2006).
ISSN 0024–7774
Savi’s pipistrelle (Hypsugo savii): bat species breeding in the Czech
Re­pub­lic (Chiroptera: Vespertilionidae)
Netopýr Saviův (Hypsugo savii): další v Česku se rozmnožující druh netopýra
(Chiroptera: Ve­sper­ti­li­o­ni­dae)
Tomáš Bartonička1 & Peter Kaňuch2,3
1
Institute of Botany and Zoology, Faculty of Science, Masaryk University, Kotlářská 2,
CZ–611 37 Brno, Czech Republic; [email protected]
2
Institute of Forest Ecology, Slovak Academy of Sciences, Štúrova 2, SK–960 53 Zvolen, Slovakia;
[email protected]
3
Institute of Vertebrate Biology, Academy of Sciences CR, CZ–675 02 Studenec 122, Czech Republic
received on 25 November 2006
Abstract. On 17 August 2006, an adult female Savi’s pipistrelle (Hypsugo savii) was mist-netted above
a park fountain in the Brno city. The female showed signs of postlactation like the presence of bare patches
around its bulgy nipples. It was measured, banded and next night released in the same park. Previous
records of the species concerned two males and a subadult female. This is the fourth record of H. savii but
the first reliable one indicating reproduction of this species in the territory of the Czech Republic.
Savi’s pipistrelle, Hypsugo savii (Bonaparte, 1837) has a West Palaearctic distribution range
and in Europe it is widespread mostly in the Mediterranean region (Horáček & Benda 2004).
Previous records of the species in the Czech Republic up to 2003 were made outside the bat’s
breeding period, thus irrelevant to the problem of reproduction of the species. First male was
found 20 km southwards of Brno in the village of Žabčice and a subadult female and an adult
male just within the territory of Brno (Gaisler 2001, Gaisler & Vlašín 2003).
At the evening of 17 August 2006, two mist-nets (7 and 12 m long) were set over a his­to­ri­cal
stone fountain (five meters in diameter) in the Lužánky park in the city of Brno. The park, ca.
400×400 meters, has a perimeter of ca. 1.8 km. Full-grown and hollow trees in the park as well
as old historical buildings surrounding the park provide high number of potential roosts for bats.
An artificial pond was built recently in the western part of the park and is also im­por­tant to the
bats. However, the shallow water body in the fountain was the only water habitat of the park
and its surroundings until the vegetation season of 2005. Mist-netting lasted between 8:00pm
and 11:30pm. We caught 13 individuals of five bat species – mostly Pipistrellus pipistrellus
(Schre­ber, 1774) s. str. and per one individual of Eptesicus serotinus (Schreber, 1774), Plecotus
auritus (Linnaeus, 1758) and Myotis daubentonii (Kuhl, 1817). The female of H. savii was
caught at 9:10 pm. She had signs of postlactation such as the presence of bare patches around
its bulgy nipples and swollen external genitals. Wings, ears and face were dark in contrast to
the brown dorsal fur and grey belly. There were two cusps on I2, between I3 and C1 was a gap,
C1 and P4 were in contact. The peak frequency of the search sequence of echolocation calls was
19
between 33.0–35.5 kHz. External measurements: body 47.8 mm, tail 32.8 mm (free tip of the tail
was 3 mm long), forearm 35.1 mm, hind foot 6.7 mm, ear 10.2, tragus 4.3 mm, the fifth finger
42.6 mm, the third finger 57 mm, body mass 6.8 g. A sample of wing membrane was preserved
in 96% ethanol for later DNA analysis. A photograph was made using a digital camera (Fig. 1).
The bat was marked with the band N. Museum Praha No X16019, kept in captivity and next
night released within the same park. It flew away without any obvious difficulties.
In addition to the two published records (Gaisler 2001, Gaisler & Vlašín 2003), an adult
male was found in March 2006 within the city of Brno (leg. P. Koutný). It was seriously injured,
with its left forearm broken. The bat was deposited in the collections of the Institute of Botany
and Zoology, Faculty of Science, Masaryk University. Fast train of records of H. savii during
the last five years shows spreading of the spe­ci­es towards the north. During the last two years
several individuals of H. savii, in winter as well as in summer, were found also in Slovakia (Le­
hotská & Lehotský 2006, Lehotská 2006, Ceľuch & Ševčík 2006). It can be assumed that the
species is an active immigrant to the Czech Republic and Slovakia. The records from Slovakia
are located more to the south that these from Brno. At present, Brno and its environs seems to
be the northernmost point of the species’ regular occurrence (Gaisler & Vlašín 2003) and the
potential reproduction. Two individuals found in northern Ger­ma­ny and another from England
were probably transported passively (Ohlendorf et al. 2000, Gaisler 2001). Low spectral parameters and, in particular, low peak frequency of pipistrelle like echolocation calls recorded
in Nitra and Brno (lower than in signals of Pi­pis­t­rel­lus nathusii) suggest more often occurrence
of H. savii in urban areas than expected (T. Bar­to­nič­ka and M. Ceľuch, unpublished). Bats can
Fig. 1. Postlactating female of Savi’s pipistrelle (Hypsugo savii) mist-netted in the Brno city.
Obr. 1. Postlaktační samice netopýra Saviova (Hypsugo savii) odchycená v Brně.
20
occupy the prefab houses similarly as Nyctalus noctula (Schreber, 1774) or Vespertilio murinus
(Linnaeus, 1758). Possible existence of a ma­ter­ni­ty colony will be investigated in near future.
SOUHRN
V centru Brna byla dne 17. 8. 2006 odchycena dospělá samice netopýra Saviova (Hypsugo savii). Samice
vykazovala jasné postlaktační znaky jako bezchlupé okolí zvětšených prsních bradavek a zduřelé vnější
genitálie. Celkově se jedná již o čtvrtý záznam druhu na území České republiky avšak jde o první doklad
jeho reprodukce na tomto území.
Acknowledgements
We are very grateful to J. Gaisler for valuable comments to the manuscript. Female was captured during
the activities involving the study of pipistrelle’s populations in central Europe. The study was supported by
the grant of Grant Agency of the Czech Republic No 206/06/0954 and the grant of Ministry of Education,
Youth and Sports of the Czech Republic No MSM0021622416.
References
Gaisler J., 2001: A mammal species new to the Czech Republic – Savi’s pipistrelle Hypsugo savii. Folia
Zool., 50: 231–233.
Gaisler J. & Vlašín M., 2003: Second record of the Savi’s pipistrelle (Hypsugo savii) in the Czech Re­
pub­lic. Vespertilio, 7: 181–182.
Horáček & Benda P., 2004. Hypsugo savii (Bonaparte, 1837) – Alpenfledermaus. Pp.: 911–941. In: Krapp
F. (ed.): Handbuch der Säugetiere Europas. Band 4: Fledertiere. Teil II: Chiroptera II. Ve­sper­ti­li­o­ni­dae
2, Molossidae, Nycteridae. AULA-Verlag, Wiebelsheim, x+605–1186 pp.
Lehotská B., 2006: Second record of the Savi’s pipistrelle (Hypsugo savii) in Slovakia. Vespertilio, 9–10:
225–226.
Lehotská B. & Lehotský R., 2006: First record of Hypsugo savii (Chiroptera) in Slovakia. Biológia,
Bra­ti­sla­va, 61: 192.
Ohlendorf B., Vierhaus H., Heddergott M. & Bodino F., 2000: Korrektur: Fund einer Nordfledermaus
(Eptesicus nilssoni) in Hamburg (ds. Z. Bd. 5, p. 220) betraf eine Alpenfledermaus (Hypsugo savii).
Nyctalus (N. F.), 7: 454.
Ceľuch M. & Ševčík M., 2006: Netopiere (Chiroptera) v meste Nitra (predbežná správa) [Bats (Chiroptera)
in the City of Nitra (Preliminary Report)]. Skupina pre ochranu netopierov, Nitra, 11 pp (in Slovak).
21
Lynx (Praha), n. s., 37: 23–44 (2006).
ISSN 0024–7774
On the occurrence of Eptesicus bobrinskoi in the Middle East
(Chiroptera: Vespertilionidae)
K výskytu netopýra turanského (Eptesicus bobrinskoi) na Blízkém východě
(Chiroptera: Vespertilionidae)
Petr Benda1,2 & Antonín Reiter3
1
Department of Zoology, National Museum (Natural History), Václavské nám. 68, CZ–115 79 Praha 1,
Czech Republic; [email protected]
2
Department of Zoology, Charles University, Viničná 7, CZ–128 44 Praha 1, Czech Republic
3
South Moravian Museum Znojmo, Přemyslovců 8, CZ–669 45 Znojmo, Czech Republic;
[email protected]
received on 23 September 2006
Abstract. A profound morphologic comparison of smaller representatives of the genus Eptesicus in­ha­
bi­ting the western Palaearctic (E. bobrinskoi, E. gobiensis, E. nasutus, and E. nilssonii) confirmed the
occur­ren­ce of Eptesicus bobrinskoi in NW Iran. Although it was previously not fully accepted, a revision
of the collection material as well as of specimens newly recorded in Iran brought evidence showing that
the doubts were unjustified.
INTRODUCTION
The Bobrinskoy’s serotine, Eptesicus bobrinskoi Kusjakin, 1935, was described on the basis
of three individuals collected by S. P. Naumov in May and June 1928 in a region close to the
north-eastern shore of the Aral Sea, central Kazakhstan, which were primarily assigned by
Bob­rin­skoi (1931) to the form Eptesicus caucasicus (Satunin, 1901) [= Hypsugo savii (Bo­
na­par­te, 1837)]. Kusjakin (1935) characterised E. bobrinskoi to be in its external appearance
close to Eptesicus alashanicus Bobrinskoj, 1926 [= Hypsugo alashanicus] but with a larger
skull, al­thou­gh both forms were of similar body size. He mentioned E. bobrinskoi to be ty­pi­cal
with its flattened braincase, a relatively small second upper incisor, and a very pale co­lo­ra­ti­on
of the pelage and naked parts. The Tjulek well in the Aral part of the Karakum Desert [= Priaralskye Karakumy D.], 65 km E of the town of Aral’skoe more [= Aral’sk], Kazakhstan, was
designated as the type locality of this species (Kusjakin 1935, Rossolimo & Pavlinov 1979).
Ellerman & Morrison-Scott (1951) provisionally arranged E. bobrinskoi as a sub­spe­ci­es of
E. nasutus (Dobson, 1877) but this view has not generally been followed (see also Harrison
1963 and Hanák & Gaisler 1971). E. bobrinskoi remains the only bat species described from
Kazakhstan (Butovskij et al. 1985).
The essential part of the distribution range of E. bobrinskoi covers the zone of colder deserts
and semi-deserts of central Kazakhstan, between the Aral Sea in the west and the western Betpak-Dala Desert in the east (Strelkov 1980), i.e. between 45° 30’ – 49° N and 59° – 70° 15’ E
23
(Butovskij et al. 1985). About 90% of the records ascribed to this species come from this region
(cf. Strelkov 1980, Strelkov & Šajmardanov 1983).
A few of records of E. bobrinskoi were described from outside this main range. Kuzjakin
(1950) reported a young male specimen from the vicinity of Faskal, North Ossetia, northern
slope of the Greater Caucasus Mts He also mentioned a female specimen deposited in the alcohol collection of the Zoological Institute of the Academy of Science [St. Petersburg] labelled
“Jakutsk, Mid­den­dorf” suggesting its origin in Yakutia [= Sakha, northern Siberia, Rus­sia].
These records seem to be unusual from the biogeographical point of view but have been largely
considered rather correctly identified (Vereščagin 1959, Strelkov 1963, 1981, Ta­vrov­skij et
al. 1971). However, Kuzjakin (1965) labelled both records with question marks (?) and noted
the preservation state of these specimens as not quite perfect and mainly, the age of the individuals as juvenile. In this way, Kuzjakin (1965) indirectly doubted his previous identifications.
Therefore, these records were considered clearly doubtful by Hanák & Ho­rá­ček (1986) who
regarded them to be of juvenile individuals of E. nilssonii (Keyserling et Blasius, 1839). This
view was followed by Nowak (1994), Borisenko & Pavlinov (1995), Pavlinov & Rossolimo
(1998), Horáček et al. (2000), and Simmons (2005), but not by Ko­op­man (1993, 1994) or Duff
& Lawson (2004). Anyway, E. bobrinskoi has not recently been considered a member of the
Yakutian bat fauna (Revin et al. 2004).
Strelkov (1980) described a finding of several individuals of E. bobrinskoi in the northern
Caspian Region in western Kazakhstan (the Mus Tomb, 18–20 km W of Kosčagyl, ca. 10 km
S of the lower Emba River). Although geographically isolated, this record has been accepted
beyond any doubt by later authors (Butovskij et al. 1985, Strelkov 1986, Rybin et al. 1989,
Ilyin 2000). Moreover, a new record of E. bobrinskoi has been recently reported from a region
close to the latter one (Davygora et al. 1998, Ilyin 2000), from a site 20–25 km S of Krasnojar, No­vo­a­lekse­e­v­ka Dist., Aktjubinskaja Region, NW Kazakhstan. This record re­pre­sents the
northwesternmost verified spot of occurrence of this species in the Palaearctic, and because the
Emba River is considered by some authors to be the Euro-Asian border in the region between
the Caspian Sea and the Ural Mts, it is now appropriate to place E. bobrinskoi on the list of
European bat species (Benda & Hutterer 2005).
The only record of E. bobrinskoi in the Middle East (and also outside the former USSR) was
published by Harrison (1963) from north-western Iran. He reported a finding of seven dead
individuals (four males, three females) in “the Sulphur caves at Guter-Su, north of Mt. Sabalan,
[…] Iranian Azerbaijan, 38° 10’ N, 47° 40’ E”. Harrison (1963) supported this spe­ci­es iden­ti­fi­
ca­ti­on by his observation of the typical morphologic characters according to Kuzjakin (1950),
and also by a comparison with the collection material of E. nilssonii and E. nasutus which both
markedly differed from the Guter-Su specimens in size and in all im­por­tant cranial traits. The
Harrison’s (1963) identification was confirmed by Hanák & Gaisler (1971) who used his data
in their broad comparison of the genus Eptesicus.
E. bobrinskoi was for a long time considered to be a part of the fauna of Iran (Lay 1967,
Etemad 1969, Hanák & Gaisler 1971, Corbet 1978, DeBlase 1980, Allison & Gaisler 1982,
Butovskij et al. 1985). However, Hanák & Horáček (1986) revised the series of skulls of E.
bobrinskoi from the Guter-Su finding, which is deposited in the collection of the Na­tu­ral History
Museum, London (BMNH), under the Nos. 63.1186–1192. They concluded that: “(a) Alle un­ter­
such­ten Stücke einschlißlich der größten (Nr. 63 1189) sind diesjährige Jungen mit auffallend
niedrigeren Werten der Schädelbreite, mit einer unvollendeten Ossifikation der Terminalränder
des Processus coronoideus und mit einer unvollendeten Eruption der Zäh­ne, u. ä. (b) Die met­ri24
s­chen Werte und proportionen des Rostrums, welche eher der Art E. bob­rin­skoi entsprechen, sind
aus den oben angeführten Gründen wenig nachweisbar; nach wich­tig­s­ten anderen Merk­ma­len
(Anwesenheit der Foramina cavernosa, Fellfährung) entspricht das Material vielmehr der Art
E. nilssoni. (c) Aus den angeführten Tatsachen ergibt sich, daß dieses Material of­fen­sicht­lich
juvenile Stücke der zentralasiatischen Population E. nilssoni darstelt. […] (d) Die an­ge­führ­te
Interpretation des Fundes aus Guter-Su steht im Einklang mit der bisherigen Ansicht über die
zoogeographischen Bewertung von E. bobrinskoi.” Ac­cor­ding to this revision, E. bobrinskoi
has no longer been regarded as a member of the bat fauna of the Middle East (although not
absolutely, see e.g. Nowak 1994, Sharifi et al. 2000), and the Harrison’s (1963) record has
been considered to be of E. nilssonii gobiensis Bob­rin­skoj, 1926 (Koopman 1993, 1994, Bori­
senko & Pavlinov 1995, Pavlinov & Rossolimo 1998, Horáček et al. 2000, Duff & Lawson
2004, Simmons 2005, cf. Rybin et al. 1989, Rydell 1993, Benda & Horáček 1998, Albayrak
2003). However, based on a profound mor­pho­lo­gic analysis, Strelkov (1986) showed the
latter form to represent a separate species, E. go­bi­en­sis. This result has been widely accepted
(Pavlinov & Rossolimo 1987, Rydell 1993, Borisenko & Pavlinov 1995, Bates & Harrison
1997, Roberts 1997, Pavlinov & Ros­so­li­mo 1998, Horáček et al. 2000, Šajmardanov 2001,
Simmons 2005, etc.).
During a field trip to Iran in the late spring of 2006 we visited the abandoned sulphuric mines
in the oriental thermal bath resort of Qutur Su (= Guter-Su by Harrison 1963). In these caverns
we found several dead birds and three partly mummified cadavers of bats. We pre­li­mi­na­ri­ly
identified these bats as small representatives of the genus Eptesicus Rafinesque, 1820, but differing from E. nilssonii or E. gobiensis by a shorter forearm in two individuals: LAt 34.4 and
36.7 mm (the third cadaver was without wings) in Qutur Su bats vs. 37–44 mm in E. nilssonii
s. l. (Hanák & Horáček 1986). These bats thus well resembled E. bobrinskoi ac­cor­ding to
Har­ri­son (1963).
The aim of this paper is to compare our specimens from Qutur Su with those identified as E.
bobrinskoi by Harrison (1963) as well as with representative samples of E. bobrinskoi, E. na­
sutus, E. nilssonii, and E. gobiensis, to identify and/or revise our and Harrison’s (1963) findings
and thus, to contribute to the knowledge of smaller Eptesicus bats in the Middle East.
THE RECORD
The three bats were found in two small abandoned sulphuric mines above the village of Qutur Su
( [qwtwr sw]), ca. 20 km ESE of Meshgin Shahr ( [mšgin šhr]), Province
of Ardabil, 38° 20’ N, 47° 51’ E; 2545 m a. s. l. (Fig. 1). Although slightly differing in the geographical
coordinates given, this site is un­doub­ted­ly identical with the locality of Guter-Su reported by Harrison
(1963). The site is situated on the northern slope of the main peak of the Sabalan Range ( [kwh
sblan]) which rises to the altitude of 4811 m from the flat steppe plateau of about 1100 m a. s. l. The village and mines lie in the zone of alpine meadows at the altitude of about 2500 m (Fig. 2). In these small
artificial rocky caverns, which act as a natural trap being filled by an unbreathable atmosphere, the bat
cadavers were found under smaller stones covering the mine floor.
MATERIAL AND METHODS
For a morphologic comparison of the newly collected bats in Qutur Su, we used museum material of all
smaller species of the genus Eptesicus occurring in the western Palaearctic; viz. E. bobrinskoi Kusjakin,
1935 (11 specimens) from central Kazakhstan (i.e. from its “topo-type” area), E. nasutus (Dobson, 1877)
(18) from the Middle East, including all currently recognised subspecies [E. n. nasutus from Afghanistan
25
Figs. 1, 2. Qutur Su, NW Iran. 1 – Abandoned sulphuric mines, the only site of (repeated) finding of E.
bob­rin­skoi in Iran (above). 2 – Alpine meadows at the altitude of ca. 2500 m sur­roun­ding the site of Fig.
1, situated on the northern slope of Mount Sabalan (4811 m a. s. l., in the background) (below).
Obr. 1, 2. Qutur Su, SZ Iran. 1 – Opuštěné sirné dobývky, jediná lokalita (opakovaného) nálezu netopýra
tu­ran­ské­ho (E. bobrinskoi) v Iranu (nahoře). 2 – Alpinské louky v nadmořské výšce zhruba 2500 m obklopující lokalitu na obr. 1, ležící na severním svahu hory Sabalan (4811 m n. m., v pozadí) (dole).
26
and from Persian Baluchestan, E. n. matschiei (Thomas, 1905) from SW Arabia, E. n. pellucens (Thomas,
1906) from Mesopotamia, and E. n. batinensis Harrison, 1968 from Oman], E. nilssonii (Keyserling et
Blasius, 1839) (24) from Central Europe, E. gobiensis Bobrinskoj, 1926 (6) from northern Kirghizia, as
well as the BMNH series of seven bats from Qutur Su identified by Harrison (1963) as E. bobrinskoi and
by Hanák & Horáček (1986) as E. nilssonii. Since Hanák & Horáček (1986) found the Qutur Su series
to be partially composed of immature individuals, we used also sets mixed from adult and subadult in­di­
vi­du­als for dimensional comparisons. For the data associated with the specimens see the Appendix.
For comparative purposes we used mainly skulls, from external dimensions we took only the forearm
length (LAt). The specimens were measured in a standard way using mechanical or optical callipers. We
evaluated 38 dimensions in each skull (17 measurements in skull and maxillar tooth-rows, 7 measurements
in the mandible and mandibular tooth-rows, see also Abbreviations) including 14 indices that described
the skull shape. The baculum was extracted with the use of a 4% solution of NaOH and coloured by the
Alizarin red. Statistical analyses were performed using the Statistica 6.0 software.
ABBREVIATIONS
Collections. BMNH = Natural History Museum, London, United Kingdom; CUP = Department of Zoology,
Charles University, Praha, Czech Republic; JOC = private collection of Ján Obuch, Blatnica, Slovakia;
NMP = National Museum (Natural History), Praha, Czech Republic; ZIN = Institute of Zo­o­lo­gy of the
Russian Academy of Science, Sankt Peterburg, Russia.
Measurements. LAt = forearm length; LCr = greatest length of skull; LCb = condylobasal length of skull;
LaZ = zygomatic width of skull; LaI = width of interorbital constriction; LaInf = rostral width between the
foramina infraorbitalia; LaN = neurocranium width; LaM = skull width at the mastiodal processes; ANc
= neurocranium height; ACr = skull height (incl. the tympanic bullae); CC = rostral width between the
canines (incl.); P4P4 = rostral width between the upper premolars (incl.); M3M3 = rostral width between
the third upper molars (incl.); IM3 = length of upper toothrow between the first incisor and the third molar,
I1M3 (incl.); CM3 = length of upper toothrow between the canine and the third molar, CM3 (incl.); P4M3 =
length of upper toothrow between the premolar and the third molar, P4M3 (incl); M1M3 = length of upper
toothrow between the first and third molars, M1M3 (incl.); CP4 = length of upper toothrow between the
canine and the premolar, CP4 (incl.); LMd = condylar length of mandible; ACo = height of the coronoid
process; IM3 = length of lower toothrow between the first incisor and the third molar, I1M3 (incl.); CM3
= length of lower toothrow between the canine and the third molar, CM3 (incl.); P4M3 = length of lower
toothrow between the second premolar and the third molar, P4M3 (incl.); M1M3 = length of lower toothrow
between the first and the third molar, M1M3 (incl.); CP4 = length of lower toothrow between the canine
and the second premolar, CP4 (incl.).
Others. m = male; f = female; ind. = individual of undetermined sex; a = adult; s = immature; A = alcohol
preparation; B = skin (balg); S = skull; Sk = skeleton.
RESULTS AND DISCUSSION
The two series of bats from Qutur Su, that published by Harrison (1963) and that newly col­lec­
ted, are very similar in dimensions (Table 1) and seem to comprise very close or even identical
morphotypes. To mention the most important characters, these specimens have si­mi­lar forearm
lengths, lengths of skull and tooth-rows and mainly, similar heights of bra­in­ca­se (ANc, ACr).
Therefore both sets of specimens from Qutur Su were compared as one group of samples.
Unlike Hanák & Horáček (1986), we found a part of the Harrison’s (1963) samples to be
adult individuals, which was also true for two of the newly collected bats (see Table 1). From
the whole set of the Qutur Su bats, at least five specimen bear signs of adult age on their skulls;
i.e. a fully ossified basilar suture in the skull base (the connection between ba­si­oc­cipi­tal and ba­
sis­phe­no­i­dal bones) and moreover, a noticeable teeth abrasion. Thus, skulls of these specimens
27
28
sex age LAt
NMP, 5 June 2006
90890
f
a 36.7 15.2414.88 9.56 4.14 4.86 7.95 8.11 4.74 5.88 4.32 6.04 5.21 3.38 1.9110.75 3.14 5.61 3.64 1.79
90891
f
s 34.4
–
– – 4.02 4.31
– – – – 3.88 5.63 5.07 3.37 1.8810.14 2.81 5.48 3.76 1.90
90892
m a
– 14.5714.43
– 3.95 4.48 7.16 7.90 4.61 5.88 4.00 5.88 5.02 3.39 1.8710.29 2.86 5.42 3.78 1.91
7.91 4.34 5.48 3.90 5.78 5.28 3.57 1.9610.51 2.91 5.62 3.85 1.84
8.22 4.47
– 3.79 5.94 5.38 3.55 2.12
– 3.12
– – –
7.77 4.52 5.51 3.74 5.61 5.08 3.47 1.8210.29 3.00 5.67 3.68 2.07
8.05 4.28 5.81
– – 5.07 3.35 1.81
– – – – –
LCr LCb LaZ LaILaInf LaN LaM ANc ACr CC M3M3 CM3 M1M3 CP4 LMd ACo CM3 M1M3 CP4
BMNH, 21 August 1961
63.1186
m a 35.0 14.6714.35
– 3.92 4.37 7.42
63.1187
m s 35.4 14.8214.28
– 4.11
– 7.82
63.1188
f
s 35.1skull broken
63.1189
m a 35.1 15.1714.87 8.38 4.19 4.28 7.64
63.1190
f
a 35.7 15.0714.53
– 4.08 4.48 7.87
63.1191
f
s 33.9skull broken
63.1192
m s 34.7skull entire but crushed
No.
Table 1. Dimensions of E. bobrinskoi collected at Qutur Su, NW Iran. For dimension abbreviations see the text. Forearm lengths (LAt) of the
BMNH specimens were taken from DeBlase (1980)
Tab. 1. Rozměry jedinců E. bobrinskoi nalezených v Qutur Su, SZ Iran. Vysvětlivky zkratek rozměrů viz text. Délky předloktí (LAt) jedinců
z londýnské sbírky (BMNH) byly převzaty z přehledu DeBlaseova (1980)
(BMNH 63.1186, 1189, 1190, NMP 90890, 90892) were available for a relevant morphologic
comparison.
As pointed out in Introduction, the Qutur Su bats are typical with relatively short forearms
(LAt 33.9–36.7 mm). In this character they are very close to the samples of E. bobrinskoi
from Kazakhstan (LAt 34.0–36.7 mm) and overlap also with the samples of E. nasutus (LAt
35.5–39.9 mm). On the other hand, the Qutur Su bats clearly differ in their forearm lengths
from the samples of E. nilssonii and E. gobiensis (Table 2).
Based on the comparison of skull dimensions, the skulls of the Qutur Su bats are most similar to the samples of E. bobrinskoi; they broadly overlap in all measurements taken (Table
2), although the Kazakh samples are on average slightly larger. The Qutur Su bats also partly
overlap in their skull lengths (LCr, LCb, CM3, P4M3, M1M3, LMd, CM3, P4M3, M1M3) and widths (LaZ, LaI, LaInf, LaN, LaM, M3M3) with E. nilssonii and slightly also with E. go­bi­en­sis,
but clearly differ from both sample sets in skull heights (ANc, ACr). However, the Qutur Su
bats clearly exceed in their skull lengts all samples of E. nasutus but overlap them in the skull
heights. These relations are shown also in Fig. 3; the Qutur Su bats are identical in skull size
(represented by its condylobasal length, LCb) with Kazakh samples of E. bobrinskoi and very
similar to the sam­ples of E. nilssonii and E. gobiensis. Concerning skull height (represented by
neurocranium height, ANc), the Qutur Su bats are identical with the samples of E. bob­rin­skoi
and also of E. nasutus. The samples of the latter species, however, completely differ by their
skull lengths. In this comparison the samples from Qutur Su and from Kazakhstan create one
Fig. 3. Scatter plot of the condylobasal length of skull (LCb) against the height of neurocranium (ANc)
in the compared samples of smaller Eptesicus species of the western Palaearctic; adopted from Strelkov
(1986). Data in millimetres.
Obr. 3. Srovnání kondylobasální délky lebky (LCb) proti výšce neurokrania (ANc) jedinců malých druhů
rodu Eptesicus západní Palearktidy; podle Strelkova (1986). Rozměry v milimetrech.
29
Table 2. Basic biometric data on the comparative samples of small representatives of the genus Eptesicus
of the western Palaearctic (see Appendix). For dimension abbreviations see the text.
Tab. 2. Základní biometrické údaje srovnávacích vzorků malých zástupců rodu Eptesicus západní Pa­leark­
ti­dy (viz Appendix). Vysvětlivky zkratek rozměrů viz text
E. bobrinskoi
NW Iran
M min max
E. bobrinskoi
C Ka­za­kh­stan
LAt
9 35.11 33.9 36.7 0.799 10 35.19 34.0 36.7 0.909
6 42.22 40.2 43.5 1.202
LCr
LCb
LaZ
LaI
LaInf
LaN
LaM
ANc
ACr
6 14.9214.5715.24 0.277
6 14.5614.2814.88 0.260
2 8.97 8.38 9.56 0.834
7 4.06 3.92 4.19 0.100
6 4.46 4.28 4.86 0.212
6 7.64 7.16 7.95 0.303
6 7.99 7.77 8.22 0.164
6 4.49 4.28 4.74 0.170
5 5.71 5.48 5.88 0.200
6 15.7515.1716.04 0.317
6 15.4314.9215.59 0.258
5 10.09 9.6110.28 0.276
6 4.11 3.92 4.28 0.136
6 4.96 4.75 5.11 0.148
6 8.07 7.82 8.42 0.216
6 8.59 8.27 8.81 0.201
6 5.20 4.97 5.47 0.207
6 6.47 6.32 6.68 0.121
CC
P4P4
M3M3
I1M3
CM3
P4M3
M1M3
6
6
6
7
7
7
7
LMd
ACo
I1M3
CM3
P4M3
M1M3
5 10.4010.1410.75 0.238
6 2.97 2.81 3.14 0.137
5 6.49 6.41 6.58 0.072
5 5.56 5.42 5.67 0.105
5 4.40 4.34 4.47 0.048
5 3.74 3.64 3.85 0.083
11 10.3610.0010.76 0.272
11 3.25 3.02 3.48 0.157
10 6.46 6.27 6.68 0.134
11 5.63 5.47 5.83 0.133
11 4.51 4.42 4.74 0.095
11 3.85 3.70 4.02 0.094
6 11.1510.6711.46 0.278
6 3.41 3.29 3.52 0.095
6 6.95 6.74 7.09 0.138
6 6.09 5.93 6.28 0.116
6 4.70 4.12 4.93 0.298
6 4.10 4.02 4.17 0.061
CM3/LCb
LaInf/LCb
LaN/LCb
LaM/LCb
ANc/LCb
ACr/LCb
M3M3/LCb
CC/CM3
M3M3/CM3
LaInf/LaI
M3M3/LaI
LaM/LaI
LaM/LaN
ACo/LMd
6 0.355
5 0.308
6 0.512
6 0.549
6 0.309
5 0.392
5 0.400
6 0.762
6 1.124
6 1.102
6 1.436
6 1.967
6 1.047
5 0.283
11 0.363
11 0.316
11 0.526
10 0.571
11 0.310
11 0.399
11 0.427
11 0.807
11 1.176
11 1.155
11 1.558
10 2.076
10 1.061
11 0.314
6 0.361
6 0.321
6 0.512
6 0.557
6 0.337
6 0.419
6 0.429
6 0.833
6 1.189
6 1.206
6 1.612
6 2.090
6 1.064
6 0.306
30
0.35
0.29
0.49
0.53
0.29
0.37
0.39
0.70
1.09
1.02
1.38
1.89
1.02
0.28
4.32 0.208
5.01 0.222
6.04 0.172
6.27 0.126
5.38 0.134
4.34 0.121
3.57 0.090
0.37 0.008
0.33 0.014
0.53 0.014
0.57 0.013
0.32 0.011
0.41 0.016
0.41 0.010
0.83 0.045
1.17 0.033
1.17 0.052
1.49 0.040
2.01 0.045
1.10 0.029
0.29 0.008
SD
11 14.7614.4215.27 0.313
11 14.3713.9714.73 0.300
11 9.54 9.20 9.86 0.228
11 3.94 3.68 4.08 0.137
11 4.55 4.33 4.88 0.180
11 7.75 7.58 7.98 0.108
10 8.23 7.93 8.56 0.199
11 4.45 4.20 4.73 0.169
11 5.73 5.52 5.94 0.126
11
11
11
11
11
11
11
4.21
5.12
6.13
6.12
5.21
4.32
3.54
4.08
4.96
6.00
5.95
5.08
4.20
3.41
0.36
0.30
0.51
0.55
0.30
0.39
0.41
0.76
1.13
1.08
1.49
1.98
1.04
0.30
4.39 0.120
5.27 0.115
6.23 0.078
6.27 0.104
5.35 0.089
4.48 0.091
3.73 0.100
0.37 0.004
0.33 0.010
0.54 0.009
0.59 0.011
0.32 0.009
0.41 0.006
0.44 0.009
0.85 0.025
1.20 0.020
1.23 0.043
1.66 0.050
2.17 0.054
1.08 0.018
0.33 0.011
n
M min max
n
3.74
4.49
5.61
5.94
5.02
3.97
3.35
n
M min max
3.94
4.73
5.81
6.08
5.16
4.16
3.44
SD
E. go­bi­en­sis
N Kirghizia
6
6
6
6
6
6
6
4.64
5.45
6.63
6.63
5.58
4.54
3.67
4.52
5.31
6.47
6.47
5.38
4.41
3.58
0.35
0.31
0.50
0.54
0.32
0.41
0.42
0.80
1.15
1.15
1.52
2.00
1.04
0.29
SD
4.83 0.110
5.61 0.108
6.78 0.116
6.74 0.105
5.71 0.115
4.64 0.094
3.77 0.061
0.37 0.009
0.34 0.012
0.52 0.009
0.57 0.009
0.35 0.012
0.43 0.008
0.44 0.005
0.87 0.024
1.23 0.029
1.30 0.059
1.70 0.061
2.21 0.080
1.10 0.027
0.33 0.013
Table 2. (continued)
Tab. 2. (pokračování)
E. nilssonii
C Eu­ro­pe
n
M min max
E. n. na­su­tus
Baluchestan
SD
n
M min max
E. n. pellucens
Mesopotamia
SD
n
M min max
SD
LAt
11 40.09 38.3 42.7 1.448
9 38.09 35.7 39.9 1.205
3 36.27 35.5 37.2 0.862
LCr
LCb
LaZ
LaI
LaInf
LaN
LaM
ANc
ACr
25 15.3514.8715.92 0.312
24 14.9914.4415.50 0.320
19 10.04 9.5110.44 0.238
25 4.05 3.71 4.28 0.173
25 4.91 4.62 5.27 0.194
25 7.91 7.57 8.38 0.229
25 8.54 8.21 8.84 0.210
25 5.19 4.96 5.54 0.151
24 6.64 6.23 6.88 0.161
8 13.0512.6513.39 0.255
8 12.7112.2813.18 0.317
6 8.79 8.53 8.98 0.151
8 2.99 2.75 3.13 0.123
8 4.30 4.02 4.43 0.131
8 6.30 6.17 6.42 0.075
8 6.90 6.75 7.08 0.105
8 4.53 4.42 4.75 0.127
8 5.68 5.49 5.87 0.139
6 13.2412.8813.95 0.393
6 12.9112.4813.65 0.443
5 8.87 8.39 9.83 0.564
6 2.89 2.69 3.13 0.165
6 4.40 4.02 5.09 0.396
5 6.34 6.00 6.64 0.269
6 7.15 6.91 7.54 0.249
6 4.52 4.34 4.74 0.158
6 5.57 5.36 5.82 0.169
5.11 0.153
5.86 0.160
6.63 0.191
6.88 0.142
5.74 0.120
4.61 0.129
3.73 0.082
8
8
8
8
8
8
8
4.24
4.85
5.90
5.61
4.88
4.00
3.38
3.95
4.68
5.68
5.23
4.42
3.63
2.97
4.40 0.167
4.94 0.101
6.17 0.175
5.84 0.210
5.13 0.227
4.16 0.175
3.64 0.209
6
6
6
6
6
6
6
4.34
4.98
6.01
5.69
4.81
4.00
3.40
4.04
4.66
5.68
5.42
4.64
3.82
3.27
25 11.0510.7311.44 0.208
24 3.30 3.03 3.57 0.139
25 6.98 6.61 7.28 0.162
25 5.97 5.65 6.31 0.150
25 4.67 4.42 4.92 0.118
25 3.96 3.75 4.22 0.111
8
8
8
8
8
8
9.31
3.20
5.64
5.13
4.13
3.58
8.64
3.08
5.21
4.64
3.72
3.33
9.59 0.300
3.37 0.098
5.91 0.233
5.40 0.252
4.45 0.225
3.78 0.163
6
6
6
6
6
6
9.53
3.28
5.69
5.16
4.16
3.59
9.1110.32 0.421
3.17 3.43 0.105
5.42 6.29 0.340
4.82 5.76 0.347
3.93 4.61 0.263
3.35 4.03 0.238
8 0.384
8 0.339
8 0.483
8 0.543
8 0.356
8 0.447
8 0.464
8 0.868
8 1.209
8 1.442
8 1.977
8 2.310
8 1.095
8 0.344
0.36
0.33
0.47
0.53
0.34
0.42
0.45
0.84
1.17
1.37
1.85
2.23
1.08
0.33
0.40 0.012
0.35 0.007
0.50 0.009
0.56 0.010
0.38 0.014
0.47 0.015
0.48 0.013
0.89 0.019
1.29 0.035
1.60 0.082
2.24 0.127
2.48 0.091
1.13 0.017
0.36 0.009
6 0.372
6 0.341
5 0.478
6 0.554
6 0.351
6 0.432
6 0.466
6 0.904
6 1.251
6 1.525
6 2.085
6 2.480
5 1.123
6 0.344
CC25
P4P4
25
M3M3
25
I1M3
25
CM3
25
P4M3
25
M1M3
25
LMd
ACo
I1M3
CM3
P4M3
M1M3
4.84
5.50
6.39
6.63
5.54
4.40
3.57
CM3/LCb 24 0.370
LaInf/LCb 24 0.328
LaN/LCb 25 0.515
LaM/LCb 24 0.570
ANc/LCb 24 0.346
ACr/LCb 23 0.444
M3M3/LCb 24 0.427
CC/CM3 25 0.875
M3M3/CM3 25 1.153
LaInf/LaI 25 1.213
M3M3/LaI 25 1.577
LaM/LaI 25 2.111
LaM/LaN 25 1.081
ACo/LMd 24 0.299
4.51
5.26
6.02
6.36
5.21
4.02
3.38
0.36
0.30
0.49
0.55
0.33
0.43
0.41
0.83
1.11
1.09
1.45
1.97
1.04
0.27
0.39 0.006
0.35 0.011
0.53 0.010
0.59 0.008
0.36 0.008
0.47 0.009
0.45 0.009
0.92 0.026
1.21 0.027
1.33 0.062
1.75 0.067
2.34 0.086
1.13 0.020
0.32 0.012
0.37
0.32
0.47
0.53
0.33
0.42
0.45
0.85
1.20
1.40
1.97
2.40
1.07
0.33
4.74 0.258
5.47 0.274
6.58 0.319
6.12 0.271
5.18 0.209
4.39 0.211
3.74 0.181
0.38 0.006
0.37 0.020
0.49 0.009
0.57 0.013
0.37 0.015
0.45 0.009
0.48 0.012
0.95 0.036
1.29 0.039
1.67 0.102
2.23 0.093
2.63 0.087
1.15 0.030
0.37 0.012
31
cluster differing from the cluster of specimens of E. nasutus and the cluster of samples of E.
nilssonii and E. gobiensis.
The same or very similar results were obtained also by the comparisons of skull indices (Figs.
4, 5). Two indices describing the shape of braincase (relative width, LaN/LCb, vs. re­la­ti­ve height,
ANc/LCb) grouped the compared samples into three clusters (Fig. 4); (1) Qutur Su bats and
E. bobrinskoi (relatively low and wide braincase), (2) E. nilssonii and E. go­bi­en­sis (relatively
high and wide braincase), and (3) E. nasutus (relatively high and narrow bra­in­ca­se). Two indices describing the shape of rostrum (relative length, I1M3/LCb, vs. relative width, CC/CM3)
grouped the samples into two main clusters (Fig. 5); (1) Qutur Su bats, E. bobrinskoi and E.
gobiensis (relatively short and narrow rostrum), and (2) E. nasutus and E. nilssonii (relative
long and wide rostrum). However, the Qutur Su bats, having a relatively shortest and narrowest
rostrum within the compared samples, showed to be most similar to the samples of E. bobrinskoi
(together making the only dimensional overlap), while the other samples displayed a relatively
much wider and mostly also longer rostrum.
Finally, the analysis of principal components (PCA) showed very similar results as the abo­ve
mentioned comparisons (Fig. 6). The PCA of all 38 skull dimensions (PC1 56.84% of variance,
PC2 20.42%) selected a group of dimensions with a higher significance for va­ri­an­ce (>70%)
within the compared set of samples; all measurements (except for ACo) and the folowing indices:
LaInf/LCb, ANc/LCb, ACr/LCb, CC/CM3, M3M3/LaI, LaM/LaI, and ACo/LMd. The PCA of
these 28 dimensions (PC1 65.85% of variance; PC2 19.00%) clearly se­pa­ra­ted three clusters of
samples (Fig. 6); (1) Qutur Su bats and E. bobrinskoi (PC2>0.55), (2) E. nasutus (PC1< –0.5),
and (3) E. nilssonii and E. gobiensis (PC1>0.1; PC2<0.55).
Fig. 4. Scatter plot of the relative width of neurocranium (LaN/LCb) against the relative height of neu­ro­
cra­ni­um (ANc/LCb) in the compared samples of smaller Eptesicus species of the western Palaearctic.
Obr. 4. Srovnání relativní šířky mozkovny (LaN/LCb) proti relativní výšce mozkovny (ANc/LCb) jedinců
malých druhů rodu Eptesicus západní Palearktidy.
32
Fig. 5. Scatter plot of the relative length of rostrum (I1M3/LCb) against the relative width of rostrum
(CC/CM3) in the compared samples of smaller Eptesicus species of the western Palaearctic.
Obr. 5. Srovnání relativní délky rostra (I1M3/LCb) proti relativní šířce rostra (CC/CM3) jedinců malých
druhů rodu Eptesicus západní Palearktidy.
Fig. 6. Results of the principal component analysis of skull dimensions in the compared samples of smaller
Eptesicus species of the western Palaearctic. For details see Results.
Fig. 6. Výsledky analysy hlavních proměnných lebečních rozměrů jedinců malých druhů rodu Eptesicus
západní Palearktidy.
33
All presented morphometric comparisons showed the Qutur Su bats to be very similar in their
skull characters to the samples of E. bobrinskoi from central Kazakhstan, and clearly di­stin­guis­hed them from all other comparative samples of Eptesicus from the western Pa­lae­ar­c­tic.
Fig. 7. Skulls of small representatives of the genus Eptesicus of the western Palaearctic; in dorsal views.
Scale bar = 5 mm.
Obr. 7. Lebky malých zástupců rodu Eptesicus západní Palearktidy; v dorsálním pohledu. Měřítko =
5 mm.
Legend / legenda: a – E. bobrinskoi (NMP 90890, female, Qutur Su, NW Iran); b – E. bobrinskoi (ZIN
61694, male, Aryskumy Desert, C Kazakhstan); c – E. nasutus (NMP 48404, female, Pir Sohrab, Ba­lu­
ches­tan, SE Iran); d – E. gobiensis (CUP CT84/25, female, Ala-Arča NR, N Kirghizia); e – E. nilssonii
(NMP 91122, female, Vrbno near Blatná, SW Bohemia, Czech Republic).
34
Figs. 7, 8 show examples of skulls of the five compared Eptesicus populations and/or taxa. The
dif­f­e­ren­ces among the morphotypes resulting from the above analyses are well ob­ser­va­ble also
in these drawings. Besides the distinctions in dimensions (Table 2), the par­ti­cu­lars in the shapes
of braincase and rostrum in the Qutur Su bats and E. bobrinskoi (an ex­tre­me­ly low braincase,
a distinct areal ratio between frontal and parietal parts of the braincase, a relatively small external auditory orifice, flattened frontal bones, narrow zygomatic arches, relative slender teeth,
a flat­te­ned facial part of the skull, a relatively very narrow mesial part of the rostrum, shapes
of supraorbital ridges, shapes of orbital processes of the zygomata) clearly differ from other
com­pa­red bats. E. nasutus is most distinct, in comparison with the previous morphotype it has
relatively much wider zygomatic arches but a much narrower bra­in­ca­se with smaller frontal and
larger parietal bones, much narrower zygomata, a more mas­si­ve lambda, more massive teeth
(and of course, a unicuspidal first upper incisor) and a very short and high ante-orbital part of
the rostrum with distinct supracanine swellings. Some of these differences were also observed
by Harrison (1963), see this paper also for distinctions in some external features. The skulls
of two very similar morphotypes, E. nilssonii and E. gobiensis, differ from the Qutur Su bats
and E. bobrinskoi mainly in the relatively and ab­so­lu­te­ly much higher braincase and rostrum,
distinct frontal concavities, a relatively large ex­ter­nal auditory orifice, relatively wider mesial
Fig. 8. Skulls of small representatives of the genus Eptesicus of the western Palaearctic; in lateral views.
Scale bar = 5 mm. For legend see Fig. 7.
Obr. 8. Lebky malých zástupců rodu Eptesicus západní Palearktidy; v laterálním pohledu. Měřítko =
5 mm. Legenda viz obr. 7.
35
parts of the rostrum, more developed orbital processes of the zygomata and more massive
teeth (Figs. 7, 8). As stressed above, these two pairs of morphotypes also clearly differ in their
lengths of forearm (Table 2).
A baculum was extracted (Fig. 9a) from the only male among the newly collected Qutur Su
bats (NMP 90892). It is a small bone, having a shape of a ‘cognac bottle’ in the dorsal aspect,
0.70 mm long and 0.45 mm broad in most wide (proximal) part; the constriction of the ‘bot­t­le-neck’ is 0.17 mm broad. The baculum of the Qutur Su series was earlier described by Harrison
(1963: 307) [No. 5] and Hill & Harrison (1987: 296) [BMNH 63.1187], however, drawings of
these descrip­ti­ons differ. The newly prepared baculum rather corresponds in its shape and size
to that de­pic­ted by Hill & Harrison (1987), and also to one of the preparations of E. bobrin­
skoi from central Kazakhstan published by Strelkov (1986, 1989) (Fig. 9). Strel­kov (1989)
gave the following length and width ranges of baculum in E. bobrinskoi: 0.62–0.72 mm, and
0.37–0.50 mm, re­spe­cti­ve­ly. However, the bacula of the Qutur Su bats (Har­ri­son 1963, Hill &
Harrison 1987, own data) as well as of E. bobrinskoi (Strelkov 1986, 1989) resemble in their
Fig. 9. Bacula of small representatives of the genus Eptesicus of the western Palaearctic; in dorsal views.
Scale bar = 1 mm. a – original, b – after Harrison (1963), c, g – after Hill & Harrison (1987), d–f and
h–q – after Strelkov (1989).
Obr. 9. Bakula malých zástupců rodu Eptesicus západní Palearktidy; v dorsálním pohledu. Měřítko =
1 mm. a – originál, b – podle Harrisona (1963), c, g – podle Hilla & Harrisona (1987), d–f, h–q – podle
Strelkova (1989).
Legend / legenda: a–c – E. bobrinskoi (Qutur Su, Iran); d–f – E. bobrinskoi (Kazakhstan); g, h – E.
nasutus (Oman, ‘Near East’); i–k – E. nilssonii (Russia, Mongolia); l–q – E. gobiensis (Mongolia, East
Turkestan, Tajikistan).
36
shape (but not in size) also those of E. gobiensis (Strelkov 1986, 1989), for which Strelkov
(1986) gave the dimension ranges 1.20–1.60 mm and 0.72–0.90 mm, respectively. On the other
hand, the bacula of E. nils­so­nii and E. nasutus com­ple­te­ly differ both in their size and shape
from the above taxa (Topál 1958, Hill & Harrison 1987, Strelkov 1989; Fig. 9).
It seems to be clear from the above gathered arguments that the series of the Qutur Su bats,
both that published by Harrison (1963) and that newly collected by us, belong to the only
morphotype. Moreover, this morphotype was found to be identical in many aspects with that of
the samples of E. bobrinskoi from central Kazakhstan. The original identification of the Qutur
Su bats as E. bobrinskoi by Harrison (1963) appears to be proved as well as the occur­ren­ce of
this species in the Middle East.
However, the most important argument from the findings by Hanák & Horáček (1986) (see
Introduction) remains, considering the geographical isolation of the region of NW Iran from
the core distribution area of E. bobrinskoi, i.e. the western and central parts of Ka­za­kh­stan (e.g.
Strelkov 1980). The shortest distance between Qutur Su and the westernmost Ka­za­kh site
(near Kosčagyl, which is, however, 400 km west of the core area, see above) is about 1100 km,
taken across the Caspian Sea. Taken round this lake, the distance can be some 1800 km along
its eastern coast or 1400 km along the western coast. However, no records of E. bob­rin­skoi are
known either in the similar landscape of eastern Azerbaijan on the western coast or in the deserts
of Turkmenistan on the eastern coast (Strelkov et al. 1978, Rahmatulina 2005).
It should be mentioned that the habitats of these distant regions are quite different. The Kazakh habitats of occurrence of E. bobrinskoi are situated in the zone of northern lowland deserts
and semi-deserts (Butovskij et al. 1985), while the Persian locality lies in the high mon­ta­ne
zone of alpine meadows in the altitude of around 2500 m (Figs. 1, 2). These regions represent
quite distinct landscape types, sharing only some general features as the absence of trees and
presumably also a harsh continental climate. The occurrence of one species in two restricted
regions differing by their habitats remains enigmatic. A profound faunal in­ves­ti­ga­ti­on in the
areas between them, i.e. in the northern parts of the Middle East as well as in Transcaucasia
and West Turkestan, is needed to shed light on this phenomenon.
However, several isolated records assigned to the smaller species of Eptesicus from the Mid­
dle East and adjacent areas have been discussed and/or doubted in the literature (see Ha­nák
& Horáček 1986, Nader & Kock 1990). It is possible that some of these records may pertain
to E. bobrinskoi.
Only two records of E. nilssonii are known from Transcaucasia*; both were published by
K. A. Satunin at the end of the 19th century and were tentatively assigned to E. n. nilssonii
by Hanák & Horáček (1986). The closer record to Qutur Su comes from the approximately
70 km distant Viljaž-Čaj River near Lenkoran in south-eastern Azerbaijan, where E. nilssonii
was found in 1888 (Satunin 1910). Although this record has been widely considered beyond
any doubt (Ognev 1927, 1928, Kuzjakin 1950, 1965, Vereščagin 1959, Strelkov 1963, Ha­
nák & Horáček 1986, Rahmatulina 1990, 2005), it may well be a misidentified spe­ci­men.
The regions of south-eastern Azerbaijan are covered by similar habitat types as the ad­ja­cent
parts of Iran, and even by hot deserts in the lowland areas of the lower Kura River. Al­thou­gh
* Ševčenko & Zolotuhina (2005) mentioned a third record of E. nilssonii from Transcaucasia: adult male,
Georgia, Teberda, 5 July 1978, leg. E. Javrujan. However, the site Teberda lies on the northern slope of the
Greater Caucasus in the Kabardino-Balkarskaja Republic, Russia, and not in Georgia. Thus, this record
belongs to the main distribution range of E. nilssonii in the Caucasus Region (see Hanák & Horáček
1986) and not to the Transcancasianone.
37
E. nilssonii, rather a boreal bat species, rarely occurs in the Greater Caucasus Range (see the
review by Hanák & Horáček 1986), south-eastern Transcaucasia belongs to a distinct faunal
zone isolated by semi-deserts and steppes from the north (Rakhmatulina 1995). The second
Transcaucasian specimen of E. nilssonii was reported by Satunin (1896) from Tiflis (= Tbilisi,
Georgia), however, this isolated record seems to be more probable from the zo­o­ge­o­gra­phi­cal
point of view than the former one. Since both individuals are kept in scientific collections, see
Bukhnikashvili et al. (2004) and Rahmatulina (2005), these records could be revised and their
species identification confirmed or disproved, respectively. In question of a possible confusion
still remains also the isolated record assigned to E. nilssonii go­bi­en­sis (= E. gobiensis) by Lay
(1967) and DeBlase (1980), collected at Sama in the Elborz Mts, N Iran. According to DeBlase
(1980), this specimen has a relatively large skull (LCr 15.9 mm) but a relatively short forearm
(LAt 37.7 mm), i.e. ratio of these dimensions in a state close to that found in E. bobrinskoi
(see above). However, this record was accepted by Hanák & Horáček (1986) under DeBlase’s
(1980) species affiliation.
Nevertheless, a real example of confusion of species identification in the genus Eptesicus
in the Middle East is a report of ‘Eptesicus nilssoni nilssoni’ by Hatt (1959) from Baghdad,
Iraq, i.e. from a semi-desert lowland region of Mesopotamia. This record was accepted under
Hatt’s (1959) species affiliation by several authors (Al-Robaae 1966, Lay 1967, Allison &
Gaisler 1982, Hanák & Horáček 1986, Rydell 1993, Duff & Lawson 2004, Simmons 2005),
although Harrison (1972) re-identifed the respective specimen as E. bottae (Peters, 1869) (see
also Nader & Kock 1990 and Harrison & Bates 1991). Similarly, Pocock (1935: 442) determined a small individual of Eptesicus from Shanna, Saudi Arabia, as “a specimen ap­pa­rent­ly
referable to Eptesicus hingstoni [= E. bottae hingstoni Thomas, 1919]”, while later on this bat
was re-examined as E. nasutus (Nader & Kock 1990, Harrison & Bates 1991, this paper). An
individual of E. anatolicus Felten, 1971 from Mala-i-Mir (= Izeh), SW Iran, was first reported
by Tho­mas (1906) under the name “Vespertilio sp., near V. serotinus”. Later on, it was considered to pertain to E. bottae by Lay (1967) and Etemad (1969) and to E. serotinus shiraziensis
(Dob­son, 1871) by Gaisler (1970). Finally, it was correctly identified by DeBlase (1980); see
also the analysis by Benda et al. (2006). Other examples of confusion in species identification
within the genus Ep­te­si­cus are mentioned by DeBlase (1980) and Nader & Kock (1990).
In conclusion, we have confirmed the occurrence of E. bobrinskoi in Iran, which seems to be
isolated from the main distribution range of the species for the time being. However, the extent
of this isolation may prove to be smaller after all specimens available are revised and/or more
profound faunal explorations are made in the areas filling the geographical gap in between
these isolated spots.
SOUHRN
Netopýr turanský (Eptesicus bobrinskoi Kusjakin, 1935) je obyvatelem především stepí a po­lo­pouš­tí se­ver­
ní části Turanské nížiny mezi Aralským jezerem a Hladovou pouští (zvanou též Betpak-Dala) ve středním
Kazachstanu (Butovskij et al. 1985) – z této oblasti pochází asi 90 % všech nálezů druhu. Několik nálezů
tohoto netopýra bylo však hlášeno i z jiných oblastí: ze Severní Osetie (severní svah Velkého Kavkazu)
a z Jakutska (Kuzjakin 1950), ze se­ve­ro­zá­pad­ní­ho Iranu (Harrison 1963) a ze zá­pad­ní­ho a severozápadního Kazachstanu (Strelkov 1980, Da­vy­go­ra et al. 1998). Zatímco poslední (ka­zaš­ské) nálezy jsou bez
výhrad akceptovány, zpo­chyb­něn byl kavkazský, jakutský a perský nález, přičemž bylo souzeno – většinou
zřejmě právem – že se jedná nejspíše o juvenilní jedince netopýra severního, E. nilssonii (Keyserling et
Blasius, 1839) (Hanák & Horáček 1986).
38
Autorům se podařilo v červnu 2006 navštívit lázeňské letovisko Qutur Su (38° 20’ N, 47° 51’ E;
2545 m n. m) v severozápadním Iranu, jedinou známou lokalitu sběru E. bobrinskoi v Iranu a na Blízkém
vý­cho­dě. Jedinci zde sebraní v roce 1961 jsou uloženi v londýnském Přírodovědeckém museu, jejich nález
a určení bylo zveřejněno Harrisonem (1963); později však byli přeurčeni jako mladí jedinci E. nilssonii
(Hanák & Ho­rá­ček 1986). Autoři tohoto příspěvku na téže lokalitě nalezli tři mrtvé kusy drobných netopýrů rodu Eptesicus, zjevně však nikoli E. nilssonii. Proto byli jedinci určení Harrisonem (1963), jakož
i nově nalezení, zevrubně porovnáni se srovnávacím materialem všech malých netopýrů rodu Ep­te­si­cus,
obývajících západní Pa­leark­ti­du: E. bob­rin­skoi, E. nilssonii, E. gobiensis a E. nasutus. Z tohoto srovnání
jasně vyplynulo, že zmíněné dvě serie (celkem 10 kusů) netopýrů z Qutur Su představují nepochybně
jedince E. bobrinskoi.
Potvrzení výskytu netopýra turanského, který představuje zejména faunový prvek nížinných stepí
a po­lo­pouš­tí, v severozápadním Iranu, navíc na lokalitě ležící v alpinském velehorském pásmu, je velmi
překvapivý. Lokalita nálezu v Iranu je nejméně 1400 km vzdálená od nejbližšího místa potvrzeného nálezu
E. bobrinskoi v Kazachstanu. V příspěvku je diskutována možnost nesprávných určení některých nálezů
rodu Eptesicus v oblasti vyplňující tuto mezeru (Satunin 1896, 1910, Lay 1967). V případě, že některé
z nich ve skutečnosti představují nález netopýra turanského, jeho známý výskyt v Iranu by nemusel být
natolik isolován, výskyt druhu v alpinské poloze velehor však záhaným zůstává nepochybně.
ACKNOWLEDGEMENTS
We thank Jiří Mlíkovský for his help in the field. We are obliged to Pauline Jenkins & Daphne Hill (BMNH)
for enabling us to revise the Persian material of E. bobrinskoi (cf. Harrison 1963) as well as of other
bats deposited in the BMNH collection, to Petr P. Strelkov (ZIN) for providing us with the comparative
samples of E. bobrinskoi from Kazakhstan, and to Ivan Horáček (CUP) for enabling us to compare the
material of E. gobiensis. We thank Ivan Horáček also for valuable comments on the topics.
This study was supported by the Grant Agency of the Czech Republic (grants Nos. 206/02/D041
and 206/05/2334) and the Ministry of Culture of the Czech Republic (grants Nos. RK01P03OMG006,
DE06P04OMG008, and MK00002327201).
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Thomas O., 1905: A new genus and two new species of bats. Ann. Mag. Natur. Hist., (7)16: 573.
Thomas O., 1906: On a collection of mammals from Persia and Armenia presented to the British Museum
by Col. A. C. Bailward. Proc. Zool. Soc. London, 1905(2): 519–527.
Thomas O., 1919: Some new mammals from Mesopotamia. J. Bombay Natur. Hist. Soc., 26: 745–749.
Topál G., 1958: Morphological studies on the os penis of bats in the Carpathian Basin. Ann. Hist.-Natur.
Mus. Natl. Hungar., 50: 331–342.
Vereščagin N. K., 1959: Mlekopitajuščie Kavkaza. Istorija formirovanija fauny [The Mammals of the
Caucasus. The History of Forming of the Fauna]. Izdatel’stvo Akademii Nauk SSSR, Moskva & Leningrad, 704 pp (in Russian).
Zukal J. & Gaisler J., 1991: K výskytu a změnám početnosti netopýra severního, Eptesicus nilssoni (Key­
ser­ling et Blasius, 1839) v Československu [On the occurrence and changes in abundance of Eptesicus
nilssoni (Keyserling et Blasius, 1839) in Czechoslovakia]. Lynx, n. s., 25[1989]: 83–95 (in Czech, with
a summary in English).
APPENDIX
List of the material examined
Eptesicus nilssonii (Keyserling et Blasius, 1839)
Czech Republic (19). 1 ma (NMP 91133 [S]), Dlouhá Ves, Franz-Franz Mine, Šumperk Dist., 30 Ja­nu­a­ry
1959, leg. V. Hanák (cf. Gaisler & Hanák 1972); – 1 fa (NMP 91144 [S+B]), Malé Karlovice, Tísňavy,
school attic, Vsetín Dist., 5 June 1973, leg. V. Bejček (cf. Bejček 1975); – 3 ma, 1 ms, 2 fa, 1 fs (NMP
91136, 91138, 91139 [S+B], 91123, 91124, 91126, 91127 [S]), Mariánská Hora, gallery near the Bílá
Desná Dam, Jablonec nad Nisou Dist., 24 February 1958, 13 February 1962, 2 December 1964, leg.
V. Hanák (cf. Ne­vr­lý 1963, Gaisler & Hanák 1972); – 1 ma (NMP 91128 [S]), Mikulov, Teplice Dist.,
13 March 1958, leg. V. Hanák (cf. Gaisler & Hanák 1972); – 1 ma (NMP 91146 [S+B]), Orlické Záhoří,
Rychnov nad Kněžnou Dist., 10 February 1977, leg. P. Rybář (cf. Bárta et al. 1981); – 1 fa (NMP 91152
[S]), Pohorská Ves, Žofín, Český Krumlov Dist., 16 June 1973, leg. V. Vohralík (cf. Anděra & Hanák
43
2004); – 1 fa (NMP 91140 [S]), Rokytnice v Orlických horách, Hanička Fortress, Rychnov nad Kněžnou
Dist., 22 January 1965, leg. J. Sklenář (cf. Sklenář 1969, Gaisler & Hanák 1972); – 1 ma (NMP 91132
[S+B]), Suchá Rudná, mine, Bruntál Dist., 30 January 1959, leg. V. Hanák (cf. Souček 1970); – 1 ind.
a (NMP 91151 [S]), Šumava Mts (SW Bohemia), leg. J. Červený; – 2 fa (NMP 91121, 91122 [S+B]),
Vrbno near Blatná, Strakonice Dist., 4 and 5 June 1956, leg. V. Hanák (cf. Hanák 1959); – 1 ma, 1 ms
(NMP 91130, 91131 [S]), Zlaté Hory, Poštovní Mine, Bruntál Dist., 29 January 1959, leg. V. Hanák (cf.
Gaisler & Hanák 1972).
Slovakia (5). 2 fa (NMP 91135 [S+B], 91134 [S]), Demänovská Dolina, Dračia jaskyňa [= Demänovská
Ice Cave], Liptovský Mikuláš Dist., 14 February 1961, leg. V. Hanák (cf. Gaisler & Hanák 1972); – 1 ma,
1 ms (NMP 91142, 91143 [S+B]), Dobšiná, Dobšinská Ice Cave, Rožňava Dist., 16 February 1968, leg.
V. Hanák (cf. Gaisler & Hanák 1972); – 1 ma (NMP 91145 [S+B]), Tatranská Javorina, Muránska jaskyňa
Cave, Poprad Dist., 13 December 1973, leg. J. Gaisler & V. Hanák (cf. Zukal & Gaisler 1991).
Eptesicus gobiensis Bobrinskoj, 1926
Kirghizia (6). 6 fa (CUP CT84/24–29 [S+A]), Ala-Arča Nature Reserve, 1990–2000 m a. s. l., ca. 40 km
S of Biškek, 30 June 1984, leg. J. Červený & I. Horáček.
Eptesicus bobrinskoi Kusjakin, 1935
Iran (10). 3 ma, 2 ms, 2 fa, 3 fs (BMNH 63.1186–1192, NMP 90890–90892 [S+A]), Qutur Su (N slope
of the Mt. Sabalan, 2545 m a. s. l.), ca. 20 km ESE of Meshgin Shahr, Ardabil Prov., 21 August 1961, leg.
Aberystwyth University Expedition (cf. Harrison 1963), 5 June 2006, leg. P. Benda & A. Reiter.
Kazakhstan (11). 1 ma (ZIN 61694 [S+B]), Aryskumy Desert, 27 km SSE of Mustafa, 160 km N of Kzyl-Orda, Kzyl-Ordinskaja Region, 25 July 1974, leg. I. I. Stogov (cf. Strelkov 1980); – 1 fa (ZIN 62247
[S+B]), 10 km NW of Čelkar, Aktjubinskaja Region, 18 June 1975, leg. P. P. Strelkov (cf. Strelkov 1980);
– 1 ma, 1 fa (ZIN 65104, 65121 [S+B]), between Džilandy and Kense, Sarysu River, Žetykanur Desert,
120 km SSE of Džezkazgan, Džezkazganskaja Region, 13 June 1977, leg. P. P. Strelkov (cf. Strelkov
1980); – 1 ma (ZIN 68618 [S+B]), Karakum Meteorologic Station, Bokdok Valley, 180 km N of Džusaly,
Džezkazganskaja Region, 21 June 1980, leg. P. P. Strelkov (cf. Strelkov & Šajmardanov 1983); – 6 fa
(ZIN 62240–62242, 62244–62246 [S+B]), Žetybaj well, 150 km N of Kzyl-Orda, Kzyl-Ordinskaja Region,
5 & 8 June 1975, leg. P. P. Strelkov (cf. Strelkov 1980).
Eptesicus nasutus (Dobson, 1877)
Afghanistan (1). 1 ma (BMNH 68.475 [S+B]), Bisut, nr. Jalalabad, 34° 26’ N, 70° 25’ E, 7 April 1967,
leg. J. Gaisler (cf. Gaisler 1970).
Iran (11). 3 m (BMNH 5.10.4.2, 5.10.4.4 [holotype of Vespertilio matschiei pellucens Thomas, 1906],
5.10.4.6 [S+B]), Ahwaz, Karun River, Arabistan [= Khuzestan Prov.], 220 ft, 28 March 1905, leg. R. Wo­
os­nam (cf. Thomas 1906); – 1 ma, 2 fa (NMP 48437, 48438 [S+A], JOC [pb1722] [Sk]), 15 km E of
Dehbarez, ca. 22 km W of Manujan, Kerman Prov., 17 April 2000, leg. P. Benda & A. Reiter; – 1 ma,
4 fa (NMP 48404–48408 [S+A]), Pir Sohrab, ca. 60 km NE of Chabahar, Baluchestan-ve-Sistan Prov.,
12 Ap­ril 2000, leg. P. Benda & A. Reiter.
Iraq (2). 1 fa (BMNH 19.3.1.2 [S+A]; holotype of Eptesicus walli Thomas, 1919), Basra, leg. F. Wall
(cf. Thomas 1919); – 1 m (BMNH 36.4.14.13 [S+B]), Zubier, Mesopotamia, 2 March 1921, leg. R. E.
Che­e­sman (cf. Harrison 1964).
Oman (1). 1 fa (BMNH 68.1356 [S+B]; holotype of Eptesicus nasutus batinensis Harrison, 1968), Har­
mul, 10 mls N of Sohar, 26 March 1967, leg. C. Seton-Browne (cf. Harrison 1968);
Saudi Arabia (2). 1 ma (BMNH 48.350 [S+B]), nr. Jedda, 200 ft., 4 July 1948, leg. G. B. Popov (cf.
Har­ri­son 1964); – 1 ind. a (BMNH 34.8.8.1 [S+A]), Shanna, Arabia, [22 February], leg. J. Philby (cf.
Pocock 1935).
Yemen (1). 1 m (BMNH 99.11.6.19. [S+B]; holotype of Vespertilio matschiei Thomas, 1905), Jimel,
W. Aden, 850 m, 16 August 1899, leg. W. Dodson (cf. Thomas 1905).
44
Lynx (Praha), n. s., 37: 45–50 (2006).
ISSN 0024–7774
Karyotype of the Small-eared dormouse (Graphiurus microtis) from
the Nyi­ka Plateau, Malawi (Rodentia: Gliridae)
Karyotyp plcha malouchého (Graphiurus microtis) z náhorní plošiny Nyika, Malawi
(Rodentia: Gliridae)
Hynek Burda1 & Wilbert N. Chitaukali2
1
Department of General Zoology, Institute of Biology, University of Duisburg-Essen,
D–45317 Essen, Germany; [email protected]
2
Biology Department, Chancellor College, Uni­ver­si­ty of Malawi, P. O. Box 280, Zomba, Malawi
received on 4 December 2006
Abstract. Karyotype of the African Small-Eared Dormouse (Graphiurus microtis) from Nganda (Nyika
Plateau, Malawi) consisted of 2n=52 chromosomes: 11 pairs of metacentrics, 14 pairs of acrocentrics and
subtelocentrics, the X-chromosome being a metacentric, Y-chromosome a small acrocentric. It represents
a new karyotype reported for African dormice (Graphiurinae). Regarding the scarity of data (thus far ka­
ry­o­typ­es of only four species representing five populations, including the present data, out of currently
re­co­gni­zed 13–14 species, distributed throughout Sub-Saharan Africa have been described) it is worth of
being published but any efforts to derive conclusions regarding chromosomal speciation in graphiurines
would be preliminary and speculative.
Introduction
Within the Gliridae, the genus Graphiurus Smuts, 1832 is unique in being an African en­de­mic
widely distributed from south of the Sahara to the Cape Province in South Africa, and in being
the most specious genus of the family (13–14 species are currently recognized, whe­re­as other
glirid genera consist of a maximum of three species; Holden 1993, Rossolimo et al. 2001).
Gra­phiu­rus constitutes one of the three major lineages identified among the Gli­ri­dae, justifying
its subfamilial rank. The lineage is assumed to have radiated rapidly, following colonization of
Africa at ca. 8–10 Myr ago (Montgelard et al. 2003). It is assumed that high rate of speciation
was driven by adaptive radiation into diverse habitats ranging from dense forests to rocky areas
(Montgelard et al. 2003). Still, the genus appears to be mor­pho­lo­gi­cal­ly quite homogeneous
(Genest-Villard 1978, Holden 1996), and the question arises whe­ther speciation may have
been driven by and/or is reflected in, chromosomal diversification, as for instance is assumed
to be the case in the adaptive radiation of blind mole-rats (Spalax Guldenstaedt, 1770 spp.; cf.
Nevo et al. 2001). So far karyotypes, ranging from 2n=40 to 2n=70 chromosomes, of only four
species, representing five populations have been reported in the literature (Table 1). These results
in­di­ca­te high karyotypic variation within the genus, and its potential to become an interesting
model for the study of chromosomal speciation accompanying either adaptive radiation (as
exemplified by the Spalax-model, cf. Nevo et al. 2001) or by vicariance and genetic drift (as
45
Table 1. Known karyotypes (diploid numbers of chromosomes) of African dormice (Graphiurus sp.)
Tab. 1. Popsané karyotypy (diploidní počty chromosomů) afrických plchů rodu Graphiurus
species
2n
country
authors
G. hueti de Rochebrune, 1883
G. mu­ri­nus (Desmarest, 1822)
G. par­vus (True, 1893)
G. microtis (Noack, 1887)
G. murinus (Desmarest, 1822)
40
70
70
52
46
Ivory Coast
Ivory Coast
Niger
Malawi
South Africa
Trainer & Dosso 1979
Trainer & Dosso 1979
Dobigny et al. 2002
Chitaukali et al. 2001, this paper
Kryštufek et al. 2004
exemplified by the African mole-rats of the genus Fukomys Kock, Ingram, Frabotta, Burda et
Honeycutt, 2006; cf. van Daele et al. 2004).
Within the context of a more general report on results of a faunistic survey of small mam­mals
in the National Park Nyika in Northern Malawi, 1997, we named the diploid number of chro­
mo­so­mes we found in two dormice captured in the region, yet we did not provide any further
in­for­ma­ti­on on this aspect (Chitaukali et al. 2001). Regarding the fact that African dormice
in general are poorly known karyologically, in contrast to most of the European species, the
karyotypes of which have been studied in detail in nany localities (Zima et al. 1994) as well as
regarding the potential importance of this kind of information (se above), it is worthy to publish
additional data on this finding.
Material and Methods
Two dormice were captured (in Sherman live traps) in Nganda (S10.26, E 33.51, grid 1033-B4, altitude
2,200–2,300 m a. s. l.), National Park Nyika, Northern Malawi, on alpine meadows in the vicinity of shrubs
and rocks in April 1997. One further animals was trapped in the Chipome Valley (S10.20, E33.50, grid
1033-B4, altitude about 1,530 m a. s. l.), surveyed in July 1998. Dominant vegetation type at Chi­po­me is
Bra­chys­te­gia woodland. In 1999, W. N. Chitaukali captured also two dormice in Mzuzu (in the garden
of Mr. R. J. Murphy) (1,350 m a. s. l.). Animals were sacrificed, examined using standard mam­ma­lo­gi­cal
procedures. Voucher specimens (preserved in ethanol) have been deposited in the Senckenberg Museum
Table 2. Measurements of small-eared dormice (Graphiurus microtis) collected in Nyika, Malawi. LC
= head and body length, LCd = tail length, LTP = length of the hind foot; under breeding state either the
number and distribution of embryos in uterine horns (L = left, R = right) or size of testes are given
Tab. 2. Rozměry plchů malouchých (Graphiurus microtis) chycených na náhorní plošině Nyika, Ma­lawi.
sex = pohlaví, mass = hmotnost, LC = délka těla, LCd = délka ocasu, LTP = délka zadní tlapky, pinna
= délka ucha; breeding state = buď počet a umístění embryí v děložních rozích (L = levý / R = pravý)
anebo rozměry varlat
ID locality
date sex
mass (g)
LC
(mm)
LCd
(mm)
LTP (mm)
69/97Ngan­da
72/97Nganda
55/98Chipome
61/99Mzuzu
62/99Mzuzu
24.0
16.0
14.0
38.0
46.0
88.6
57.4
61.4
64.5
72.6
93.2
101.0
100.0
15.6
15.0
16.8
12.0
18.0
46
10.04. 1997
10.04. 1997
22.07. 1998
09.08. 1999
09.08. 1999
F
M
M
F
F
pinna breeding state
(mm)
12.0
14.0
13.2
11.5
15.0
L2 / R1
5.6×2.6
in Frankfurt am Main, Germany. Both individuals from Nganda were karyotyped using the standard
preparation of bone marrow chromosomes from long bones (Lee 1969, Lee & Elder 1977). The slides
were stained by Giemsa.
Results and Discussion
Basic biometric data are given in Table 2. Dormice from Nyika were preliminarily identified (on
the base of morphological features and biogeography) as Graphiurus microtis (Noack, 1887)
(determination by Dr. D. Kock, Frankfurt am Main). The specimens from Mzuzu are distinctly
larger (Table 2) and may belong to a different species.
Finding of a pregnant female (with two embryos) in April in Nganda suggests that the bre­e­
ding season involves at least the end of the rainy season.
The karyotype of two specimens from Nganda consisted of 2n=52 chromosomes (Fig. 1): 11
pairs of metacentrics, 14 pairs of acrocentrics and subtelocentrics, the X-chromosome being
a metacentric, Y-chromosome a small acrocentric.
Fig. 1. Karyotype of a male Graphiurus microtis from Nganga, Nyika Plateau, Malawi.
Obr. 1. Karyotyp samce plcha malouchého (Graphiurus microtis) z lokality Nganga, náhorní plošina
Nyika, Malawi.
47
The karyotypes of dormice are characterized by the prevalence of biarmed autosomes (Zima
et al. 1994). This is also the case in G. murinus studied by Kryštufek et al. (2004). In this respect, karyotypes of G. microtis in our study are somewhat unusual being composed of more
acro­cent­ric and telocentric chromosomes rather than by clearly biarmed chro­mo­so­mes. Sex
chro­mo­so­mes (larger metacentric X, and dot like Y) correspond to the pattern known also from
other dormouse taxa.
In absence of comparative data on karyotypes of other populations of G. microtis, no conclu­
si­on about the taxonomic status of the Nyika population can be made at the present. Regarding
the fact that so far, only five populations (representing at least four different species) of Gra­
phiu­rus were karyotyped, and in absence of any chromosome banding studies, it would be
very speculative to discuss differences between karyotypes from taxonomical or evo­lu­ti­o­na­ry
point of views.
Souhrn
Karyotyp afrického plcha malouchého (Graphiurus microtis) z lokality Nganda (Nyika Plateau, Malawi)
(obr. 2) sestává z 2n=52 chromozomů: 11 párů metacentrických, 14 párů akrocentrických a subtelocentrických, X-chromozom je metacentrický, Y-chromozom malý akrocentrický. Vzhledem k tomu, že v rámci
Fig. 2. Small eared dormouse (Graphiurus microtis) from the Nyika Plateau, Malawi.
Obr. 2. Plch malouchý (Graphiurus microtis) z náhorní plošiny Nyika, Malawi.
48
pod­če­le­di Graphiurinae byl karyotyp dosud popsán (včetně této práce) jen u čtyř druhů z pěti oblastí, přičemž
je uzná­vá­no 13 až 14 druhů rozšířených v celé subsaharské Africe, představuje každá nová studie cenný
příspěvek k poznání ne zcela jasné systematiky a evoluce celé čeledi. Diskuse chromozomové speciace
afrických plchů na základě dostupných omezených dat by ale byla předčasná a spekulativní.
Acknowledgement
The work could not have been done without organisation and logistic support provided by Peter C. Over­
ton, organiser of the expeditions surveying biodiversity of the National Park Nyika. Assistance in the
field by Jana Burda, Marianne J. Overton, Katie Ried, and scouts of the Nyika National Park is highly
appreciated. Dieter Kock, Forschungsinstitut und Museum Senckenberg, Frankfurt am Main, Germany,
is to be thanked for checking determination of dormice. Last but not least, we thank to the Malawi De­
part­ment of National Parks and Wildlife for the permission to work in the Nyika National Park and for
all the assistance provided.
References
Chitaukali W. N., Burda H. & Kock D., 2001: On small mammals of the Nyika Plateau, Malawi. Pp.:
415–425. In: Denys C., Granjon L. & Poulet A. (eds.): African Small Mammals. IRD Editions, Col­le­
cti­on Colloques et seminaires, Paris, 570 pp.
Dobigny G., Nomao A. & Gautun J.-C., 2002: A cytotaxonomic survey of rodents from Niger: im­pli­ca­ti­
ons for systematics, biodiversity and biography. Mammalia, 66: 495–523.
Genest-Villard H., 1978: Revision systematique du genre Graphiurus (Rongeurs, Gliridae). Mammalia,
42: 391–426.
Holden M. E., 1993: Family Myoxidae. Pp.: 763–770. In: Wilson D. W. & Reeder D. M. (eds.): Mammal
Species of the World. A Taxonomic and Geographic Reference. Second Edition. Smithsonian In­sti­tu­ti­on
Press, Washington, 1206 pp.
Holden M. E., 1996: Systematic revision of sub-Saharan African dormice (Rodentia: Myoxidae: Gra­
phiu­rus). I. An introduction to the generic revision, and a revision of Graphiurus surdus. Am. Mus.
Novit., 3157: 1–44.
Kock D., Ingram C. M., Frabotta L. J., Burda H. & Honeycutt R. L., 2006: On the nomenclature of
Bathyergidae and Fukomys n. g. (Mammalia: Rodentia). Zootaxa, 1142: 51–55.
Kryštufek B., Haber W., Baxter R. M. & Zima J., 2004: Morphology and karyology of two populations
of the woodland dormouse Graphiurus murinus in the Eastern Cape, South Africa. Folia Zool., 53:
339–350.
Lee M. R., 1969: A widely applicable technique for direct processing of bone marrow for chromosomes
of vertebrates. Stain Technol., 44: 155–158.
Lee M. R. & Elder F. F. B., 1977: Karyotypes of eight species of Mexican rodents (Muridae). J. Mammal.,
58: 479–487.
Montgelard C., Matthee C. A. & Robinson T. J., 2003: Molecular systematics of dormice (Rodentia:
Gli­ri­dae) and the radiation of Graphiurus in Africa. Proc. R. Soc. Lond. B, 270: 1947–1955.
Nevo E., Ivanitskaya E. & Beiles A., 2001: Adaptive Radiation of Blind Subterranean Mole Rats: Na­ming
and Revisiting the Four Sibling Species of the Spalax ehrenbergi Superspecies in Israel: Spalax galili
(2n=52), S. golani (2n=54), S. carmeli (2n=58), S. judaei (2n=60). Backhuys Pu­b­lis­hers b.v., Leiden,
The Netherlands, 204 pp.
Rossolimo O. L., Potapova E. G., Pavlinov I. Ya., Kruskop S. V. & Voltzit O. V., 2001: Soni (Myoxi­
dae) mirovoj fauny [Dormice (Myoxidae) of the World]. Moscow Univ. Publ., Moscow, 233 pp (in
Russian).
Trainer M. & Dosso H., 1979: Recherches caryotypiques sur les rongeurs de Côte d’Ivoire: résultats
préliminaire pour les milieux fermés. Mammalia, 43: 254–256.
49
Van Daele P. A. A. G., Dammann P., Kawalika M., Meier J.-L., Van De Woestijne C. & Burda H., 2004:
Chromosomal diversity in Cryptomys mole-rats (Rodentia: Bathyergidae) in Zambia; with the descrip­
ti­on of new karyotypes. J. Zool. Lond., 264: 317–326.
Zima J., Macholan M. & Filippucci M. G., 1994: Chromosomal variation and systematics of myoxids.
Hys­trix, n. s., 6(1–2): 63–76.
50
Lynx (Praha), n. s., 37: 51–66 (2006).
ISSN 0024–7774
Notes on the genus Ochotona in the Middle East
(Lagomorpha: Ocho­to­ni­dae)
Poznámky k rodu Ochotona na Blízkém východě (Lagomorpha: Ochotonidae)
Stanislav Čermák1, Ján Obuch2 & Petr Benda3,4
Department of Geology and Paleontology, Charles University, Albertov 6, CZ–128 43 Praha 2,
Czech Republic; [email protected]
2
Botanical Garden, Comenius University, SK–038 15 Blatnica, Slovakia; [email protected]
3
Department of Zoology, National Museum (Natural History), Václavské nám. 68, CZ–115 79 Praha 1,
Czech Republic; [email protected]
4
Department of Zoology, Charles University, Viničná 7, CZ–128 44 Praha 2, Czech Republic
1
received on 14 September 2006
Abstract. New records of Ochotona from Iran and Turkey are reported. The finding of a bone remain
of the Recent age (from the Eagle owl pellet) from Turkey represents the first record of the genus and
family in this country and exdends the range of the genus in the Middle East by approximately 600 km
to the northwest. All new records from Iran belong to O. rufescens (Gray, 1842), the only known species
in the region. Based on the comparison of the Persian material, the pikas from the Zagros and Kopetdag
Mts are similar in size and probably do not represent two different subspecies. The newly reported pikas
of the Recent age from the Armenian Highlands of Turkey and Iran slightly differ from those inhabiting
the rest of Iran and are reported as O. cf. rufescens.
INTRODUCTION
Only one species of ochotonid lagomorph, the rufescent or collared or Afghan pika, Ocho­to­na
rufescens (Gray, 1842), is known to occur in the Middle East (i.e. the region comprising Arabia, Iran, and the Asian part of Turkey). Its distribution range covers the mountainous areas of
Afghanistan, north-western Pakistan, Iran and south-western Turkmenistan (Ellerman & Mo­r­
ri­son-Scott 1951, Gromov & Erbaeva 1995). Concerning the Middle East proper, the Afghan
pika has been reported to reach the mountains of Iran.
Taghizadeh (1964) and Lay (1967) summarised the distribution of the Afghan pika in Iran.
Lay (1967: 151) using the data by Blanford (1876), Murray (1884), and Misonne (1956) and
his new records, suggested that: “the rufescent pika inhabits all of the mountainous re­gi­ons of
Iran”. However, the westernmost Persian record of pika came from the vicinity of Agbolagh
Morched, Kurdistan (Lay 1967), in the central Zagros Mts, and represented also the westernmost
Recent record of the species. No records have been reported from the wes­tern Elborz Mts, the
Talysh Mts or from Armenian Highlands, i.e. from all the mountains of the northwestern part
of Iran (see Fig. 1 and/or Taghizadeh 1964: 25, Fig. 12). The last new records of the Afghan
pika from Iran were referred by de Roguin (1988) and Obuch & Kriš­tín (2004), including the
easternmost Persian record of the species, from Mount Taftan, Baluchestan.
51
Although the talus/steppe-dwelling pikas (sensu Smith et al. 1990) are known to be re­gu­
lar­ly distributed in the mountains from Iran and Turkmenistan to the east, several records of
Ocho­to­na have been mentioned from Transcaucasia. However, at least some of them actually
are the records of fossil or sub-fossil bone fragments found in cave deposits, and their spe­ci­es
af­fi­li­a­ti­on is often not fully convincing. Sosnihina (1947, 1948) reported findings of Ocho­to­na
from Armenia; a lower jaw of sub-fossil age from a gorge of the Čirahan River and other bones
from the vicinity of the Aknalič Lake. She identified these bone remains to come from pellets of
the Eagle owl (Bubo bubo) and specifically attributed both records to O. rufescens according to
the geographically nearest known species. Another Armenian records of pikas of sub-fossil age
were reported by Dal’ (1954, 1957) from caves of the Urcskij Range. However, Dal’ (1957)
Fig. 1. Records of Ochotona in the Middle East and Transcaucasia; numbers correspond with the sites
numbered in the list of new records (see the text); squares – fossil or sub-fossil records (of the Upper Biharian-Toringian age), circles – Recent records, closed symbols – new records, open symbols – literature
data (based on Sosnihina 1947, 1948, Coon 1951, Dal’ 1957, Gadžiev & Aliev 1966, Lay 1967, de Ro­
guin 1988, Montuire et. al. 1994, and Obuch & Krištín 2004). For the extralimital data on occurrence of
Ocho­to­na rufescens in Turkmenistan see Sokolov et al. (1994: 59, Fig. 18), in Afghanistan see Hassinger
(1973: 34, Fig. 6), and in Pakistan see Roberts (1997: 310, map 80).
Obr. 1. Místa nálezů pišťuchy (Ochotona) na Blízkém východě a v Zakavkazí; čísla odpovídají číslování
lokalit v textu (kapitola New Records); čtverce – fossilní, nebo subfossilní nálezy (stáří svrchní bihar až
toring), kroužky – recentní nálezy, plné symboly – nové nálezy, prázdné symboly – literární údaje (podle
Sosnihiny 1947, 1948, Coona 1951, Dal’a 1957, Gadžieva & Alieva 1966, Laye 1967, de Roguina 1988,
Montuireové et. al. 1994 a Obucha & Krištína 2004). Extralimitní rozšíření pišťuchy rezavé (Ochotona
ru­fes­cens) v Turkmenii viz Sokolov et al. (1994: 59, Fig. 18), v Afghanistanu viz Hassinger (1973: 34,
Fig. 6) a v Pakistanu viz Roberts (1997: 310, map 80).
52
placed his own and Sosnihina’s records, vernacularly named as an Armenian pika (‘ar­me­nij­ska­
ja piščuha’), into morphological proximity of Ochotona eximia (Khomenko, 1914), descri­bed
from the Upper Mi­o­ce­ne site of Taraklia in Moldavia. Vereščagin (1959) summarised these
Transcaucasian records of Armenian pikas as of the Holocene age and considered the ‘species’
to be extinct in the Recent.
Vekua & Šidlovskij (1958) reported on a record of a large sized ochotonid from the Mous­
te­ri­an, Middle Palaeolithic, of Georgia (Copi Station, Marneuli Dist., E Georgia). Later, based
on an additional material from this site, Vekua (1967) described the respective pika as a new
species, Ochotonoides transcaucasica (now included in the genus Ochotona, see below).
Another pika species from Transcaucasia, Ochotona azerica, is reported from the Acheulean/
Mousterian deposits of the Azyh Cave, Azerbaijan (Aliev 1969, Gadžiev et al. 1979). A se­cond
Azerbaijan record of Ochotona was published by Gadžiev & Aliev (1966) from Taglar, also
of the Mous­te­ri­an age.
All the above mentioned published records of Ochotona from Transcaucasia are reported
to be of fossil to sub-fossil age (the Upper Biharian-Toringian age sensu Fejfar & Heinrich
1983; the Akhalkalaki-Akhstyr faunal units sensu Baryshnikov 2002). However, some Tran­s­
cau­ca­si­an records were mentioned to come from the Eagle owl pellets but without spe­ci­fying
their age, i.e. whether they come from fresh pellets or from deposits of bones from supposed
owl pelleting places.
Vinogradov & Gromov (1952) suggested possible occurrence of O. rufescens in Ar­me­nia,
based on findings of pika bones in Eagle owl pellets in the Daralegezskij Range, Mikojan [=
Ehednadzor] District. Šidlovskij (1976: 72) mentioned: “remains of lower jaws of pikas have
been found in the last years in southern Armenia, in the Eagle owl pellets in a cave near the
village of Amagu in the Sarajbulagskij Range and in Georgia, on the eastern slope of the Džavahetskij Range (Dmanisi Dist.)” [translated from Russian].
Probably based on these reports, Hoffmann (1993) and Hoffmann & Smith (2005) men­ti­o­ned
Armenia among the countries inhabited by O. rufescens. On the other hand, Russian authors
(i.e., Ognev 1940, Gureev 1963, 1964, Erbaeva 1988, Sokolov et al. 1994, Gromov & Erbaeva
1995, Pavlinov et al. 1995, Pavlinov & Rossolimo 1998) did not mention any Recent representative of the genus Ochotona from Transcaucasia. Moreover, Gureev (1981: 73) wrote in the
paragraph concerning O. rufescens: “obviously, this species or a species close to it existed in the
Caucasus at the end of the Pleistocene” [translated from Russian]. Thus, the occurrence of pikas
in Transcaucasia and adjacent regions of the Middle East in the Recent remains indefinite.
Here, we report new findings of Ochotona from several parts of the Middle East in attempt
(1) to draw the Recent distribution range of the genus in the region of southwestern Asia, and
(2) to evaluate morphologic characteristics and systematic status of particular populations of
the Middle East.
MATERIAL AND METHODS
The most of ochotonid material described in this paper comes from owl pellets (see the list of new records
below). The bone remains from owl pellets are fragmented and mostly belong to juvenile or sub-adult
in­di­vi­du­als. Only non-juvenile specimens (according to Lissovsky 2004) were included in the metric
comparison. The following abbreviations are used in the text: P – premolar; M – molar; L – length; W
– width; n – sample size; X – arithmetic mean; OR – observed range of the sample; MR – mandible ratio
[mandible height at P3 ×100 / alveolar length of P3–M3]; PR – posteroconid ratio [posteroconid length × 100
/ total tooth length]. Descriptive dental terms and metrics are used according to López-Martínez (1974),
53
Erbaeva (1988), Sen (1998), and Čermák (2004). Dimensions and drawings were made using binocular
microscope. Me­a­su­re­ments are given in millimetres and ratios in percentages.
For comparative purposes, the morphometric characteristics of O. pusilla (Pallas, 1768), O. pallasi (Gray,
1867), O. rutila Severtzov, 1873, O. macrotis (Günther, 1875), and O. roylei (Ogilby, 1839) provided
by Ognev (1940), Gureev (1964), and Erbajeva (1988) were taken into account. The studied material is
de­po­si­ted partly in the collection of the second author (JO), partly in the collection of the Department of
Zoology, National Museum, Prague (NMP).
NEW Records
Turkey
1. 3 km SE of Güzyurdu, small cave in a mountain pass, sub-fossil deposite 2A* (Gümüşhane Prov;
39° 54’ N, 39° 34’ E, ca. 2300 m a. s. l.), 15 September 1995, isolated right M1 and a fragment of left
maxilla of Ochotona cf. pusilla collected, leg. I. Horáček.
2. Ishak Paşa Sarayi, 6 km E of Doğu Bayazit, rock outcrop (Ağri Prov.; 39° 31’ N, 44° 08’ E, ca. 1950 m
a. s. l.), 1 October 2002, right mandible of Ochotona cf. rufescens (NMP 90157) from a pellet of Bubo
bubo collected, leg. J. Obuch.
Iran
3. Bastam, 10 km W of Qarah Ziya’oddin, archaeological site (Azerbaijan Gharbi Prov., 38° 53’ N,
44° 58’ E, ca. 1250 m a. s. l.), 30 September 1998, right mandible of Ochotona cf. rufescens (NMP
90156) from a pellet of Bubo bubo collected, leg. J. Obuch.
4. rocks above Gazanak (Mazandaran Prov., 35° 55’ N, 52° 15’ E, ca. 1650 m a. s. l.), 15 May 1997, re­ma­
ins of three inds. of Ochotona rufescens from pellets of Bubo bubo (one left and three right mandibles,
two maxillar fragments) collected, leg. J. Obuch.
5. 3 km E of Tangeh, 10 km SW of Raz (Khorasan Shamali Prov., 37° 54’ N, 56° 56’ E, ca. 1200 m
a. s. l.), 12 May 1997, several colonies of Ochotona rufescens observed, five individuals caught, an
adult male (NMP 90153 [S+A]), sub-adult male and a female (NMP 90154, 90155 [A]) collected, leg.
P. Benda, J. Sádlová & R. Šumbera.
6. Qarloq, 15 km W of Bojnurd (Khorasan Shamali Prov., 37° 30’ N, 57° 26’ E, ca. 890 m a. s. l.), 12 May
1997, remains of five inds. of Ochotona rufescens from pellets of Bubo bubo (four left and five right
mandibles, fragments of five skulls) collected, leg. J. Obuch.
7. two rocky valleys ca. 2 and 5 km E of Emam Qoli, ca. 30 km N of Quchan (Khorasan Razni Prov.;
37° 22’ N, 58° 32’ E, ca. 1710 m a. s. l.), 11 May 1997, remains of 30 inds. of Ochotona rufescens
from pellets of Bubo bubo (30 right and 30 left mandibles, remains of 16 skulls), leg. J. Obuch; several
colonies of O. rufescens observed, one individual caught, leg. J. Sádlo; 25 May 2006, colonies of O.
rufescens observed, leg. A. Reiter (cf. Fig. 2).
* Güzyurdu 2A: surface layer of a loamy debris infilling of a narrow karstic fissure in a small rock overhang
ca. 3 km SE of the Güzyurdu village which provided the following assemblage (MNI): 1 ind. of Bufo cf.
viridis, 1 Rana cf. temporaria, 1 Lacerta sp., 1 Aves indet., 1 Erinaceus sp., 1 Spermophilus cf. xan­tho­
prym­nus, 1 Apodemus cf. uralensis, 3 Mesocricetus cf. brandtii, 2 Cricetulus migratorius, 1 Allactaga
cf. williamsi, 1 Arvicola cf. terrestris, 4 Chionomys nivalis, 7 Microtus cf. obscurus, 2 Microtus cf. sub­
ter­ra­ne­us, 3 Spalax sp., 1 Ochotona cf. pusilla, 1 Lepus sp. The sample obtained from the lower layer
of the infilling was of a similar composition (except for Ochotona, Spalax, and Erinaceus). The bones
exhibit signs of fossilisation and undoubtedly are not of the Recent age. Tentatively, the assemblage is
con­si­de­red to be of the Holocene age (I. Horáček in litt.). The accompanying species do occur in a wider
sur­roun­dings of the site even recently (cf. e.g. Kryštufek & Vohralík 2001).
54
8. dry valley ca. 5 km S of Mina, ca. 25 km SW of Dargaz (Khorasan Razni Prov.; 37° 18’ N, 58° 58’ E,
ca. 1075 m a. s. l.), 22–23 May 2006, a colony of Ochotona rufescens observed, leg. A. Reiter.
9. rocks 12 km E of Bazangan, 18 km NNW of Mazdavand (Khorasan Razni Prov., 36° 17’ N, 60° 33’ E,
ca. 650 m a. s. l.), 11 May 1997, remains of 16 inds. of Ochotona rufescens in pellets of Bubo bubo
(14 right and 16 left mandibles, fragments of 9 skulls) collected, 8 October 2002, remains of 11 inds.
of O. rufescens from pellets of Bubo bubo (7 right and 11 left mandibles) collected, leg. J. Obuch.
10. wadi under Emam Sadeh, ca. 12 km W of Kashan (Esfahan Prov., 33° 59’ N, 51° 17’ E, ca. 1130 m
a. s. l.), 6 April 2000, a fragment of a left mandible of Ochotona rufescens from a pellet of Bubo bubo
collected, leg. J. Obuch.
11. Deh Zireh, 25 km NNW of Natanz (Esfahan Prov., 33° 46’ N, 51° 42’ E, ca. 850 m a. s. l.), 27 April
1996, remains of five inds. of Ochotona rufescens from pellets of Bubo bubo (three left and four right
mandibles, remains of one skull) collected, leg. J. Obuch.
12. Qamishlu, ca. 40 km W of Shahreza (Es­fa­han Prov., 32° 02’ N, 51° 29’ E, ca. 2200 m a. s. l.), 28 April
1996, remains of one skull and two mandibles (right and left) of Ochotona rufescens from a pellet of
Bubo bubo collected, leg. J. Obuch.
13. 5 km NE of Deh Bakri, ca. 40 km W of Bam (Kerman Prov., 29° 05’ N, 57° 56’ E, ca. 1930 m a. s. l.),
7–8 April 2000, remains of two inds. of Ochotona rufescens from pellets of Bubo bubo (two left mandibles and one right mandible, fragments of one skull) collected, leg. J. Obuch.
Fig. 2. A valley at Emam Qoli, ca. 30 km N of Quchan, Kopetdag Mts, northeastern Iran. Rocky slopes
represent a typical habitat abundantly dwelled by Ochotona rufescens (photo by A. Reiter).
Obr. 2. Horské údolí nedaleko Emam Koli, asi 30 km na sever od města Kučan, pohoří Kopetdag, severovýchodní Írán. Skalnaté svahy údolí vytvářejí biotop hojně obývaný pišťuchou rezavou (Ochotona
rufescens) (foto A. Reiter).
55
NOTES ON BIOGEOGRAPHY
New records of Ochotona of the Recent age from the Middle East come from five sub-regions
(see Fig. 1); (1) northeastern Iran, the range of the Kopetdag Mts along the border with Turk­
me­nistan [records Nos. 5–9]; (2) southeastern Iran, the easternmost extent of the Zagros Mts
in the Kerman Province [13]; (3) central Iran, central parts of the Zagros Mts and northern part
of the Qohrud Range [10–12]; (4) central Iran, central part of the Elborz Mts [4]; (5) the border
region of Iran and Turkey, the eastern part of the Armenian Highlands [2, 3].
Most of the new records come from Iran, where they cover mainly the known distribution
range as described by Lay (1967). They come from all main mountainous areas of the Iran
Plateau, including the central Persian mountains (Zagros Mts, Elborz Mts, Qohrud Mts) as well
as the southeasternmost parts of the Zagros system, the Jebal Barez Mts, bordering the Baluchestani deserts. Records from the latter area were given already by Lay (1967) and de Roguin
(1988). Only one record has been published before from the Persian side of the Kopedag Mts
(Akhlamad, Khorasan Razni Prov.; Missone 1956). Newly reported data sug­gest the occur­ren­ce
of O. rufescens in the whole main range of these mountains, similarly as it is known from the
Turkmen part of the region (Sokolov et al. 1994).
The new records of pikas from the border area between the northwestern corner of Iran and
eastern Turkey [2, 3] are of particular significance because the extant representatives of the
family had not been reported from this region before (see e.g. Lay 1967, Kumerloeve 1975).
In Turkey, the finding at Ishak Paşa Sarayi represents the first record of the genus and family
in this country (Kumerloeve 1975, Kryštufek & Vohralík 2001). The pika jaw was obtained
from a pellet of the Eagle owl, at the site where material from pellets had been collected at least
three times before, during visits in April 1996, April 1997, and September 1998, and no remnants
of Ochotona were found. The Recent age of this finding is therefore undoubted and possible
confusion with fossil or sub-fossil material is unlikely. Concerning the Iran side of the region,
one pika record comes from Bastam, an archaeological site used as a rest place by the Eagle owl.
These records shift the known Recent distribution range of the genus Ocho­to­na some 600 km
to the northwest and verify the Recent distribution of this genus in the Cau­ca­sus region (see the
Introduction chapter). Moreover, they suggest that the records from Transcaucasian countries
formerly considered as of rather uncertain age may also be of the Recent age (see Fig. 1).
The other Turkish finding of Ochotona [record No. 1] comes from a sub-fossil deposition and
its interpretation as an extant occurrence of pika in central Anatolia is irrelevant.
Notes on morphology
The ochotonid material collected in various parts of the Middle East is quite homogenous in
morphology and dimensions. It is represented by a large sized pika (alveolar length P3–M3:
X=9.33, n=27; alveolar length P2–M2: X=8.97, n=14) with a relatively long and low mandible
(MR: X=59, n=27). The lower incisors extend to below the area between P4 and M1. The posterior mental foramen is located ventrally to M3, or even more posteriorly. The skull is narrow.
Frontal bones with well-developed crests form the a narrow interorbital region. The relatively
short rostrum possesses a large preorbital foramen of the triangular type. Nasal bones are wider
anteriorly than posteriorly. The incisive-palatal foramen is not closed.
Third lower premolars (P3; 9 specimens), the most characteristic teeth, are available almost
only in non-adult stages; based on the morphology, they belong to not overwintered animals
younger than 4 months (sensu Lissovsky 2004). These permanent teeth are worn, ne­ver­the­less
56
they still retain their conical structure. So, occlusal dimensions and morphology of the tooth
(the main diagnostic features) are not stable during conversion of the crown to its de­fi­ni­ti­ve
prismatic state, therefore evaluation of demonstrative conclusions is difficult. Dif­f­e­ren­ces
between occlusal and root morphologies are shown on Fig. 3 (1–4, 1a–4a). Only in one tooth,
a well developed lingual fold divides the posteroconid and forms a three-seg­men­ted juvenile
appearance of the tooth (Fig. 3: 5).
Owing to the hypselodont teeth, the alveolus of P3 corresponds in its shape with occlusal
outline of P3, therefore the ratios of visible structures of both alveolus and P3 are the same and
their comparison possible (see Čermák 2004). According to morphology of the available P3
Fig. 3. Morphology of P3 (non-adult) and P3 teeth in Ochotona rufescens from Iran; all teeth are figured
as left specimens (1–2, 5–6, 9–10 are reversed); 1–10 – occlusal views, 1a–4a – root views; scale bar
– 1 mm.
Obr. 3. Morfologie třenových zubů (P3 – nedospělí jedinci; P3) u pišťuchy rezavé z Íránu; všechny zuby
jsou zobrazeny jako levé (1–2, 5–6, 9–10 jsou stranově převráceny); 1–10 – okluzální pohled, 1a–4a
– kořenový pohled; měřítko – 1 mm.
Legend / legenda: P3: 1–3, 1a–3a, 5 – Emam Qoli, 4, 4a – Bazangan; P3: 6–8 – Emam Qoli, 9 – Qarloq,
10 – Bazangan.
57
Fig. 4. Morphology of P3 alveoli in Ochotona from the Middle East (Iran, Turkey); all alveoli are figured
as left specimens (1–4, 8, 9, 21–23, 25 are reversed); scale bar – 1 mm.
Obr. 4. Morfologie alveolů třetího třenového zubu (P3) u pišťuchy rezavé z Blízkého východu (Írán,
Turecko); všechny alveoly jsou zobrazeny jako levé (1–4, 8, 9, 21–23, 25 jsou stranově převráceny);
měřítko – 1 mm.
Legend / legenda: 1 – Ishak Paşa Sarayi, 2 – Bastam, 3, 4 – Qarloq, 5 – Gazanak, 6–12 – Emam Qoli,
13–19 – Bazangan, 20–22 – Deh Zireh, 23 – Qamishlu, 24–25 – Deh Bakri.
58
alveoli (L×W = 1.80×2.00, OR = 1.55–2.00 × 1.70–2.30; n=33), the tooth is almost tri­an­gu­lar
(Fig. 4). The posteroconid is large and long (the relative length is determinable from entoconid
pro­tru­si­on on the lingual side of P3 alveolus – see Fig. 4). The anteroconid is small (PR of P3
alveoli: X=74, n=33), rounded, often with asymmetrical position. The P2 is (ac­cor­ding to the
shape of alveoli, n=6) oval, in its shape, its lingual side is almost straight. The P3 (Fig. 3: 6–10)
is tra­pe­zoi­dal in the occlusal outline (L×W = 1.27×2.61, OR = 1.08–1.44 × 2.04–3.00; n=7).
The mesial hypercon of P3 is relatively large (mean of hyperconal width is 1.51); its width is
quite variable. There is also often an additional protrusion on the anterior part of the P3 metacon
(Fig. 3: 6–8).
All the above mentioned features of the compared material are characteristic for Ochotona
rufescens and fall clearly within the known variation range of the species (see e.g., Ognev 1940,
Erbaeva 1988). At the same time, it differs from the other species inhabiting the sur­roun­ding
regions. The studied ochotonids from the Middle East, as well as all specimens of O. rufescens,
are well distinguishable from O. pusilla by their much larger size, and from all of the other known
Fig. 5. Skull and mandible of Ochotona rufescens from the Kopetdag Mts., NE Iran (NMP 90153) (above) and mandible of O. cf. rufescens from Ishak Paşa Sarayi, E Turkey (NMP 90157) (below); scale bar
– 5 mm.
Obr. 5. Lebka a spodní čelist pišťuchy rezavé (Ochotona rufescens) z pohoří Kopetdag, severovýchodní
Írán (NMP 90153) (nahoře) a spodní čelist pišťuchy O. cf. rufescens z letoviska Ishak Paşa Sarayi, vý­
chod­ní Turecko (NMP 90157) (dole); měřítko – 5 mm.
59
species by their much smaller P3 anteroconid; the mean PR of P3 alveoli of geographically close
species (i.e., O. pallasi, O. rutila, O. macrotis, and O. roylei) falls wi­thin the range from 62 to
65. These species differ also from the pikas from Iran in many skull features: e.g., in their closed incisive-palatal foramen (O. pallasi and O. rutila), in their dis­tinct­ly wider interorbital part
(O. rutila, O. macrotis, and O. roylei), or in their presence of foramina in the frontal bones (O.
macrotis and O. roylei), etc. (see Ognev 1940, Gureev 1964 and Erbaeva 1988 for details).
Based on the above discussed dimensions and morphological characters, the pikas from Ishak Paşa Sarayi (Turkey; record No. 2) and Bastam (NW Iran; No. 3) correspond very clo­se­ly
with O. rufescens and fall clearly into its variation range. Nevertheless, they differ in cer­tain
respect. These specimens are somewhat larger than other Persian pikas under study (Figs. 5, 6).
However, taking into account the small size of the sample, it is difficult to evaluate the actual
meaning of the difference. It seems possible either that (1) Ochotona from the border region of
Iran and Turkey falls within the variation range of populations from northeastern and central
Iran, while accidentally only the larger individuals were available for comparison, or (2) these
western pikas are actually larger than Ochotona from northeastern and central Iran. However,
the second possibility does not have to imply the existence of a separate taxon, as there could
also be a cline increase in body size in pikas from east to west (the phenomenon known in such
geographical arrangement in other mammals, e.g. bats, see Benda & Horáček 1995). In P3 an­
te­ro­co­nid proportions, the pikas from Ishak Paşa Sarayi and Bastam lie at the upper margin of
variation at the Persian pika samples; PR of P3 alveoli are 83 and 81, re­spe­cti­ve­ly. However,
as there are no mandibles or additional dentition available (particularly P3), it is quite difficult
to make a more conclusive comparison and taxonomical identification of the pikas from the
region at the border between Iran and Turkey. Thus, we assign these specimens tentatively to
O. cf. rufescens.
Using the available material from Iran, the expected differences in body size between po­pu­
la­ti­ons from northeastern Iran (the Kopetdag Mts, an area of occurrence of O. rufescens regina
Thomas, 1911) and central Iran (central parts of the Zagros Mts, an area of occur­ren­ce of O. r.
vizier Thomas, 1911) were not proved (Fig. 6).
In any case, based on both morphological and metric characteristics, all the other Persian
samples surveyed in this paper can be referred to O. rufescens, with an exception of the ma­te­rial
from Emam Sadeh (record No. 10), where the lack of available features does not allow a precise
species assignment. However, taking into account the geographical position of the locality, it
seems very probable that the mandibular fragment belongs also to that species.
NOTES ON PALEONTOLOGY
The fossil record of O. rufescens, including the fossils placed in close proximity of this species,
is quite poor to date (Erbaeva 1988). The ochotonid remains, almost identical with the extant O.
rufescens, known from the Acheulean Palaeolithic site of Sel’ Ungur, Kirghizia, slightly differ
from the Recent species by their longer diastema and wider P3 (Erbaeva 1988). Besides the
ochotonid from Kirghizia, some large sized pikas referred to O. rufescens have been reported
from Transcaucasia (see above). Unfortunately, no detailed characteristics of these specimens
are available in the literature, and a relevant comparison is difficult. Dal’ (1957) placed the
Armenian findings into proximity of Ochotona eximia (Khomenko, 1914) based on the size
and morphological similarities in P3 and mandible. In his opinion, these ochotonids represent
an Upper Tertiary relic derived from a heterogeneous group of species from the “eximia-gigas”
60
Fig. 6. Scatter plot of the alveolar length of the lower tooth-row (P3–M3) against the buccal height of
the mandible at P3 in the samples of Ochotona coming from the Middle East (Iran, Turkey); only adult
spe­ci­mens were used.
Obr. 6. Srovnání alveolarní délky dolní zubní řady (P3–M3) proti bukální výšce spodní čelisti u třetího
tře­no­vé­ho zubu (P3) jedinců pišťuch z Blízkého východu (Írán, Turecko); použity byly rozměry pouze
dospělých jedinců.
range (for details see Argyropulo & Pidopličko 1939, Agardžan­jan & Erbaeva 1983, Erbaeva
1988, Erbajeva 1994, and Sen 2003). According to Averianov & Baryšnikov (1992), findings
from the Čirahan River and Urcskij Range are likely to belong to Ochotona transcaucasica.
This species was first described by Vekua (1967) from the Copi Station, Georgia, and due to its
large size and general structure of P3 as Ochotonoides transcaucasica, i.e., belonging to a fossil
genus, which was known that time from China and Hungary (see Boule & Teilhard de Chardin
1928, Teilhard de Chardin & Young 1931, Teilhard de Chardin 1940, and Kretzoi 1962).
With more fossil evidence at hand, Erbaeva (1988) assigned the pika from the Copi Station to
the genus Ochotona. In spite of very similar morphology (small and rounded anteroconid in P3,
posterior mental foramen in mandible below M3, etc.), this species differs from the ocho­to­nids
under study by its larger size; OR of the P3–M3 length is 11.00–12.00 in O. transcaucasica
(Vekua 1967) compared to 8.30–9.85, n=27, in the studied ochotonids of the Middle East. Ocho­
to­na transcaucasica was also reported from the Azih cave in Azerbaijan; the available teeth
are very close in size and morphology to those of pika from the Copi Station (Markova 1982,
Erbaeva 1988). A smaller pika form found at the Azih cave is referred to Ochotona azerica
(Aliev 1969, Gadžiev et al. 1979). This pika is of a similar size (OR of P3–M3 length: 8.7–9.5)
61
as O. rufescens presented in this paper, nevertheless it has a more robust mandible and a larger
anterokonid of P3 (Erbaeva 1988).
Besides the large sized ochotonids from the Middle East, a small sized form was found at the
Güzyurdu 2A site in central Turkey (for details see New Records). Poorly preserved remains
do not allow a precise species assignment. Nevertheless, in its size (L×W of the avai­lable M1
is 1.28×2.44), the pika from Güzyurdu 2A resembles rather O. pusilla than O. rufescens: OR
of L×W of M1 is 1.15–1.45 × 2.00–2.40, n=20 in the extant O. pusilla in contrast to 1.50–1.85
× 2.60–3.15, n=11 in the studied O. rufescens. According to the shape of alveolus, the mesial
hyperloph of P3 is relatively wide and flat; its width is ap­pro­xi­ma­te­ly 75 per cent of the tooth
width. Based on body size, we assign tentatively the material from Güzyurdu 2A to O. cf.
pusilla. A similarly sized pika, referred to Ochotona sp., was reported by Vekua et al. (1981)
from the Toringian site of Bronzovaja in western Georgia; according to its size, it is also placed
into systematical proximity of O. pusilla (Vekua et al. 1981, Averianov & Baryšnikov 1992).
A pika of an uncertain taxonomical position, referred to Ochotona sp., is known from the Middle Pleis­to­ce­ne (the Lower Toringian) site of Emir­ka­ya-2 from the south of Central Anatolia
(Montuire et. al. 1994). Its P3 shares transitional features between O. pusilla and O. rufescens,
and therefore this pika is considered to be an early representative of the former species by some
authors (e.g., Montuire et. al. 1994, Er­ba­je­va 2001), by others it is referred to the latter species
(e.g., Averianov 2001).
Because of the limited data on fossil record, phylogenetic relationships of the taxa under
study with other extant species are not yet clear. Nevertheless, from the morphological point
of view it can be mentioned that O. rufescens as well as the large sized fossil ochotonids from
Transcaucasia and O. pusilla share archaic dental features (i.e., P3 with small and rounded
anteroconid, wide confluence between anteroconid and posteroconid, shallow proto- and pa­ra­f­
le­xid, rounded P2 with a short anteroflexus) and their dentition is much less differentiated than
in other extant species.
Phylogenetic relationships within the genus Ochotona reconstructed using the mi­to­chon­d­rial
gene for cytochrome b (Niu et al. 2004) do not support the current subgeneric classifications (see
Allen 1938, Ellerman & Morrison-Scott 1951, Erbaeva 1988, Erbajeva 1994, and others).
Niu et al. (2004) recognised five groups; they placed O. rufuscens into the Sur­roun­ding Quing­
hai-Tibet Plateau group, i.e. into phylogenetic proximity of the rock-talus-dwel­ling species (O.
macrotis, O roylei, O. iliensis, O. himalayana, O. rutila, O erythrotis, O. gloveri, O. brookei,
and O. muliensis) and intermediate types between the talus- and steppe-dwelling species (O.
ladacensis, O. forresti and O. koslowi). The species divergence occur­red most probably in the
Early Pleistocene and was closely related to the uplifting of the Quinghai-Tibet Plateau and
subsequent climatic changes. Due to the relatively stable en­vi­ron­ment, the differentiation was
not so strong within the Central Asian group, including O. pu­sil­la (Niu et al. 2004). Based on
fossil remains, the structure of teeth, and molecular data, O. pusilla represents a very distinct
and ancient species, which is often considered to be a relic of the Late Pliocene (Erbajeva
1994, Niu et al. 2004).
SOUHRN
V příspěvku jsou prezentovány nové nálezy pišťuch (12 recentních a 1 subfosilní) z území Turecka a Íránu.
Většina recentních nálezů, řazených zde do druhu Ochotona rufescens (Gray, 1842) – pišťucha rezavá,
pochází z horských oblastí Íránu; naše nálezy většinou spadají do dosud známého areálu rozšíření tohoto
druhu. Nález kosterních pozůstatků pišťuchy ve východním Turecku představuje první recentní nález
62
v této zemi a významně posouvá hra­ni­ci dosud známého areálu rozšíření rodu (i celé čeledi Ocho­to­ni­
dae) na Blíz­kém východě. Nálezy ve východním Turecku a severozápadním Íránu (Arménská vysočina)
spadají mor­fo­me­t­ric­ky do variační šíře pišťuchy rezavé, přesto se liší od studovaných pištuch z Íránu
větší velikostí a trochu odlišnou morfologií třetího třenového zubu. Nicméně vzhledem k nepočetnému
materiálu z Arménské vy­so­či­ny je věrohodné srovnání problematické a studované pišťuchy jsou pro­vi­zor­
ně determinovány jako O. cf. rufescens. Nově nálezený subfosilní jedinec pišťuchy ze střední Anatolie je
výrazně menší než předešlé pištuchy a jedná se s největší pravděpodobností o pišťuchu stepní – O. pusilla
(Pallas, 1768), rozšířenou v současné době nejblíže v Předkavkazí. Uvedené nálezy byly též morfologicky
srovnány s ostat­ní­mi pišťuchami regionu, a to jak v kontextu neontologickém, tak pa­le­on­to­lo­gic­kém.
ACKNOWLEDGEMENTS
We are very grateful to Ivan Horáček and Vladimír Vohralík (Charles University, Prague) for providing
us with the material and information related to Güzyurdu (Turkey) and for numerous valuable comments.
We thank A. Reiter, J. Mlíkovský, J. Sádlová, J. Sádlo, and R. Šumbera for their help with data and
material collection in the field. The research was partly supported by grants GA ČR 206/05/2334 and
MK00002327201.
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Lynx (Praha), n. s., 37: 67–78 (2006).
ISSN 0024–7774
Bats of the Čerchovský les Mts. and the first record of the Greater horseshoe
bat (Rhinolophus ferrumequinum) in western Bohemia (Czech Republic)
(Chiroptera)
Netopýři Čerchovského lesa a první nález vrápence velkého (Rhinolophus
ferrumequinum) v západních Čechách (Chiroptera)
Jaroslav Červený1,2, Václav Fišr3, Petr Faschingbauer4 & Luděk Bufka5
Institute of Vertebrate Zoology AS CR, Květná 8, CZ–603 65 Brno, Czech Republic;
Czech Agriculture University, Faculty of Forestry and Environment, Kamýcká 1176,
CZ–165 21 Praha 6, Czech Republic; [email protected]
3
Domažlice Municipal Forests, Tyršova 611, CZ–344 01 Domažlice, Czech Republic; [email protected]
4
Stará Huť 14, Nemanice, CZ–344 01 Domažlice 1, Czech Republic; [email protected]
5
PLA and NP Šumava Administration, Sušická 399, CZ–342 69 Kašperské Hory, Czech Republic;
[email protected]
1
2
received on 8 December 2006
Abstract. The study summarises results of bat survey in the southern part of the Český les Mts. in the
period 2004–2006. The following 17 species of bats (73.9% of bat fauna of the Czech Republic) have
been confirmed to occur in the study area: Rhinolopus ferrumequinum, Myotis mystacinus, M. brandtii, M.
bechsteinii, M. nattereri, M. myotis, M. daubentonii, Vespertilio murinus, Eptesicus serotinus, E. nilssonii,
Pipistrellus pipistrellus s. l., P. nathusii, Nyctalus noctula, N. leisleri, Barbastella barbastellus, Plecotus
auritus, and P. austriacus. Five species were recorded there for the first time: R. ferrumequinum, E. sero­
tinus, P. nathusii, N. noctula and N. leisleiri. Numerous records of the “forest species”: M. bechsteinii, V.
murinus, E. nilssonii, P. nathusii, N. leisleri, and B. barbastellus are of particular importance. Since the
record of R. ferrumequinum in the study area represents the only second record of this species in Bohemia
and the seventh record in the Czech Republic after 1950, is seems to be of high importance.
INTRODUCTION
Although western Bohemia has been subjected to a considerable chiropterological survey (e.g.,
Hůrka 1973, 1986, 1989, Bufka et al. 2001, Dvořák et al. 2003), the Český les Mts. have been
rather neglected. The comparison with the situation in the neighbouring Šumava Mts., which
ranks among the best chiropterologically studied regions in the Czech Republic (Anděra &
Červený 1994), is particularly striking. The reasons may be not only in the absence of easily
controlable winter roosts and in difficult detectability of bats in forest habitats, but especially in
very constrained opportunities for zoological research in this area at times, when chiropterology
was flourishing in the then Czechoslovak Socialist Republic. Till 1989 much of this geomorphologically very narrow and relatively long stripe at the Czech-German border was closed for
political and military regions. As a result, only 12 bat species, representing just 52.2% of the
currently known Czech bat fauna were recorded there. However, neither the bordering area of
67
the Oberpfälzer Wald in Bavaria has been satisfactorily studied from the given point of view
(Meschede & Rudolph 2004).
Regarding these facts we decided to focus on subsequent complementation of our knowledge
on the bat occurrence in a broader region of the Český les Mts. In this paper we present results
of the bat survey in the southern part of the region, namely in the Čerchovský les Mts.
STUDY AREA
The studied area of the Čerchovský les Mts. (Fig. 1) encompasses the whole southern third of the Český
Les Protected Landscape Area, with the total area of about 250 km2, and is delimited by the state border
and the Oberfälzer Wald in the west and south, by Kateřinská kotlina (a basin) and Přimdský les Mts in
the north, by Chodská pahorkatina (hills) and Českokubická vrchovina (highlands) in the east. The area
is characterized by several elongated mountain ridges, separated by deep valleys. It is composed of three
highlands: Haltravská hornatina, Nemanická vrchovina, and Ostrovská vrchovina. The highest point is
Mt. Čerchov (1042 m a. s. l.), the average altitude of the area being 633.8 m a. s. l. The climate is mostly
mild warm, above 700 m a. s. l. cold. The average annual temperature ranges – according to the altitude
– from 8 to 4.5 °C, annual precipitation from 700 to 1000 mm.
Most of the area (almost 80%) is covered by submontane or montane forests, with prevailing beech,
e.g. acidophilous beechwood and fir-beechwood like Luzulo-Fagion or Eu-Fagetion, locally watered
firs (Equiseto-Abietetum) and locally limited also watered spruces (Mastigobryo-Piceetum). Currently,
secondary spruce monocultures prevail, yet still many mixed forests (spruce, beech, and fir, with intermixed ash, maple and sycamore), are preserved. The proportion of beech in the whole region makes up
about 15%, in the southern part of Čerchovský les, and in the very region of Čerchov Mt. almost 50%.
The average age of the forests is about 70 years, except for the area of Čerchov where more than 50% of
trees are 111–180 years old.
Remark. The locality ‘Černá Řeka’ (gallery) in this study is identical with the locality name ‘Jindřichova
Hora’ used by L. Hůrka in his papers.
METHODS
During the period 2004–2006, bats were mainly observed in their summer and winter roosts. Bats were
also caught using mistnets along their flight corridors in the forest or above water.
The distribution of each bat species is given using the standard grid map (KFME system), as usual in
faunistic surveys (Anděra & Červený 1994, Bufka et al. 2001). The individual findings are specified as
follows: square number, site name, altitude, date of finding, and habitat or shelter. For each species, the
complete list of localities arranged alphabetically in particular squares is given.
Abbreviations. M – male, F – female, F gr – gravid female, F lact – lactating female, ex. – individual(s),
s.i. – sex indetermined, S – summer record (15 April – 15 October), W – winter record (16 October
– 14 April).
RECORDS AND DISCUSSION
Greater horseshoe bat, Rhinolophus ferrumequinum (Schreber, 1774)
6542: Černá Řeka (560 m a. s. l.), 4 December 2006, gallery, 2 M.
Remarks. The finding of this bat species represents the first record of its occurrence in the
study area, but mainly the second record in Bohemia and seventh record in the whole Czech
Republic after 1950. At the same time, it represents the highest situated locality of its occurrence in the country. From the Czech Republic, there have been so far only few less reliable
68
41
42
43
63
1
3
2
64
4
CZE
5
GER
65
7
10
8
9
6
11
12
14
13
16
17
19
18
20
27
66
22
21
23
24
28
25
26
29
30
31
Fig. 1. Schematic map of the study area (grey – forested area; dashed line – border of the area under
study; bigger closed circle – important settlement; smaller closed circle – locality under study; cross
– important hill).
Obr.1. Schematická mapa sledované oblasti (šedé pozadí – lesnatá oblast, přerušovaná linie – hranice
Čerchovského lesa, větší plný kruh – důležité sídlo, menší plný kruh – sledovaná lokalita; křížek – důležitý vrchol).
Sites / lokality: 1 – Železná, 2 – Smolov, 3 – Bělá nad Radbuzou, 4 – Újezd u Svatého Kříže, 5 – Rybník,
6 – Hraničná, 7 – Závist, 8 – Pivoň, 9 – Vranov, 10 – Mnichov, 11 – Lučina, 12 – Nemanice, 13 – Stará
Huť, 14 – Díly, 15 – Klenčí pod Čerchov, 16 – Černá Řeka, 17 – Jindřichova Hora, 18 – Capartice,
19 – Výhledy, 20 – Chodov, 21 – Čerchov Mt., 22 – Čerchov, Hánovka (Fig. 2), 23 – Čerchov, Na zlomu,
24 – Čerchov, Zámeček, 25 – Čerchov, Rajská, 26 – Bystřice (Fig. 3), 27 – Pec, 28 – Babylon, 29 – Česká
Kubice, Zelená chýše, 30 – Česká Kubice, Česká studánka, 31 – Česká Kubice.
69
data on presence of this species from the 19th and beginning of the 20th centuries; after 1950
singular occurrence is known only from six hibernating sites (Hanák & Anděra 2005). The
last finding originates from the Moravian Karst in 1979 (Gaisler 1997). From Bohemia, so
far only one record has been published, namely from the Mořina mine near Karlštejn in 1962
(Hanák 1962). In the neighbouring parts of Bavaria, approximately 80 km from the presented
locality, in the Danube valley (between Ingolstatd and Regensburg), a stable population of this
species occurs and local transitions are known there (Meschede & Rudolph 2004). The size
of the Bavarian population has been found to grow (S. Morgenroth ad verb.) and hence, it is
well possible that the record described here represents a winter excursion. The ability of long
distance wandering in this species has been evidenced out of the Czech Republic by means of
ringing (Gaisler et al. 2003).
Whiskered bat, Myotis mystacinus (Kuhl, 1817)
6442: Bělá nad Radbuzou (480 m a. s. l.), 18 May 2006, shutter of a hut, 2 M; Rybník (530 m a. s. l.),
17 May 2006, shutter of a cottage, 1 M; 3 October 2006, shutter of a cottage, 2 M; 6542: Černá Řeka
(560 m a. s. l.), 27 January 2004, gallery, 4 ex. s.i.; Jindřichova Hora (670 m a. s. l.), 3 October 2006,
shutter of a cottage, 1 F M; Stará Huť u Nemanic (560 m a. s. l.), 22 June 2005, shutter of a hut, 3 M;
12 August 2005, shutter of a hut, 1 M; 6642: Čerchov – Na zlomu (875 m a. s. l.), 17 May 2006, wooden panelling of a hut, 1 M; 6643: Babylon – Černý rybník (470 m a. s. l.), 7 May 2006, netting above
a stream near the fishpond, 2 M.
Previous records. 6542: Černá Řeka (560 m a. s. l.), gallery, W, 1979–2002 (Hůrka 1986, Bufka et al.
2001, Dvořák et al. 2003); Díly (600 m a. s. l.), loft of a barn, S, 1968 (Hůrka 1973); 6642: Čerchov,
1932 (Gaisler 1956).
Remarks. The whiskered bat was found at nine summer localities (470–875 m a. s. l.) in the
study area. Summer records are often related to human residences at forest edges or close to
water bodies, like in the Šumava Mts. (Anděra & Červený 1994). No breeding colony has
been found so far. Individual bats were recorded to occupy different fissures of buildings, mainly behind window shutters. The only regular winter roost is in the gallery near Černá Řeka
(560 m a. s. l.). Another record of this species is known from the nearby region – the locality
Diana (Kůs 1999). The species is considered to be common in western Bohemia (Hůrka 1989,
Bufka et al. 2001).
Brandt’s bat, Myotis brandtii (Eversmann, 1845)
6442: Rybník (525 m a. s. l.), 17 May 2006, netting above the Radbuza river, 1 M; 6542: Stará Huť
u Nemanic (560 m a. s. l.), 22 June 2005, shutter of a hut, 1 M.
Previous records. 6542: Černá Řeka (560 m a. s. l.), gallery, W, 2002–2003 (Dvořák et al. 2003).
Remarks. Two records of the Brandt’s bat represent the first summer occurrence (525 and
560 m a. s. l.) of this species in the study area. The only winter roost is known in the gallery
near Černá Řeka (560 m a. s. l.). Habitats occupied by this species are often identical with those
of M. mystacinus. Infrequent records indicate rare occurrence not only in the area under study,
but also in western Bohemia in general (Hůrka 1989, Anděra & Červený 1994, Bufka et al.
2001), as well as in the whole Czech Republic (Hanák & Anděra 2006).
70
Bechstein’s bat, Myotis bechsteinii (Kuhl, 1917)
6442: Rybník (525 m a. s. l.), 17 May 2006, netting above the Radbuza river, 1 M; 6542: Černá Řeka
(560 m a. s. l.), 27 January 2004, gallery, 5 ex. s.i.; 6 February 2006, 1 ex. s.i., 4 December 2006, 2 ex s.i.;
6642: Bystřice (535 m a. s. l.), 11 August 2005, netting above a small fishpond in the forest near an abandoned village, 1 M; Čerchov – Zámeček (670 m a. s. l), 23 June 2005, netting near a small fishpond in the
forest, 1 M; 6643: Česká Kubice – Zelená chýše (642 m a. s. l.), 4 July 2005, netting in the forest, 1 M.
Previous records. 6542: Černá Řeka (560 m a. s. l.), gallery, W, 1971–2003 (Hůrka 1973, Bufka et al.
2001, Dvořák et al. 2003).
Remarks. Four records of the Bechstein’s bat represent the first summer occurrence of this
species in the study area. A low number of records reflects difficult detectability of this species,
rather than its relative rarity (as discussed in Hůrka 1989, Anděra & Červený 1994, Bufka et
al. 2001, Hanák & Anděra 2006). The records come from middle altitudes (525–670 m a. s. l.),
always from habitats with the presence of deciduous forests, mainly beech, or mixed forests.
The only regular winter roost is in the gallery near Černá Řeka (560 m a. s. l.). Another record
of this species is known from the nearby region – the locality Přimda (Kůs 1999). The species
is considered to be rare in western Bohemia (Hůrka 1989, Bufka et al. 2001).
Natterer’s bat, Myotis nattereri (Kuhl, 1817)
6442: Rybník (530 m a. s. l.), 17 May 2006, shutter of a cottage, 1 M; 6542: Capartice (760 m a. s. l.),
12 August 2005, shutter of a cottage, 1 M; Černá Řeka (560 m a. s. l.), 27 January 2004, gallery, 5 ex. s.i.,
4 December 2006, 1 ex. s.i.; 6 February 2006, 5 ex. s.i.; 6642: Bystřice (555 m a. s. l.), 11 August 2005,
netting in front of the cellar of a school in an abandoned village, 2 M; Bystřice (535 m a. s. l.), 11 August
2005, netting above a small fishpond in the forest near an abandoned village, 1 M; 6643: Česká Kubice
(505 m a. s. l.), 13 August 2005, netting above a fishpond, 1 F.
Previous records. 6542: Černá Řeka (560 m a. s. l.), gallery, W, 1974–2003 (Hůrka 1986, Bufka et al.
2001, Dvořák et al. 2003).
Remarks. The Natterer’s bat is typical for lower altitudes with plenty of wetlands (Hůrka 1973,
1989, Anděra & Červený 1994, Bufka et al. 2001, Hanák & Anděra 2006). Six localities of
occurrence (505–760 m a. s. l.) were found in the study area, but only some of them were situated
near water. No breeding colony has been found so far. Individual bats were hidden in different
fissures of buildings, mainly behind window shutters. The only regular winter roost is known
in the gallery near Černá Řeka (560 m a. s. l.). The species is considered to be quite common
in western Bohemia (Hůrka 1989, Bufka et al. 2001, Hanák & Anděra 2006).
Greater mouse-eared bat, Myotis myotis (Borkhausen, 1797)
6442: Bělá nad Radbuzou (440 m a. s. l.), 18 May 2006, town square, 1 M dead; 6542: Černá Řeka
(560 m a. s. l.), 27 January 2004, gallery, 11 ex. s.i.; 6 February 2006, 18 ex. s.i., 4 December 2006, 12 ex.
s.i.; Pivoň (590 m a. s. l.), 8 November 2006, basement of the monastery, 2 ex. s.i.
Previous records. 6542: Černá Řeka (560 m a. s. l.), loft of a school, S, 1967 (Hůrka 1973), gallery,
W, 1969–1998 (Hůrka 1973, Hůrka 1986, Bufka et al., 2001), Pivoň (590 m a. s. l.), basement of the
monastery, W, 1971–2003 (Hůrka 1973, Bufka et al. 2001, Dvořák et al. 2003).
Remarks. The greater mouse-eared bat is one of the most common species of western Bohemia
(Hůrka 1973, Anděra & Červený 1994, Bufka et al. 2001, Hanák & Anděra 2006), however,
this does not apply for the study area. Here it is known mainly from two winter roosts (560 and
71
590 m a. s. l.). The only two summer records (440 and 560 m a. s. l.) come form the foothills.
This species was recorded also in the nearby region – the locality Světce near Tachov (Hůrka
1988).
Daubenton’s bat, Myotis daubentonii (Kuhl, 1817)
6442: Bělá nad Radbuzou (485 m a. s. l.), 18 May 2006, netting above a small fishpond, 1 M; Rybník
(525 m a. s. l.), 17 May 2006, netting above the river Radbuza, 1 F; 6542: Černá Řeka (560 m a. s. l.),
27 January 2004, gallery, 11 ex. s.i.; 3 September 2005, netting in front of a gallery, 1 M; 6 February 2006,
18 ex. s.i., 4 December 2006, 6 ex s.i.; Lučina-Grafenried (600 m a. s. l.), 7 November 2006, cellar of
a destroyed building, 1 M; 6642: Bystřice (540 m a. s. l.), 23 June 2005, netting above a small fishpond
in the forest, 1 M; Bystřice (535 m a. s. l.), 11 August 2005, netting above a small fishpond in the forest
near an abandoned village, 1 F; Bystřice (545 m a. s. l.), 7 November 2006, the cellar of a building in
an abandoned village, 1 ex. s.i.; 6643: Babylon – Černý rybník (470 m a. s. l.), 7 May 2006, netting
above a stream near the fishpond, 2 M, 3 F; Česká Kubice (505 m a. s. l.), 13 August 2005, netting above
a fishpond, 2 F.
Previous records. 6542: Černá Řeka (560 m a. s. l.), gallery, W, 1971–2003 (Hůrka 1973, Bufka et al.
2001, Dvořák et al. 2003).
Remarks. A common species, recorded at nine localities (470–600 m a. s. l.). The Daubenton’s
bat occurs in various habitats, but forages mainly above water, and its distribution is significantly
influenced by the presence of various types of water bodies. No breeding colony or summer
shelters have been found so far. The only regular winter roost is known in the gallery near Černá
Řeka (560 m a. s. l). The species is also considered to be common in western Bohemia (Hůrka
1973, 1989, Anděra & Červený 1994, Bufka et al. 2001, Hanák & Anděra 2006).
Parti-coloured bat, Vespertilio murinus Linnaeus, 1758
6441: Smolov (485 m a. s. l.), 18 May 2006, shutter of a hut, 2 M; 6442: Rybník (530 m a. s. l.), 3 October
2006, shutter of a cottage, 1 M; 6542: Stará Huť u Nemanic (560 m a. s. l.), 12 August 2005, shutter of
a hut, 1 M; 3 September 2005, 2 M; 6642: Bystřice – třešňovka (540 m a. s. l.), 9 November 2006, wooden
hunting lookout, 1 M dead; 6643: Pec (510 m a. s. l.), 11 June 2005, shutter of a cottage, 1 M.
Previous records. 6542: Díly (600 m a. s. l.), room in a building, S, 1967 (Hůrka 1971a).
Remarks. The parti-coloured bat was found at six summer localities (485–600 m a. s. l.) in
the study area. No breeding or summer male colonies have been found so far, only individual
bats hidden in different fissures of buildings (mainly behind window shutters) or fissures in
a wooden hunting lookout. This species was formerly considered to be very rare in western
Bohemia (Hůrka 1973, 1989, Bufka et al. 2001), but it is relatively common in some habitats
of the Šumava Mts. (Anděra & Červený 1994).
Serotine Eptesicus serotinus (Schreber, 1774)
6442: Rybník (530 m a. s. l.), 17 May 2006, loft of a house, 1 M; 6542: Nemanice (530 m a. s. l.), 23 June
2005, loft of the church, 1 ex. s.i.
Remarks. Two records of the serotine represent the first occurrence of this species in the study
area. Both are situated at rather middle altitudes (530 m a. s. l.), in an open landscape and in
human settlements in lofts of houses. The species is common throughout western Bohemia
(Hůrka 1973, Bufka et al. 2001).
72
Northern bat, Eptesicus nilssonii (Keyserling et Blasius, 1839)
6442: Bělá nad Radbuzou (485 m a. s. l.), 18 May 2006, netting above a small fishpond, 1 F gr; 6542:
Černá Řeka (560 m a. s. l.), 27 January 2004, gallery, 1 ex. s.i.; Pivoň (580 m a. s. l.), 8 May 2006,
netting above a small fishpond, 1 F; Stará Huť u Nemanic (560 m a. s. l.), 22 June 2005, netting above
a small fishpond, 1 F lact.; 6 May 2006, shutter of a hut, 1 M; Vranov (650 m a. s. l.), 7 May 2006, loft
of a building, breeding colony of approx. 50 ex. (2 F gr were examined); Závist (590 m a. s. l.), 7 May
2006, wooden panelling of a building, 1 M; 6642: 1 M; Čerchov (1040 m a. s. l.), 17 May 2006, panelling
of a building on the top of the hill, 1 M; Čerchov – Zámeček (670 m a. s. l.), 23 June 2005, netting near
a small fishpond in the forest, 1 F; 6643: Pec (520 m a. s. l.), 11 June 2005, loft of the building, breeding
colony of approx. 80 ex. (1 F gr was examined).
Previous records: 6542: Černá Řeka (560 m a. s. l.), gallery, W, 1983–2003 (Hůrka 1986, Bufka et al.
2001,); Díly (600 m a. s. l.), loft of a dwelling house, breeding colony of 70 ex., S, 1976 (Hůrka 1986).
Remarks. The northern bat had been considered to be very rare for a long time, but now it seems
to be a relatively common species not only in the study area, but also in the rest of western Bohemia (Anděra & Červený 1994, Bufka et al. 2001). Morever, its occurrence was confirmed at
10 localities (485–1040 m a. s. l.), mainly in woodlands. Breeding colonies were found (50–80
individuals) in lofts of dwelling houses or weekend cottages, individual bats were hidden in
different fissures of buildings. The only winter roost is in the gallery near Černá Řeka (560 m
a. s. l.). Other findings of this species come from the nearby region – the localities Chodovská
Huť (Hůrka 1973) and Přimda (Kůs 1999). In the neighbouring Oberpfälzer Wald, a breeding
colony is also known (Merkel – Wallner et al.1987) and many records were made using bat
detectors (Skiba 1987).
Common pipistrelle, Pipistrellus pipistrellus (Schreber, 1774)
6542: Výhledy (680 m a. s. l.), 11 June 2005, 3 October 2006, shutter of a building, 2 M; 6642: Čerchov-Hánovka (900 m a. s. l.), 29 August 2005, netting in the forest, 1 M, 1 F; 6643: Pec (510 m a. s. l.),
11 June 2005, shutter of a building, 3 M.
Previous records. 6441: Železná (525 m a. s. l.), loft of the school, S, 1968 (Hůrka 1973); 6542: Klenčí
pod Čerchovem (495 m a. s. l.), loft of a house, S, 1965 (Hůrka 1973).
Remarks. Probably a common species in the study area, its occurrence was confirmed at five
localities (495–900 m a. s. l.). Breeding colonies have not yet been found, individual bats were
hidden in fissures of buildings. Although several winter roosts are known from the foothills
of the Čerchovský les (Hůrka 1973, Bufka et al. 2001), no hibernating bats were recorded in
the study area. The species is considered to be relatively common in western Bohemia (Hůrka
1989, Bufka et al. 2001). However, confusion with a very similar species Pipistrellus pygmeus
is possible.
Nathusius’ pipistrelle, Pipistrellus nathusii (Keyserling et Blasius, 1839)
6642: Bystřice (535 m a. s. l.), 4 July 2005, netting above a small fishpond in the forest near an abandoned
village, 1 F lact; 6643: Česká Kubice (505 m a. s. l.), 13 August 2005, netting above a fishpond, 1 M;
Česká Kubice – Zelená chýše (642 m a. s. l.), 4 July 2005, shelter in the roof of a hut, 1 M.
Remarks. Three findings of the Nathusius’ pipistrelle represent the first records of this species
in the study area. All are situated at rather middle altitudes (505–642 m a. s. l.) in forested areas
near water bodies. The species is widespread in some parts of the Czech Republic (Jahelková
73
et al. 2000), however, this does not apply for western Bohemia where it is very rare (Červený
& Bufka 1999, Bufka et al. 2001).
Noctule, Nyctalus noctula (Schreber, 1774)
6542: Pivoň (580 m a. s. l.), 8 May 2006, netting above a small fishpond, 1 F; 6643: Babylon – Černý
rybník (470 m a. s. l.), 7 May 2006, netting above a stream near a fishpond, 1 M.
Remarks. Two findings of the noctule represent the first record of this species in the study area.
The species is quite common at lower altitudes of western Bohemia (Hůrka 1989, Bufka et al.
2001). Both our records (470 and 580 m a. s. l.) come from forest habitats with water bodies.
Similarly as in the lower Šumava Mts. (Červený & Bürger 1989), a sympatric occurrence of
N. nocula and N. leisleri was recorded at one locality (Pivoň).
Leisler’s bat, Nyctalus leisleri (Kuhl 1817)
6442: Rybník (525 m a. s. l.), 17 May 2006, netting above the river Radbuza, 1 F; 6542: Pivoň (580 m
a. s. l.), 8 May 2006, netting above a small fishpond, 1 M, 2 F; 6642: Čerchov-Rajská (830 m a. s. l.),
13 November 2005, wooden hunting lookout, 2 F; 6643: Česká Kubice – Zelená chýše (642 m a. s. l.),
4 July 2005, netting in a forest, 1 F lact.
Remarks. Four captures of the Leisler’s bat represent the first record of this species in the
study area. The records were made at relatively higher altitudes (525–830 m a. s. l.), always in
habitats with deciduous forests, mainly beech, or with mixed forests. Similarly as in the lower
Šumava Mts. (Červený & Bürger 1989), a sympatric occurrence of N. nocula and N. leisleri
was recorded at one locality (Pivoň). This species is considered to be extremly rare in western
Bohemia (Hůrka 1989, Bufka et al. 2001).
Barbastelle, Barbastella barbastellus (Schreber, 1774)
6441: Smolov (485 m a. s. l.), 18 May 2006, shutter of a cottage, 1 M; 6542: Černá Řeka (560 m a. s. l.),
27 January 2004, gallery, 7 ex. s.i.; 6 February 2006, 9 ex. s.i.; Stará Huť u Nemanic (560 m a. s. l.),
6 May 2006, shutter of a cottage, 1 M; 6642: Bystřice (555 m a. s. l.), 11 August 2005, netting in front of
a cellar of the school in an abandoned village, 1 M.
Previous records. 6542: Černá Řeka (560 m a. s. l.), gallery, W, 1976–1980 (Hůrka 1986); Pivoň, (590 m
a. s. l.), basement of the monastery, W, 1971–1975 (Hůrka 1973, Bufka et al. 2001).
Remarks. The barbastelle was found at three summer localities (485–560 m a. s. l.) in the study
area. Summer findings are often related to human residences at forest edges, like in the Šumava Mts. (Anděra & Červený 1994). No breeding colony has been found so far. Individual bats
were hidden behind window shutters. The only two winter roosts are situated at the altitudes of
560 and 590 m a. s. l. This species is considered to be relatively common in western Bohemia
(Hůrka 1989, Anděra & Červený 1994, Bufka et al. 2001, Hanák & Anděra 2005).
Brown long-eared bat, Plecotus auritus (Linnaeus, 1758)
6442: Bělá nad Radbuzou (450 m a. s. l.), 18 May 2006, shutter of a building, 4 F; Rybník (530 m a. s. l.),
23 June 2005, loft of a house, breeding colony of approx. 15 ex.; 6541: Hraničná (685 m a. s. l.), 8 November 2006, cellar of a building, 2 F; 6542: Černá Řeka (560 m a. s. l.), 27 January 2004, gallery, 6 ex.
74
Figs. 2, 3. Sites of bat netting in the Čerchovský les Mts. 2 – Mixed forest on Mount Čerchov, Hánovka
(photo by V. Fišr) (above). 3 – Deforested area near the abandoned village Bystřice with Mt. Čerchov in
the horizon (photo by J. Červený) (below).
Obr. 2, 3. Lokality nettingu netopýrů v Čerchovském lese. 2 – Smíšený lesní porost na lokalitě Čerchov,
Hánovka (foto V. Fišr) (nahoře). 3 – Odlesněná oblast u bývalé obce Bystřice s vrcholem Čerchova
v pozadí (foto J. Červený) (dole).
75
s.i.; 3 September 2005, netting in front of a gallery, 1 M; 6 February 2006, gallery, 9 ex. s.i, 4 December
2006, 1 ex. s.i.; Nemanice (530 m a. s.l.), 23 June 2005, loft of the church, breeding colony of approx.
8–10 ex.; Pivoň (590 m a. s. l.), 8 November 2006, basement under the monastery, 1 F; Závist (590 m
a. s. l.), 7 May 2006, loft of a cottage, breeding colony of approx. 10 ex.; 6642: Bystřice (555 m a. s. l.),
11 August 2005, netting in front of a cellar of the school in an abandoned village, 2 M, 1 F; 6642: Bystřice
(545 m a. s. l.), 11 August 2005, netting in front of a cellar of a building in an abandoned village, 1 F;
Bystřice (535 m a. s. l.), 11 August 2005, netting above a small fishpond in the forest near an abandoned
village, 1 F; Čerchov (1040 m a. s. l.), 17 May 2006, panelling of a building on the top of the hill, 1 M;
Čerchov – Zámeček (670 m a. s. l.), 23 June 2005, netting near a small fishpond in the forest, 1 F; 6643:
Česká Kubice – Česká Studánka (650 m a. s. l.), 17 July 2005, bird nestbox in the forest, 8 F.
Previous records. 6442: Újezd u Svatého Kříže (450 m a. s. l.), loft of the church, S, 1971 (Hůrka 1973);
6542: Černá Řeka (560 m a. s. l.), gallery, W, 1971–2003 (Hůrka 1973, Bufka et al. 2001, Dvořák et
al. 2003); Díly (600 m a. s. l.), loft of the chapel, S, 1967 (Hůrka 1971b); Mnichov (725 m a. s. l.), loft
of the church, S, 1974 (Hůrka 1986); Pivoň, 590 m a. s. l., basement of the monastery, W, 1971–1972
(Hůrka 1973).
Remarks. The brown long-eared bat is the most common species in the study area. It was found
at 14 summer localities (450–1040 m a. s. l.) and in three winter roosts (560–685 m a. s. l.).
Summer findings are often related to the periphery of human settlements in forested areas, like
in the Šumava Mts. (Anděra & Červený 1994). Breeding colonies (8–15 individuals) were
found in lofts of dwelling houses or in a bird nestbox, individual bats were hidden in different
fissures of buildings. Other findings of this species are known from the nearby parts of the
Český les Mts. – the localities Halže and Chodovská Huť (Hůrka 1973), Mílov and Přimda
(Kůs 1999). This species is considered to be very common in western Bohemia (Hůrka 1989,
Anděra & Červený 1994, Bufka et al. 2001).
Grey long-eared bat, Plecotus austriacus (Fischer, 1829)
6542: Pivoň (580 m a. s. l.), 8 May 2006, netting above a small fishpond, 1 M; 8 November 2006, basement
of the monastery, 2 M; Stará Huť u Nemanic (560 m a. s. l.), 7 November 2006, cellar of a destroyed
building, 1 M; 6643: Česká Kubice (505 m a. s. l.), 13 August 2005, netting above a fishpond, 1 M.
Previous records: 6542: Díly (600 m a. s. l.), loft of the chapel, S, 1967 (Hůrka 1973), Chodov (500 m
a. s. l.), loft of the school, S, 1966 (Hůrka 1971).
Remarks. The grey long-eared bat was found only at four summer localities (500–600 m a. s. l.)
and in two winter roosts (560 and 580 m a. s. l.). Summer findings are situated in an open landscape and in human settlements in lofts of houses. Other records of this species come from the
nearby region – the locality Halže (Hůrka 1973). This species is considered to be very common
throughout western Bohemia (Hůrka 1989, Anděra & Červený 1994, Bufka et al. 2001).
SOUHRN
Práce shrnuje výsledky výzkumu netopýrů Čerchovského lesa, který zaujímá celou jižní třetinu Chráněné krajinné oblasti Český les o rozloze přibližně 250 km2 a je vymezen státní hranicí (resp. Oberfälzer
Wald) na západě a jihu, Kateřinskou kotlinou a Přimdským lesem na severu, Chodskou pahorkatinou
a Českokubickou vrchovinou na východě. Území je charakteristické několika podélnými hřbety oddělenými hlubokými údolími. Skládá se ze tří geomorfologických okrsků: Haltravské hornatiny, Nemanické
vrchoviny a Ostrovské vrchoviny. Nejvyšší kótou je Čerchov (1042 m n. m.), střední výška území je
633,8 m n. m. Podnebí je převážně mírně teplé (MT 3), nad 700 m n. m. chladné (CH 7). Průměrná roční
teplota se podle nadmořské výšky pohybuje od 8 do 4,5 °C, průměrné roční srážky od 700 do 1000 mm.
76
Většinu území pokrývají lesy submontánního až montánního stupně, lesnatost území dosahuje hodnoty
téměř 80 %. Rekokonstrukčně převládají acidofilní bučiny a jedlobučiny (Luzulo-Fagion), podmáčené
jedliny (Equiseto-Abietetum), místy květnaté bučiny (Eu-Fagetion), omezeně podmáčené smrčiny (Mas­
tigobryo-Piceetum). V současnosti sice převládají stanoviště sekundárních monokultur smrku, přesto se
však zde zachovalo více smíšených porostů (smrk, buk a jedle s vtroušeným jasanem, javorem a klenem),
než jinde v České republice. Podíl buku v celé oblasti činí okolo 15 %, v jižní části Čerchovského lesa,
a zvláště pak v oblasti samotného Čerchova téměř 50 %. Průměrný věk lesních porostů je přibližně 70 let,
v oblasti Čerchova je však více než 50 % ve věku 111–180 let.
V letech 2004–2006 byl ve sledované oblasti potvrzen výskyt 17 druhů netopýrů, což představuje 73,9 %
netopýří fauny celé České republiky. Zjištěni byli: vrápenec velký (Rhinolophus ferrumequinum), netopýr
vousatý (Myotis mystacinus), netopýr Brandtův (Myotis brandtii), netopýr velkouchý (Myotis bechsteinii),
netopýr řasnatý (Myotis nattereri), netopýr velký (Myotis myotis), netopýr vodní (Myotis daubentonii),
netopýr pestrý (Vespertilio murinus), netopýr pozdní (Eptesicus serotinus), netopýr severní (Eptesicus
nilssonii), netopýr hvízdavý (Pipistrellus pipistrellus), netopýr parkový (Pipistrellus nathusii), netopýr
rezavý (Nyctalus noctula), netopýr stromový (Nyctalus leisleri), netopýr černý (Barbastella barbastellus),
netopýr ušatý (Plecotus auritus) a netopýr dlouhouchý (Plecotus austriacus). Druhy R. ferrumequinum,
E. serotinus, P. nathusii, N. noctula a N. leisleri byly zjištěny v oblasti poprvé. Nejhodnotnější údaje
představují početné nálezy tzv. “lesních druhů”: M. bechsteinii, V. murinus, E. nilssonii, P. nathusii, N.
leisleri a B. barbastellus, které jsou v České republice obecně považovány za druhy vzácné, řídké, nebo
alespoň méně běžné.
Nález Rh. ferrumequinum je zcela ojedinělý a zárověň velmi významně přispívá k poznání výskytu
tohoto druhu u nás. Je teprve druhým prokázáným výskytem v Čechách a sedmým v celé České republice
po roce 1950. Zároveň je svou nadmořskou výškou 560 m naší nejvýše položenou lokaltu výskytu. Nález
pravděpodobně představuje zálet (cca 80 km) na zimoviště z nejbližší známé letní populace v údolí Dunaje
mezi Ingolstatdem a Regensburgem v sousedním Bavorsku.
ACKNOWLEDGEMENTS
We express our thanks to P. Cehláriková and J. Kadera from the Administration of the Český les Protected Landscape Area and H. Burda who kindly revised the manuscript. Special thanks go to M. Anděra
(National Museum, Prague) and L. Dvořák (Šumava National Park) who took part in some field work.
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a summary in English).
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Westböhmen. Folia Mus. Rer. Natur. Bohem. Occid., Zool., Plzeň, 27: 35–55.
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Merkel-Wallner G., Mühlbauer H.& Heller K.-G., 1987: Ein Wochenstubennachweis der Nordfledermaus, Eptesicus nilssoni (Keyserling et Blasius, 1839) in der Oberpfälz. Myotis, 25: 37–40.
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78
Lynx (Praha), n. s., 37: 79–93 (2006).
ISSN 0024–7774
Species composition, spatial distribution and population dynamics
of bats hibernating in Wisłoujście Fortress, Polish Baltic Sea Coast
(Chiroptera: Vespertilionidae)
Skład gatunkowy, rozmieszczenie przestrzenne i dynamika populacji nietoperzy
zimujących w Twierdzy Wisłoujście na polskim Pobrzeżu Bałtyku
(Chiroptera: Ve­sper­ti­li­o­ni­dae)
Mateusz Ciechanowski1, Agnieszka Przesmycka1 & Konrad Sachanowicz2
1
Academic Chiropterological Circle of Polish Society for Nature Protection “Salamandra”,
Department of Vertebrate Ecology and Zoology, University of Gdańsk, Legionów 9,
PL–80-441 Gdańsk, Poland; mat­ci­[email protected] [MC]; [email protected] [AP]
2
Department of Animal Ecology, Ni­co­laus Copernicus University, Gagarina 9, PL–87-100 Toruń,
Poland; [email protected]
received on 8 June 2006
Abstract. The structure and dynamics of bat assemblage in winter and transitional period was studied in
the Wisłoujście Fortress (16th–19th century) located in Gdańsk city, northern Poland. The object was used
partially as a commercial store (until 1995), later as a summer tourist attraction, protected in winter from
human penetration. Serious threats for existance of this monument forced recently the local officials to the
intensive renovation project, leading to deterioration of roosting conditions for bats. In total, nine species
were recorded: Myotis myotis, M. nattereri, M. mystacinus, M. brandtii (rare in Pomerania region), M.
da­s­yc­ne­me (EN in Poland), M. daubentonii, Eptesicus serotinus, Pipistrellus nathusii (first winter record
in Poland) and Plecotus auritus. M. daubentonii predominated among bats counted in autumn (43.4%)
and netted in the entrances to the casemates (60%). M. nattereri was the most numerous species during
winter (67.3%) and spring censuses (71.0%). M. dasycneme composed 3.5% of all bats counted inside of
the Fortress (n=2256 records) but about 18% of individuals captured in mist nets (n=67). Only 8–12 bats
were counted during winter 1993/1994. The number of bats increased significantly (r=0.86, p<0.02) until
2005, reaching the total number of 313 individuals in the whole object; however this trend was severely
interrupted by restoration works. The adaptation of formerly unprotected roost in the Fortress area (installation of bat grill, walls of air-bricks etc.) compensated this decline only in a limited way. In winter
2002/2003 we studied seasonal dynamics in bats’ numbers. The highest number of M. daubentonii and M.
dasycneme was re­cor­ded in September. The number of M. nattereri, after a peak in a half of September,
declined almost to zero in October and increased again, reaching its maximum value in February. Natura
2000 site was established in the Wisłoujście Fortress, although its future remains un­cer­tain.
INTRODUCTION
Several species of vespertilionid bats of the temperate zone spend winter in underground roosts
maintaining optimal thermal conditions and humidity (Althringam 1996). Such ro­osts are mainly
natural caves, widely distributed in mountain and upland areas. However, al­most no shelters of
that kind are available in the lowlands of Central Europe, thus many spe­ci­es need to hibernate
79
in their anthropogenic substitutes: cellars (Lesiński et al. 2004), wells (Bernard et al. 1998)
and – among the most important – various military structures, often considered as historical
mo­nu­ments (Bogdanowicz 1983, Urbańczyk 1990, Bernard et al. 1991, Fuszara et al. 1996,
Hebda & Nowak 2002, Sachanowicz & Zub 2002). The latter ones may serve as crucial sites
for ta­xo­no­mi­cal­ly rich bat assemblages, consisting of unu­su­al­ly large numbers of individuals
(e.g. Ur­bańc­zyk 1990). Forts, castles, air-raid shelters, bunkers and tunnels of underground
factories make an oportunity for long-therm monitoring of bat populations (Urbańczyk 1989),
similar to that conducted in caves and abandoned mi­nes of upland areas (for comparision see:
Fig. 1. Plan of the Wisłoujście Fortress. Explanations: I – the guard room of the postern tunnel, II – The
Południowo-Wschodni (South-Eastern) Bastion, III – The Ostroróg Bastion, IV – The Artyleryjski (Ar­til­le­
ry) Bastion, V – The Furta Wodna Bastion, VI – The Wreath (Arabic numbers refers to the controlled cellars),
VII – the powder-magazine of The Eastern Entrenchment. Objects I–VI are parts of the Fort Carré.
Rys. 1. Plan Twierdzy Wisłoujście. Objaśnienia: I – wartownia poterny, II – Bastion Południowo-Wschod­ni,
III – Bastion Ostroróg, IV – Bastion Artyleryjski, V – Bastion Furta Wodna, VI –Wieniec (cyfry arabskie
odnoszą się do kontrolowanych piwnic), VII – prochownia Szańca Wschodniego. Obiekty I–VI stanowią
część Fortu Carré.
80
Kowalski & Lesiński 1991, Weinreich & Oude Voshaar 1992, Řehák & Gaisler 1999). Although
selection of roosts by some bat species in military objects is well known recently (Bogdanowicz
& Urbańczyk 1983, Lesiński 1986), the factors affecting their population dynamics remained
unex­plai­ned, even if patterns of this dynamics were described in many localities (Bagrowska­
-Ur­bańc­zyk & Urbańczyk 1983, Fuszara & Kowalski 1995, Fuszara et al. 1996). Several
con­struc­ti­ons have stopped to ser­ve for military purposes just in the recent times, they became
utilized commercially, aban­do­ned or demolished; restoration works are started in some of them
in order to save their his­to­ri­cal values, however causing serious threats for wintering bats (e.g.
Mi­t­chell-Jones 2004). In contrast, unguarded sites are often under a serious pressure of un­con­
trol­led human pe­ne­tra­ti­on, intensifying spontaneous bat arousals (Thomas 1995) or even acts
of vandalism (Mi­t­chell-Jones 2004). Thus, intensity of human disturbance in bat hibernacula
may vary in spa­ce and time, however no case studies were published to document any effects
of this factor on bat numbers.
The Polish Baltic Sea Coast is the area of special biogeographic significance. It is known not
only as an important bat migratory path (Jarzembowski 2003), but also as a northern limit of
distribution for some European species (Baagøe 2001, Güttinger et al. 2001, Bog­da­nowicz
& Ruprecht 2004). However, the state of knowledge about distribution and size of winter
bat colonies in that region remains insufficient, restricted only to a small portion of available
un­der­g­round constructions (Jarzembowski et al. 2000, Wojtaszyn et al. 2001, Dzię­gi­e­lewska
2002). No detailed informations were published regardind bat hibernation in one of the most
valuable monuments of military architecture – 16th-century Wisłoujście Fortress (Gdańsk City).
Scattered data from transitional periods revealed significance of the object for some species
recognized as endangered in Poland (Myotis dasycneme – Ciechanowski & Przesmycka 2001)
or extremely rare in the region (Myotis brandtii – Ciechanowski & Sa­cha­nowicz 2003). Single
winter records of only four species have been published until re­cent­ly (Jarzembowski et al.
2000, Ciechanowski & Przesmycka 2001, Ciechanowski & Ko­ku­rewicz 2004).
The aim of this study was to describe the structure of bat assemblage using the Wisłoujście
Fortress during winter and transitional period, as well as long-term and seasonal changes in the
number of bats roosting in the monumental military object. The additional goal of the following
paper was to document the effect of temporally varying human disturbance on the largest bat
hibernaculum of Polish Baltic Sea Cost.
Study area and history of the site
The Wisłoujście Fortress is located in the Gdańsk city, northern Poland (18° 40’ 43” E, 54° 23’ 39” N).
Geographically, it lies in the eastern part of Baltic Sea Coast region – a border between Vistula Spit and
Żuławy Wiślane (delta of the Vistula river). The main object – Fort Carré – was built in the 16th century,
as a sea-fortress protecting the port of Gdańsk. It is composed of four Bastions: Południowo-Wschodni
(South-Eastern), Ostroróg, Artyleryjski (Artillery) and Furta Wodna. They contain large casemates, built
of bricks and covered by earth embankements. The lateral walls of the bastions are built of bricks and
stones as well. The fortress was built almost on the sea level, thus neither the casamates nor most of the
remaining bat hibernacula are the typical underground roosts, however the thick, brick walls made them
well insulated, at least before the main renovation works in 2004–2005. Originally, the casemates were
opened on the lateral sides, however the secondary walls were built in these openings in the 20th century,
making the insides dark and partially free of frost; the effect was increased when wooden doors were
installed in the main casemate gates. The remaining bat hibernacula of the Fort Carré are small cellars
under the officers’ houses in so called Wreath and small guard-room in the fort’s postern. Outside of the
central fort, the belt of external earthwork fortifications (The Eastern Entrenchment) was built in the 17th
81
century, enriched by the 19th century powder-magazine. The latter object, containing the narrow corridor
with some ventilation shafts in the ceiling, maintain a significant bat hibernaculum as well. In February,
the casemates and cellars were relatively cool (mean=3.7 °C, SD=2.2, range 0.6–9.0 °C, n=33) and moderately humid (mean=66.9%, SD=10.4, range 39.0–81.0%, n=19), although these figures may poorly
reflect conditions inside of the crevices or ventilation shafts. Mean external temperature in the same period
amounted 2.8 °C (SD=4.3, range –7.0–8.0 °C, n=8) (authors’ unpublished data).
Both Fort Carré and The Eastern Entrenchment are surrounded by wide moats, filled by brackish waters
from neighbouring Martwa Wisła – the dead branch of the Vistula River. Vegetation on The Eastern En­
tren­chment consists of shrubs, as well as unmanaged tree stand of maples Acer platanoides, old, partially
dying black poplars Populus nigra and lane of monumental horse-chestnuts Aesculus hip­po­casta­nus. The
earthworks of Fort Carré were densly overgrown by Lycium halimifolium, removed com­ple­te­ly in 2003. The
moat banks are covered by scattered beds of Phragmites australis and Schoenoplectus tan­baer­mon­ta­ni.
The Wisłoujście Fortress was demilitarised after the First World War and heavily demolished in 1945.
After a partial restoration it was used extensively as a store of paints, windows and vegetable preserves.
The vertical ventilation shafts were closed by concrete plates or even fulfilled with soil and rubble what
increased an insulation of internal parts of the casemates. The monument faced an increasing devastation
as an effect of sea waves, penetration of walls by moat waters, precipitation of salt and air-polution by
neighbouring “Siarkopol” company, which stored and processed large amounts of sulphur. Deep crevices
appeared in the walls and ceilings of all casemates of Fort Carré, causing a serious threat to the buliding,
although – both with loose fragments of plaster – maintaining crucial bat shelters during hibernation. After
1995, the Fortress stopped to be used as a commercial store. The monument, managed by The City of
Gdańsk History Museum, became utilised only in summer, as a seasonal tourist attraction. The Fort Carré
remained closed and guarded in winter, contrary to the powder magazine of the Eastern Entrenchment,
facing uncontrolled human penetration and disturbance.
The Fortress was included on the list 100 most endangered cultural monuments by World Monument
Watch in 2000–2001; the situation of the site resulted in starting of wide-scale restoration projects. First
renovation works were performed since September 2003, in the same period, when bats used to aggregate
in winter roosts. They included archeological excavations, prospecting drillings and removal of earth em­
ban­ke­ments in Bastion Artyleryjski, as well as temporary opening of ventilation shafts in all bastions (the
latter was performed in October). In the same period the Fortress became recognized as an important bat
site by Polish Ministry of Environment (established as Special Area of Protection PLH 220037 in Natura
2000 European network). All works were stopped during the winter 2003/2004, while the ven­ti­la­ti­on
shafts were closed provisionally (with spreadsheets of plastic foil) after an intervention of en­vi­ron­men­tal
officials. Restoration started again in spring 2004, resulting in fulfillment of all crevices in internal walls
of Bastion Artyleryjski with cement; in summer 2005 they became completely covered with de­sa­li­na­ti­on
compress. The small room in its casemates was separated from their remaining parts with a provisional
synthetic curtain (only in winter 2004/2005). Additionally, a secondary wall in South-Eastern Bastion
was partially removed in 2005, resulting in appearance of ice-cover in one of the corridors. All works
were stopped for the two last winter periods. As a partial compensation for negative results of renovation
works, the powder-magazine of the Eastern En­tren­chment was modified in order to increase its capacity
as bat hibernaculum. Its entrance was closed with bat grills (Mitchell-Jones 2004), while walls of air
bricks and water reservoir were built inside in summer 2005.
Material and methods
All four bastions of the Fort Carré and cellar no. 12 of the Wreath were checked every February between
2000 and 2006 (7 checks). In winter season 2002/2003 these objects were checked every month (since
September until April), in order to reveal seasonal dynamics in the number of bats (9 checks). Additional
objects (cellars no. 2 and 6, guard-room of the postern tunnel) were checked irregularly until 2005, when
all potential bat hibernacula in the Fortress area were controlled for the first time. Fort Carré was controlled
also 5 times in winter 1993/1994 (December–March), on 27 November 1999, three times in the autumn
82
2003 (29 October, 18 September, 19 December) and on 8 April 2006. The powder-magazine of the Eas­
tern En­tren­chment has been checked during every visit since September 2003. In total, 28 controls of the
Fortress were conducted between 1993 and 2006.
During each visit all visible bats were counted in the controlled objects, including those hidden in deep
crevices and ventilation shafts. Potential shelters were checked using torches and head-lamps. We avoided
handling of bats, except for species needing a careful examination of dental features or taking some me­a­
sur­ments (Pipistrellus spp., Myotis mystacinus, Myotis brandtii).
Five times (6 October 2000, 21 October 2000, 14 September 2002, 14 September 2003, 15 October 2006)
a mist netting was performed at the entrances of bastions of the Fort Carré. Two or four ECOTONE mist
nets were set up since dawn until midnight. Captured bats were determined to the species, sexed, aged (in
the case of M. daubentonii – Richardson 1994), weighed and their forearms measured prior to release.
Results
In total, nine bat species were recorded in autumn-spring period: Large mouse-eared bat Myo­
tis myotis (Borkhausen, 1797), Natterer’s bat Myotis nattereri (Kuhl, 1817), Whiskered bat
Myotis mystacinus (Kuhl, 1817), Brandt’s bat Myotis brandtii (Eversmann, 1845), Pond bat
Myotis dasycneme (Boie, 1825), Daubenton’s bat Myotis daubentonii (Kuhl, 1817), se­ro­ti­ne
Eptesicus serotinus (Schreber, 1774), Nathusius’ pipistrelle Pipistrellus nathusii (Key­ser­ling
et Blasius, 1839) and brown long-eared bat Plecotus auritus (Linnaeus, 1758). Among them,
M. nattereri was the most numerous, with a significant share of M. daubentonii (Table 1). Both
species were found during almost every control since the beginning of the study. M. dasyc­
neme appeared to be the third most numerous species, amounting not more than 5.2% of all
bats, but regularly encountered since its first winter observation on 11 February 2002. Single
M. myotis appeared irregularly (10 February 2001, 17 April 2003) until 2004, when it became
a permanent element of bat assemblage wintering in Fort Carré. The remaining spe­ci­es were
noted only sporadically: P. auritus (29 October 2003, 19 December 2003, 24 Febru­a­ry 2004,
18 February 2005), E. serotinus (21 November 2002, 21 January 2003, 18 February 2005), M.
mystacinus (1 male, 21 November 2002: Bastion Ostroróg), M. brandtii (1 fe­ma­le, 17 April
Table 1. The number and percentage of bat records from Wisłoujście Fortress. Autumn: 14 IX – 20 XII,
winter: 21 XII – 20 III, spring: 21 III – 17 IV
Tab. 1. Liczba stwierdzeń i udział procentowy gatunków nietoperzy w Twierdzy Wisłoujście. Jesień (au­
tumn): 14 IX – 20 XII, zima (winter): 21 XII – 20 III, wiosna (spring): 21 III – 17 IV
species
autumn
n
%
winter
n
%
spring
n
%
n
total
%
Myotis myo­tis
Myotis nat­te­re­ri
Myotis mys­ta­ci­nus
Myotis brand­tii
Myotis da­s­yc­ne­me
Myotis dau­ben­to­nii
Eptesicus se­ro­ti­nus
Pipistrellus na­thu­sii
Plecotus au­ri­tus
indeterminate
2
225
1
–
31
260
1
–
1
78
0.3
37.6
0.2
0.0
5.2
43.4
0.2
0.0
0.2
13.0
10
927
–
–
41
239
2
1
2
156
0.7
67.3
0.0
0.0
3.0
17.3
0.1
0.1
0.1
11.3
4
198
–
1
8
50
–
–
–
18
1.4
71.0
0.0
0.4
2.9
17.9
0.0
0.0
0.0
6.5
16
1350
1
1
80
549
3
1
3
252
0.7
59.8
<0.1
<0.1
3.5
24.3
0.1
<0.1
0.1
11.2
total
599
100.0
1378
100.0
279
100.0
2256
100.0
83
Table 2. The number and species composition of bats hibernating in Wisłoujście Fortress, counted on
18.02.2005. Explanations: I – the guard room of the postern tunnel, II – The Południowo–Wschodni (South-Eastern) Bastion, III – The Ostroróg Bastion, IV – The Artyleryjski (Artillery) Bastion, V – The Furta
Wodna Bastion, VI – cellars of the Wreath, VII – the powder-magazine of The Eastern Entrenchment
Tab. 2. Liczebność i skład gatunkowy nietoperzy zimujących w Twierdzy Wisłoujście dnia 18.02.2005. Ob­
jaś­ni­e­nia: I – wartownia poterny, II – Bastion Południowo-Wschodni, III – Bastion Ostroróg, IV – Bastion
Ar­ty­le­ryj­ski, V – Bastion Furta Wodna, VI – piwnice Wieńca, VII – prochownia Szańca Wschod­nie­go
species
I
II
III
IV
V
12
VI
2
6
VII
total
%
1.3
78.9
3.5
9.6
0.3
0.3
6.1
M. myo­tis
–
M. nat­te­re­ri
3
M. da­s­yc­ne­me
–
M. dau­ben­to­nii
–
E. se­ro­ti­nus
–
P. au­ri­tus
–
indeterminate
2
53
10
11
1
1
4
–
8
–
2
–
–
4
–
143
–
2
–
–
3
–
13
–
4
–
–
1
–
3
–
–
–
–
1
–
–
2
4
–
–
–
–
–
–
–
–
1
2
18
1
11
–
–
5
4
247
11
30
1
1
19
total
number of species
82
6
14
2
148
2
18
2
4
1
3
1
37
4
313 100.0
6
–
3
1
4
1
2003: Bastion Artyleryjski) and P. na­thu­sii (1 male, 16 February 2003: Bastion Furta Wodna).
In total 2256 bat records were noted (Table 1) and 313 bats were counted maximally in the
whole Fortress (Table 2).
The species composition varied in time. In autumn, M. daubentonii predominated, but in winter
and spring M. nattereri was a dominating taxon. The proportion of M. daubentonii compared
to M. nattereri was significantly higher in autumn than in winter (Chi-square test with Yates’
correction, χ2=176.50, p<0.0001), in autumn than in spring (χ2=73.85, p<0.0001), but did not
differ between winter and spring (χ2=0.00, p>0.9).
The total number of hibernating bats in the Fortress has changed significantly since 1994.
After closing of the commercial stores it began to increase progressively (r=0.86, p<0.02; y=
–41415.7626 + 20.7553957x) until 2005, when it reach the maximum. This trend was interrupted
only in winter 2003/2004, when first renovation works, excavations and pro­spe­cting drillings
were performed in autumn. However, a year later, the number of bats roosting in the Fort Carré
dropped about 77% in association with removal of secondary wall in South-Eastern Bastion and
covering of the walls with desalination compress in Artillery Bastion. Although the number of
bats counted in newly adapted powder-magazine of the Eastern En­tren­chment increased almost
twice, it did not compensate a population decline in the re­ma­i­ning objects.
Hibernating bats were unevenly distributed in the Fortress area, with 85% of the as­sem­bla­ge
concentrated in just three objects. The highest species richness was observed in South-Eastern
Bastion and powder-magazine of the Eastern Entrenchment, while the highest number of in­di­vi­
du­als appeared in the Artillery Bastion. Only a few bats were noted in the remaining six objects
(Table 2). With respect to the species composition, the first two objects appeared to be the most
vaulable bat hibernacula, where most individuals of M. dasycneme spend win­ter and mate in
autumn (observations of copulating pond bats in 2002). Progressing re­s­to­ra­ti­on works strongly
affected the spatial distribution of bats. After their first stage in 2003, performed in the autumn,
bats number decreased more than one third in the Artillery Bastion, but increased in the South84
-Eastern Bastion (Fig. 3). However, the fulfillment of crevices in the Artillery Bastion did not
cause any decline of bats number in the next winter season (2005). Moreover, they aggregated
almost all in one place – a small chamber, separated by a plastic curtain, where three, large
clusters of M. nattereri (59, 51 and 17 individuals) hanged directly on its walls and composing
a majority (88%) of all bats wintering in this bastion. Such a behaviour was formerly unknown
in the Fortress, where most of the animals hibernated hid­den in ceiling cracks, dispersed among
se­ve­ral, smaller clusters. Only after the second stage of restoration works, bats almost completely
abandoned the Artillery Bastion, where not only crevices were filled, but also the temporary
curtain was removed (Fig. 4).
The number of bats varied strongly through the season as well (Fig. 5). The largest ag­gre­ga­ti­
ons of M. daubentonii appeared in early September. Contrary to the winter period (see above),
they did not hide in the crevices, but covered the ceilings and walls of ventilation shafts in dense
clusters. The number of M. daubentonii declined visibly in the third decade of September, later
increasing again, but not reaching the earlier level. It remained relatively stable between October
Fig. 2. Long-therm changes in the number of bats hibernating in the Wisłoujście Fortress in 1994–2006,
based on February counts. Explanations: 1. Autumn works in Artillery Bastion (prospecting drillings,
archeological excavations, opening of ventilation shafts, removal of earth embankements); 2. Summer
renovation works (fulfillment of crevices and laying of desalination compress on walls in Artillery Bastion,
partial removal of secondary wall in South-Eastern Bastion).
Rys. 2. Długoterminowe zmiany liczebności nietoperzy zimujących w Twierdzy Wisłoujście (lata 1994–
2006), w oparciu o liczenia w lutym. Objaśnienia: 1. Jesienne prace w Bastionie Artyleryjskim (wiercenia
roz­po­znawc­ze, wykopaliska archeologiczne, udrożnienie przewodów wentylacyjnych, usunięcie na­sy­pów
zi­em­nych znad sklepienia); 2. Letnie prace remontowe (wypełnienie szczelin i pokrycie kompresem odsalającym wewnętrznych ścian Bastionu Artyleryjskiego, częściowe usunięcie wtórnego zamurowania
w Bastionie Południowo-Wschodnim).
85
Fig. 3. Effect of works conducted in Artillery Bastion in autumn 2003 on bats number in particular parts
of Fort Carré, expressed as a differrence in bat number between December 2002 and 2003.
Rys. 3. Wpływ prac prowadzonych w Bastionie Artyleryjskim jesienią 2003 na liczebność nietoperzy
w poszczególnych częściach Fortu Carré, wyrażony jako różnica w liczbie osobników między grudniem
2002 i 2003.
Fig. 4. Effect of works conducted in Artillery and South-Eastern Bastions in 2005 on bats number in
par­ti­cu­lar parts of Wisłoujście Fortress, expressed as a differrence in bat number between February 2005
and 2006.
Rys. 4. Wpływ prac prowadzonych w Bastionach Artyleryjskim i Południowo-Wschodnim w roku 2005
na liczebność nietoperzy w poszczególnych częściach Twierdzy Wisłoujście, wyrażony jako różnica
w liczbie osobników między lutym 2005 i 2006.
86
Table 3. The species and sex composition of bats netted in autumn in the entrances of casemates and pow­
der-magazine of the Wisłoujście Fortress
Tab. 3. Skład gatunkowy i struktura płciowa nietoperzy odławianych w sieci jesienią przy wylotach z ka­
za­mat i prochowni Twierdzy Wisłoujście
species
males
females
total
%
Myotis myotis
Myotis nattereri
Myotis mystacinus
Myotis dasycneme
Myotis dau­ben­to­nii
1
8
1
10
24
2
3
–
2
16
3
11
1
12
40
4
16
1
18
60
total
43
22
67
100
and February, decreasing finally in spring. A similar seasonal dy­na­mics was revealed by M.
dasycneme, however basing on a very small sample (n=39 records). The number of M. nattereri
– after an early peak in the beginning of season – dropped almost to zero in October. However,
it increased about 60 times until January, exceeding the number of M. daubentonii already in
the late autumn. The highest number of M. nattereri was re­cor­ded in March (declining almost
six times in the next month), but the highest total number of bats – in February.
Fig. 5. Seasonal dynamics in the number of bats roosting in Wisłoujście Fortress in winter 2002/2003.
Rys. 5. Sezonowa dynamika liczebności nietoperzy wykorzystujących Twierdzę Wisłoujście jako kry­
jówkę zimą 2002/2003.
87
Among bats netted in the autumn (n=67), M. daubentonii predominated, but the high per­
cen­ta­ge of M. dasycneme (18%) was noticeable (Table 3). Most captured individuals were
males, however a sex ratio did not differ significantly from 1:1 (χ2=2.74, p>0.09). Among M.
daubentonii 31 juveniles (yearlings) and 9 adults were captured; difference from 1:1 age ratio
was statistically significant (χ2=5.41, p=0.02). When the mist netting was conducted, several
individuals of small Myotis were observed flying inside of casemates.
Discussion
The number of bats roosting in the Wisłoujście Fortress during winter period placed it among
the largest bat hibernacula in the Polish Baltic Sea Coast, at least in some seasons. Only one
site – ruins of amunition factory in Police near Szczecin – appears larger, accumulating more
than 1300 hibernating individuals (K. Drabińska, J. Żejmo, M. Dzięgielewska in Kepel et
al. 2005). About 200 individuals were counted maximally in the castle of Teutonic Knights
in Malbork (Stec & Kasprzyk in Kepel et al. 2005) and 120–150 individuals in bunkers of
Kołobrzeg-Stadion and Szczecin-Zdroje (Wojtaszyn et al. 2001, Dzięgielewska 2002). No
other bat hibernaculum located in Gdańsk city has ever been used by more than 90 animals
(Jarzembowski et al. 2000).
Among the species recorded in Wisłoujście, some of them are of special significance, even
if only exceptionaly encountered. M. brandtii, although considered as widely distributed in Po­
land, is extremely rare in the northern part of the country. It is only the second known locality
of this species in the Baltic Sea Coast region (Ciechanowski & Sachanowicz 2003). P. nathusii
is a long-distance migrant, known to leave Central Europe for winter and hi­ber­na­ting in hollow
trees or houses located in milder climate of western Europe (Strelkov 1969). The Wisłoujście
Fortress is the first known winter site of P. nathusii in Poland and quite an unusual type of
hibernaculum, although the recent winter record of the species from Czech Republic is also
associated with a similar object (a castle cellar, Benda & Hotový 2004). Among the regular
inhabitants of the Fortress, M. dasycneme should be considered as the most interesting species.
Regarded as globally vulnerable (Hutson et al. 2001) and nationally endangered (Wołoszyn
2001), it is in­clu­ded in Annex II of EU Habitat Directive, justifying designation of Natura
2000 site in Wisłou­jś­cie Fortress. Polish hibernacula do not con­cen­tra­te large numbers of M.
dasycneme. Its share in winter bat assemblages usually do not ex­ce­ed 0.5% (Bagrowska-Ur­
bańczyk & Urbańczyk 1983, Lesiński 1986, Kowalski & Lesiński 1991, Fuszara et al. 1996)
and even in some large brick forts no pond bats are encountered (Bogdanowicz 1983, Fuszara
& Kowalski 1995, Hebda & Nowak 2002). Wisłoujście Fortress is one of the largest winter sites
of M. dasycneme in Poland, along with Fort Osowiec in Biebrza River Valley (at maximum
34 individuals – Lesiński & Kowalski 2002) and four other localities (including three fortification
complexes), each con­cen­tra­ting no more than 6–15 individuals (Ciechanowski & Kokurewicz
2004, T. Kokurewicz in litt.). In fact, the real number of pond bats roosting in the Fortress can
be even higher, what may be presumed from regular captures of M. dasycneme in autumn and
observations of some in­di­vi­du­als hiding deeply in narrow crevices. Bogdanowicz (1983) stated that M. dasycneme avo­i­ded winter quarters located in heavily urbanized areas, thus being
a bioindicator of the chan­ges in a habitat subjected to urbanization pressure. This hypothesis
can recently be re­jec­ted, as the Wisłoujście Fortress – intensively used by this species both in
autumn and winter – is located on the edge of densely populated city, where two large chemical
factories have pol­lu­ted the neighbouring area since 1969.
88
The species composition of bats wintering in Wisłoujście Fortress differed from that re­cor­ded
in the other large, brick forts of Poland. In central and northeastern part of the country, forts are
inhabited mainly by a barbastelle Barbastella barbastellus (Schreber, 1774) com­po­sing 50.8%
of all bats (Fuszara et al. 1996). Also in the forts of Nysa (Lower Silesia) barbastelle is the most
numerous species (Hebda & Nowak 2002), while in the forts of Po­znań it composed at least
14.1–20.9% of winter bat colonies (Bogdanowicz 1983). Bar­bas­tel­le is an oligothermophilous
species, hibernating in 0.0–3.0 °C (Bogdanowicz & Ur­bańc­zyk 1983) and large forts are considered rather cool roosts with intensive air circulation (due to many entrances and windows),
what explains high numbers of B. barbastellus spending winters there. Thus, the lack of any
bar­bas­tel­les hibernating in Wisłoujście Fortress seems astonishing, however the species does
not occur anywhere in Gdańsk (Jarzembowski et al. 2000) and it is a very rare bat in the whole
Pomerania region (Kowalski & Szkudlarek 2003). Instead, the winter bat assemblage of the
studied locality is dominated by moderately ther­mo­phi­lous species, i.e. M. nattereri and M.
daubentonii. However, they are known to find shel­ters in the deep, highly located crevices and
ventilation shafts (Bogdanowicz & Urbańczyk 1983, Lesiński 1986), where the temperature
may be significantly higher and more stable than in the corridors themsleves.
Low number of bats in the period, when object was intensively used as a commercial store,
confirms their sensibility to repetitive disturbance (light, noise) during hibernation (Thomas
1995). Later increase in bat numbers appeared similar to that observed in roosts newly ope­ned
for these animals, e.g. the old mines, where exploitation came to an end (Lutsar et al. 2000,
Bihari 1998 in Güttinger et al. 2001). Also in winter quarters, where bats were in­ten­si­ve­ly ban­
ded, their numbers increased several times after the end of marking programm (Le­siński 1990,
Řehák & Gaisler 1999). Spatio-temporal dynamics of bat numbers in the Wisłou­jś­cie Fortress
suggest however, that human pressure restricted only to the part of fortification complex forced
at least some bats to change roost for the nearest available, not to leave completely the area.
Movements among hibernacula in one and the same winter season (ap­pa­rent­ly associated with
flying outside) were reported during some banding programmes (Bog­da­nowicz & Urbańczyk
1983, Gaisler et al. 2003). Bats may change their shelter not only directly due to the human
activity, but its subsequent effects on internal conditions. At least some elements of restoration
works (removal of secondary wall, opening of ventilation shafts) might strongly affect micro­c­
li­ma­te in the casemates, causing bats not only hiding deeper in the crevices (reaction observed
frequently in M. daubentonii when ambient temperature de­cre­a­sed – Kokurewicz 2004), but
also moving into unaffected parts of the Fortress. The latter behaviour can be compared with
that recorded in B. barbastellus wintering in small, concrete bunkers and regularly moving
from the upper floor to the dried wells in the coldest months (Sachanowicz & Zub 2002). Our
ob­ser­va­ti­ons showed however, that alternative roosts could only partially compensate the effect
of human pressure on hibernaculum, likely due to un­pre­dictable character of this factor.
The phenology of bat occurrence in Fortress resembles that observed in the other for­ti­fi­ca­ti­
ons of lowland Poland. M. daubentonii is a bat appearing earlier in the undergrounds that M.
nattereri and the highest numbers of this species are observed in the autumn, decreasing later
during winter months (Bagrowska-Urbańczyk & Urbańczyk 1983, Jurczyszyn 1998). Howe­ver,
the mentioned peak in the number of Daubenton’s bats was reported a bit later – from October
(Lesiński 1986, Fuszara & Kowalski 1995, Fuszara et al. 1996) or even from No­vem­ber (Jur­
c­zys­zyn 1998). Thus, the mass occurrence of M. daubentonii in the Fortress in September is
worth noticing, possibly representing a short-therm phenomenon shifting in time among years
– we observed it on 14 September 2002 but not on 18 September 2003. The species composition
89
of bats netted in autumn resembled that recorded inside of the ca­se­ma­tes, although it may not
represent the latter ones (they remained in a deep torpor) but the animals visiting the Fortress
in order to take part in a swarming behaviour (sensu Parsons et al. 2003). Several individuals
of M. daubentonii visited the Modlin Fortress near Warsaw even in August, however they
number dropped almost to zero in the first week of September and increased again. Most of the
in­di­vi­du­als appearing then in Modlin remained active, flying inside of the forts, and only 4%
of bats ringed in the autumn (including those found in le­thar­gy) stayed there in winter (Lesiński
1990). Thus it was possible that high numbers of M. dau­ben­to­nii roosting in the Fortress in
September did not refer to the animals counted later in mid-winter but to the bats using the site
as a transitional shelter during autumn movements.
Contrary to M. daubentonii, the maximum numbers of M. nattereri in hibernacula are usu­al­ly
observed in mid-winter, mostly in January–February (Bagrowska-Urbańczyk & Ur­bańc­zyk
1983, Fuszara et al. 1996), sometimes in March (Fuszara & Kowalski 1995) or De­cem­ber
(Jur­c­zys­zyn 1998). The earlier results are, thus, consistent with our observations, however an
early peak in the number of Natterer’s bats, revealed in the half of September, was recorded
for the first time. These animals could also use the Fortress as a transitional roost, leaving it
after a short time, what may be judged from the very low numbers of M. nattereri in October,
confirmed also in the other sites (Lesiński 1986, Fuszara et al. 1996, Jurczyszyn 1998). There
are almost no earlier studies to compare with our data on phenology of M. dasycneme. The two
series of censuses conducted by Le­siński & Kowalski (2002) in northeastern Poland revealed
the numbers of pond bats at least 2–10 times higher in November–December than in February.
No such phenomenon was observed in Wisłoujście and the September peak in the number of
M. dasycneme seems to reveal the same aspect of annual life cycle as in M. nat­te­re­ri and M.
daubentonii. It should be noted however, that some changes in the number of bats counted in
underground hibernacula could be an artifact, unrelated to arrival or departure from roosts, but
caused by movements of some in­di­vi­du­als into the deep crevices (Jurczyszyn 1998). This effect
in M. daubentonii was explained as a reaction to unfavorable thermal con­di­ti­ons (Kokurewicz
2004).
The future of the Wisłoujście Fortress as an important bat site remains uncertain. It would be
hard to preserve such a large winter colony in the casemates of Fort Carré, because the pro­spe­
cting restoration works – necessary to preserve the Fortress itself – will change the micro­c­li­
ma­te, as well as the number and quality of micro-shelters. Adaptation of the powder-magazine
on Eastern Entrenchment may compensate these losses only in a limited way and there would
be a need for a significant delay (about 2–4 years) in renovation of the South-Eastern Bastion,
until more bats accept a new roost. The main purpose for the creation of Natura 2000 site on the
Fortress’ area – regular occurrence of M. dasycneme – seems possi­ble to be maintained, as the
powder-magazine became recently the most important shelter of this species (5 of 7 individuals
were noticed there on 8 February 2006).
Streszczenie
W latach 1994–2006 badaliśmy strukturę i dynamikę zgrupowania nietoperzy wykorzystujących Twierd­zę
Wisłoujście (Gdańsk, północna Polska) zimą i w okresach przejściowych. Obiekt, budowany w XVI–XIX
wieku, do 1995 roku był częściowo użytkowany jako magazyny, w ostatnich latach zaś jako letnia atrakcja turystyczna, zabezpieczona przed ludzką penetracją w okresie zimowym. Poważne zagrożenia dla
substancji zabytkowej Twierdzy, skłonily lokalne władze do intensywnej renowacji obiektu, co pro­wad­zi
jednak do ob­niże­nia jego wartości jako kryjówki nietoperzy.
90
Łącznie na terenie Twierdzy stwierdzono 9 gatunków nietoperzy: nocek duży Myotis myotis, nocek
Nat­te­re­ra M. nattereri, nocek wąsatek M. mystacinus, nocek Brandta M. brandtii (rzadki na Pomorzu),
nocek łydkowłosy M. dasycneme (kategoria EN w Polsce), nocek rudy M. daubentonii, mroczek późny
Ep­te­si­cus serotinus, karlik większy Pipistrellus nathusii (pierwsze w Polsce stwierdzenie zimowe) i gacek
bru­nat­ny Plecotus auritus. Nocek rudy dominował wśród nietoperzy liczonych jesienią wewnątrz Twierd­
zy (43,4%) i odławianych w sieci przy wejściach do kazamat (60%). Nocek Natterera był najliczniejszym
gatunkiem podczas liczeń zimowych (67,3%) i jesiennych (71,0%). Nocek łydkowłosy stanowił 3,5%
ws­zyst­kich nietoperzy liczonych wewnątrz Twierdzy (n=2256 stwierdzeń), ale około 18% osobników
odławi­a­nych w sieci (n=67).
Zimą 1993/1994 na terenie Twierdzy znajdowano zaledwie 8–12 nietoperzy. Łączna liczba tych zwierząt
istotnie wzrastała do 2005 roku (r=0,86, p<0,02), osiągając wartość 313 osobników w całym obiekcie;
jednak trend ten został przerwany na skutek prac renowacyjnych. Adaptacja wcześniej niechronionej
Prochowni Szańca Wschodniego (instalacja kraty zabezpieczającej, ścian z cegły-dziurawki) tylko częś­
ci­owo skom­pen­so­wała spadek liczebności nietoperzy między latami 2005 i 2006.
Zimą 2002/2003 badaliśmy sezonową dynamikę liczebności nietoperzy. Najwięcej nocków rudych
i noc­ków łydkowłosych odnotowaliśmy we wrześniu. Liczba nocków Natterera, po osiągnięciu szczytu
w połowie września, spadła niemal do zera w październiku, po czym ponownie wzrosła, osiągając mak­
sy­mal­ną wartość w lutym. Twierdza Wisłoujście uznana została za Specjalny Obszar Ochrony w sieci
Natura 2000, jednak jego pr­zyszłość, jako jednego z największych zimowisk nietoperzy na Pomorzu
Gdańskim, po­zos­ta­je ni­e­pew­na.
Acknowledgements
Authors are cordially thankful to all who have contributed to the following study and conservation of bats
in the Wisłoujście Fortress. Members of The City of Gdańsk History Museum staff: Adam Koperkiewicz,
Halina Trzebiatowska and Zdzisław Balewski for making access to the study area possible and for their
decision to adapt a powder-magazine for bat hibernaculum. Members and co-workers of Academic Chi­ro­
p­te­ro­lo­gi­cal Circle of PTOP “Salamandra” in Gdańsk: Urszula Anikowska, Anna Biała, Piotr Chybowski,
Tomasz Jar­zem­bowski, Monika Kulas, Anna Miotk, Anna Nalewaja, Justyna Naumowicz, Anna Pawlik,
Katarzyna Pazio, Paulina Piasecka, Weronika Rogalska, Aleksandra Rynkiewicz, Katarzyna Woj­c­zu­la­nis,
Tomasz Za­jąc and Aneta Zapart for their help in the field studies.
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ISSN 0024–7774
The Early Miocene mammalian assemblages in Jebel Zelten, Libya
Spodnomiocénní společenstva savců z Džebelu Zelten, Libye
Oldřich Fejfar1 & Ivan Horáček2
Department of Geology and Palaeontology, Charles University, Albertov 6, CZ–128 43 Praha 2,
Czech Republic; [email protected]
2
Department of Zoology, Charles University, Viničná 7, CZ–128 44 Praha 2, Czech Republic;
[email protected]
1
received on 8 December 2006
Abstract. The small mammal assemblage obtained from six sites of the Miocene deposits in Jebel Zelten,
Libya, in 1982, 1983 and 1997, is composed of 10 species of rodents (seven different clades), 2 spp. of
ochotonid Lagomorphs and one vespertilionid bat. A special attention is given to the forms characterising
the uppermost part of the series: they belong (mostly as new species) to the genera Potwarmus (Muridae),
Mellalomys (Myocricetodontinae), Heterosminthus (Lophiocricetinae), Sayimys (Ctenodactylidae) and
a new genus of Vespertilionidae. The most parsimonious dating for them is the early Middle Miocene (ca.
15 My) as the cricetids and ctenodactylids are apparently less advanced than their congeneric forms in Beni
Mellal (14 My) while Potwarmus, representing the first wave of murid expansion from Asia to Africa, is
more advanced than the congeneric forms from Banda daud Shah in Pakistan (16 My). All these forms
reveal close relations both to the clades known from several other African localities and to those found
in the Early or Middle Miocene of Pakistan and illustrate the intesive s. c. “southern” faunal interchange
between north Africa and Asia in time of the Early and Middle Miocene.
Introduction
Small mammal remains from seven localities were collected during two geological and paleontological field campaigns (1982, 1983 and 1997). The assemblages are small, but the twelve
species recognized and described represent seven rodent families, one lagomorph and one bat
family. The Jebel Zelten large mammal fauna was considered in most literature to represent
one time-slice, although the inter­pretation of its age has been diverse. However, on basis of the
evolutionary stage of the small mammal species, the faunal compositions and the stratigraphic
sequence the Jebel Zelten assemblages represent three periods in time and cover approximately
4 million years. Three assemblages can be assigned to the Middle Early Miocene (18–19 Ma),
one to the Late Early Miocene (16–17) and two to the Middle Miocene (14–15 Ma) (Wessels
et al. 2003).
From sites with a high concentration of vertebrate remains at the sur­face, the fine cross-bedded (estuarine – fluviatile) sands were extensively dry sieved by O. Fejfar in 1982 and 1983.
On two sites, the “Measured Section 2” (MS 2) in the middle part of the escarpment and Wadi
Umm Ash Shatirat (WS) in the most southern part of the escarpment (Fig.1), isolated molars of
several taxa of rodents were collected. Site MS 2 corresponds with the vertebrate site “H – area
6409” and the site of Wadi Shatirat (WS) corresponds with the Vertebrate site “LP – areas 641295
16” of Savage & Hamilton (1973). Each assemblage is derived from a different stratigraphical
level, e.g., MS2 belongs to a stratigraphi­cally lower level than Wadi Shatirat.
A geological and paleontological campaign El Arnauti-Daams in 1997, resulted both in
collection of large mammals and further small collection of rodents and lagomorphs. In many
localities of the Jebel Zelten escarpment three to four fossilliferous units of sandsto­nes were
recognised. The lowermost fossiliferous unit consists of shallow channel deposits containing
rust-coulored sands, small clay lenses, reworked clay pebbles, remnants of bioturbation, wood
(stumps) and large mammal bones. The second unit is a channel depo­sit also, consisting of coarse
green sands and large bones. The third unit consists mostly of white (bleached) sands intersected
by small pebble layers. Bioturbation and large bones are common. The fourth unit is composed
of coarse sands with large bones. These units, however, are not continuous and the correlation
of the localities in dif­ferent sections is therefore mainly based on fossil content. Large mammal
remains were recoverd from many localities. After wet screening of sediment (with water from
an oil well) rodent and lagomorph remains were found in only five localities.
SYSTEMATIC LIST
Vespertilionidae Gray, 1821
Vespertilionidae gen. nov. et sp. nov.
Despite respresented by a single specimen only (found in the washed residuum of site MS2 by
Fejfar in 1983), the chiropteran record from Jebel Zelten is of a considerable significance. It is
a right mandible of a robust vespertilionid bat (CM3 alv. 7.2 mm) with well preserved myotodont
molars (M1 length 1.76 mm, width 1.06/1.20 mm, M2 length 1.85 mm, width 1.20/1.22 mm).
Preliminary we reported it as Scotophilus sp. (Wessels et al. 2003) as it corresponds to Recent
Table 1. Species list and sequence of localities, dry sieved in the field in 1983 at sites A, B, and washed
at the other sites in 1997 (from Wessels et al. 2002)
Tab. 1. Soupis druhů/forem a přehled lokalit, ze kterých byl materiál nasucho prosíván roku 1983 a vymýván roku 1997 (převzato z Wesselse et al. 2002)
Sites / lokality: 1 – ATH7A2 (2), 2 – ATH7A3 (2), 3 – ATH5A1 (1), 4 – ATH4B (3), 5 – QAB1C (4),
6 – MS2 (A), 7 – Wadi Shatirat (B)
(sub)family Cricetidae Cricetidae Murinae Myocricetodontinae Myocricetodontinae Rhizomyinae Lophocricetinae Ctenodactylidae Thryonomyidae Ochotonidae Ochotonidae Vespertilionidae 96
sites:
?Cricetidae gen. et sp. indet. gen. et sp. indet. Potwarmus sp. nov. Mellalomys sp. cf. Myocricetodon sp. Prokanisamys sp. Heterosminthus sp. indet. Sayimys nov. sp. gen. nov. et sp. nov. Alloptox sp. ?Kenyalagomys sp. gen. nov. et sp. nov. 1
2
3
4
5
6
7
–
–
–
–
–
+
–
–
+
–
–
–
–
–
–
–
–
–
–
–
+
–
+
–
–
+
–
–
–
+
–
–
–
–
–
–
–
–
–
+
+
+
–
+
+
–
–
–
–
–
–
+
–
–
–
–
–
–
–
–
+
–
+
+
–
–
+
+
–
+
–
+
–
–
+
–
–
–
–
+
–
–
–
–
Scotophilus kuhlii Leach, 1821 or S. viridis (Peters, 1852) in degree of reduction of unisuspid
teeth and robust construction of mandibular body and molars i.e. the characters by which it
differs from other extant genera like Eptesicus Rafinesque, 1820, Otonycteris Peters, 1859 or
Scotomanes Dobson, 1875. Of course, as demonstrated elsewhere (Horáček et al. 2006) it cannot
be directly coidentified with extant Scotophilus Leach, 1821 because of less advanced form of
molars and with the combination of these characters it does not respond to diagnostic criteria
of any other genus, both extanct and extinct, except for partial agreement with the Paleogene
African clade Philisidae Sigé, 1985 (Philisis sphingis Sigé, 1985 from Oligocen of Fayum,
Dizzya exultans Sigé, 1991 from early Eocene of Chambi, Tunisia, Phylisis sevketi Sigé, 1994
from Oligocene of Taqah, Oman) from which it differs by more advanced degree of unicuspid
reduction. Thus, it cannot be excluded that the Jebel Zelten bat represents a transitional stage
between Phylisinae and the extant genus Scotophilus. The latter genus is now distributed throughout Africa and Madagascar (with 14 spp.) and in southern Asia. The Asiatic Scotophilus kuhlii
is considered the most primitive both in morphological and molecular respects, its divergence
time from the African clades can be estimated (based on molecular clock) to Middle or Late
Miocene. For more details see Horáček et al. (2006).
Fig. 1. Geographical map of the middle part of northern Libya with localities of fossil mammals in the
Jebel Zelten escarpment (modified after Savage & Hamilton 1973).
Obr. 1. Geografická mapka střední části severní Libye s nalezišti fosilních savců v Džebelu Zelten (upraveno podle Savage & Hamiltona 1973).
Legend / legenda: sites / lokality 1983: A – Measured Section 2 / zaměřený profil 2 (MS2), B – Wadi Umm
Ash Shatirat (WS); sites / lokality 1997: 1 – ATH5A, 2 – ATH7A, 3 – ATH4B, 4 – QAB1C.
97
Murinae Gray, 1821
Potwarmus Linsay, 1988
Potwarmus sp. nov.
(Fig. 3: 1–4)
The material of Potwarmus sp. nov. was recorded at the sites and layers: 1. locality 1 (MS 2),
in lower part of the Qarat Jahannam member of Maradah Formation, middle part of the SW
Jebel Zelten escarpment, and 2. locality at Section of Wádí Shatírát.
The record of Potwarmus from Jebel Zelten is characterized with inflated median cusps (t5,
t8), absence of anterior and posterior mure, enterostyle (t4) at the level of t6, and long posterior
cingulum on M1. It shows some light differences in size and morphology of the cusps from
Potwarmus primitivus Wessels et al., 1982 (Chinji, Pakistan), P. thailandicus Jaeger et al., 1985
(Li, Thailand) and P. minimus Wessels et al., 1987 (Pakistan). Potwarmus sp. nov. from Jebel
Zelten seems to be more advanced than P. primitivus and P. thailandicus. Generally the molars
of the north African form are less bunodont than other species of Potwarmus which could represent a more advanced stage of evolution in comparison with the asiatic forms.
The occurrence of the genus Potwarmus in North Africa indicates a migra­tion of this genus
from southern Asia to Africa. Its migration route is unknown since primitive murines are not
known from Asia minor or the Arabian penin­sula. Potwarmus sp. nov. is slightly more evolved
than Potwarmus from Banda daud Shah in Pakistan (Wessels et al. 1982; dated ca. 16 Ma), excluding a migration during the Early Miocene times (18 Ma) (Wessels et al. 2003). Considering
the uncertainties of its relationship(s) to African muroid subfamilies, Potwarmus is regarded as
a primitive murid, as is Antemus Jacobs, 1978, although both genera lack the (for true murids)
characteristc chevrons of three cusps is the first upper molar.
Myocricetodontinae Lavocat, 1962
Mellalomys Jaeger, 1977
Mellalomys sp. (nov. ?)
(Fig. 3: 7–9)
The material from Jebel Zelten is clearly dis­tinct from Mellalomys lavocati Wessels, 1996 from
Sind, Pakistan, Lower-Middle Miocene and is except for the poorly divided anterocone, clearly
more evolved. The pre­sence of a double anterocone, the short ‘normal’ lon­gitudinal crest, an

Fig. 2. Representants of selected species of rodents from the Jebel Zelten sites MS2 and Wadi Shatirat.
Occlusal views of the molars, figured as left. 1–4 – Potwarmus sp. nov., M1 sin. (1); M1 (2, 3); M1 and M2
from one individual (4, inv.); 5, 6 – Heterosminthus sp. indet., M2 dext. (inv.); 7, 8 – Mellalomys sp. nov.;
7 – M1 dext. (inv.); 8 – M2 sin.; 9, 10 – ?Cricetidae gen. et sp. indet., 9 – M2 dext.; 10 – M2 dext.; 11–14
– Sayimys sp. nov., 1a – M2 dex; 1b – anterior view; 2a – M3 (inv.); 13, 14 – posterior views. Scale A to
the Figs. 1–10, scale B to the Figs. 11, 12, scale C to the Figs. 13, 14.
Obr. 2. Zástupci vybraných druhů hlodavců z nalezišť v Džebelu Zelten: MS2 a Wadi Shatirat. Okluzální
plochy molárů jsou vyobrazeny jako levé. 1–4 – Potwarmus sp. nov., M1 sin. (1); M1 (2, 3); M1 a M2
z jediného kusu (4, inv.); 5, 6 – Heterosminthus sp. indet., M2 dext. (inv.); 7, 8 – Mellalomys sp. nov.; 7
– M1 dext. (inv.); 8 – M2 sin.; 9, 10 – ?Cricetidae gen. et sp. indet., 9 – M2 dext.; 10 – M2 dext.; 11–14
– Sayimys sp. nov., 1a – M2 dex; 1b – pohled zpředu; 2a – M3 (inv.); 13, 14 – pohledy odzadu. Měřítko A
k obr. 1–10, měřítko B k obr. 11, 12, měřítko C k obr. 13, 14.
98
99
elongated anteroconid are charac­teristic for Mellalomys Jaeger, 1977. A short meso­lophid (M1)
is known to occur in primitive Mella­lomys species from Pakistan (Wessels 1996). Mellalomys
sp. (nov. ?) is smaller than Mellalomys atlasi (Lavocat, 1961), has lower cusps and ridges, the
anterocone less well divided, the anterior ledge on the anteroco­ne smaller (or absent) and the
longitudinal crest is not oblique. Mellalomys from Jebel Zelten could rep­resent a new species, related to and more primitive than Mellalomys atlasi from Beni Mellal (14 Ma; Jaeger 1977).
Dipodidae Fischer von Waltheim, 1817
Lophocricetinae Savinov, 1970
Heterosminthus Schaub, 1930
Heterosminthus sp. indet.
(Fig. 3: 5)
This specimen shows similarity with Heterosmint­hus which has four roots on the M1 and M2,
a pro­minent cusp on the postero-lingual edge of the proto­cone and lacks the lingual branch of
the anteroloph. It differs from Heterosminthus in lacking the lingual branch of the posteroloph
and having the metaloph connected to the posteroloph (Qiu 1996). It differs from the more
progressive genus Arabosminthus Whybrow et al., 1982 by its elongate shape, the less robust
cusps and anterior arm of the hypocone, the wide first labial syncline, the presence of a low
connection bet­ween protocone and paracone and the strong connec­tion between metacone
and posteroloph. Heterosminthus is known from the Late Oligoce­ne and the Miocene of Asia
(Daxner-Höck 2001).
Ctenodactylidae Gervais, 1853
Sayimys Wood, 1937
Sayimys sp. nov.
The upper molars are similar to Sayimys inter­medius De Bruijn et al., 1989 from the Middle
Miocene of Pakistan (De Bruijn et al. 1989), and differ from an African miocene Ctenodactylid
Africanomys pulcher Lavocat, 1961 (in Jaeger 1971) in having a transverse metalophule (the
metacone is connected to the labial part of the posteroloph in A. pulcher) while the posterior
lobe of the M3 is more reduced in A. pulcher. The Sayimys sp. nov. from the sites MS2 and Wadi
Shatirat can be regarded as the predecessor of Africanomys pulcher.
Ctenodactylids, known from the Lower Miocene of Turkey, Lower and Middle Miocene of
Pakistan and Middle Miocene of North Africa (Africa­nomys pulcher, Beni Mellal) and Israel
(Metasa­yimys) occur in the same Jebel Zelten localities as the Myocricetodontinae. The Jebel
Zelten Ctenodactyli­dae are more primitive than those from Beni Mellal, they seem to have
entered Africa at about the same time as the Myocricetodontinae or earlier.
Myocricetodontinae Lavocat, 1962
Myocricetodon Lavocat, 1952
cf. Myocricetodon sp.
In Myocricetodon cherifiensis Lavocat, 1952 and Myocricetodon parvus (Lavocat, 1961) the
cusps are more voluminous and the anterior arm of the hypo­cone is in most M1 and M2 obli100
Fig. 3. Important remains of the Miocene rodents from Jebel Zelten, Libya (occlusal views of molars).
Obr. 3. Významné nálezy miocénních hlodavců z Džebelu Zelten. Okluzální pohledy molárů.
Legend / legenda: 1–3 – Potwarmus sp. (nov. ?), 1 – M1; 2, 3 – M1; 4 – Mellalomys sp., M1; 5 – Sayimys sp., M2.
101
quely directed towards the paracone, with a ‘new’ longitudinal crest formed between hypocone
and paracone in M. par­vus (Wessels 1996). Our specimens seem to be more primitive in these
characters. Several primitive Myocricetodon species appear in the middle Miocene of Pakistan
(Wessels 1996). The specimens from Jebel Zelten are similar to Myocricetodon cf. M. parvus
from HGSP 8224 (Wessels et al. 1987) which shows a weakly developed anterior arm of the
hypocone in the M2. Our specimens seems to be more evolved. – The origin and migration pattern of the Myocrice­todontinae is not yet fully understood, but primitive Myocricetodontinae
are known from the Lower Mio­cene of Turkey (Wessels et al. 2003; MN3) and other, more
derived, Myocricetodontinae are known from Pakistan (18–13.7 Ma), Turkey (Yeni Eskihisar)
and Saudi Arabia (16 Ma). The origin and initial development of the Myocricetodontinae may
have been on the Arabian Peninsula. Mellalomys sp. is more primitive than Mellalomys atlasi
from Beni Mellal (14 Ma) and is thus considered to be older. The Myocricetodon and Mella­
lomys from Jebel Zel­ten are more primitive than those of Beni Mellal and Berg Aukas. These
Jebel Zelten localities are there­fore considered to be older than Beni Mellal (14 Ma) and Berg
Aukas (13 Ma).
Rhizomyinae Winge, 1887
Prokanisamys De Bruijn, Hussain et Leinders, 1981
Prokanisamys sp.
Prokanisamys cheek-teeth are characterised by their small size, low crowns, the cuspidate
cheekte­eth. The short mesolophid and short or absent meso­loph are regarded as primitive in
the Rhizomyinae. The teeth from Libya are similar to the rhizomyids from the Lower Miocene
of Pakistan. The oldest known rhizomyid comes from Pakistan (20 Ma; Lindsay, 1996), either
derived from a (yet unknown) Pakistani cricetodontine or migrated into Pakistan from an
unknown area. Prokanisamys sp. from the Jebel Zelten faunas is close to Prokanisamys major,
known from Pakistani assemblages dated bet­ween 19.5 and 16.4 Ma. The Rhizomyinae from
Jebel Zelten are similar to the Early Miocene taxa from Pakistan (Wessels & De Bruijn 2001)
and not to the Middle Miocene forms, therefore the immigration of the Rhizomyinae into North
Africa must have taken place in Early Miocene times. Prokanisamys sp. is considered by us to
be ancestral to Pronakalimys from Fort Ternan (14 Ma; Tong & Jaeger 1993).
Thryonomyidae Pocock, 1922
Thryonomyidae gen. nov. et sp. nov.
The morphology of six specimens from localities of 1997 collection, exclude our material
from the Thryonomyid-like genera: Paraulacodus Hinton, 1933; Neosciuromys Stromer, 1926;
Paraphiomys Andrews, 1914; Apodecter Hopwood, 1929 and Kochalia De Bruijn & Hussain,
1985. The M3 of Rodentia indet. (only one M3 and one M3) from the Middle Miocene site from
the Hadrukh For­mation of eastern Saudi Arabia (Whybrow et al. 1982) is very similar to the M3
of our material. This spe­cimen seems to represent a more evolved species of the Jebel Zelten
thryonomyid. The Thryonomyidae from Jebel Zelten are consi­dered to be more closely related
to Late Eocene Phiomyids from Algeria, and less closely to the Oli­gocene forms of Libya and
Egypt and the Miocene Phiomyids and Thryonomyidae from Eastern Africa (Lavocat 1973,
Denys 1992, Winkler 1992). The Phiomyidae become extinct after the Early Miocene, the Thryonomyidae are known from the Middle Mio­cene of Africa, Saudi Arabia, Pakistan and India.
102
Conclusions based on the record of Rodents
(Fig. 4)
The presence of Potwarmus sp. and Mellalomys sp. in the assemblages of MS2 and WS places
these assemblages betweeen 16 and 14 Ma. In conclusion, the fauna od small mammals of Jebel
Zelten localities spans approximately 4 Millions years, from 19 Ma to 15 Ma. The differences
between the particular assemblages suggest that the Jebel Zelten mammal associations represent
at least three different time periods.
Fig. 4. Paleogeographic map of Europe, North Africa and the Middle East during the Early Miocene
around 17 Ma (late Burdigalian, Ottnangian): (1) the Eastern Mediterranean seaway is closed producing
a landbridge which enabled the faunal (mammalian) interchange between Africa and Asia (B). (2) in the
West the Atantic Ocean communicated with the Tethys-Mediterranean and Paratethys (A). The northern
faunal (mammalian) interchange (C) was of different character and apparently separated from the southern one in the region of the today Balkan peninsula (B). The symbol of the star indicates the position of
Jebel Zelten.
Obr. 4. Paleogeografická mapa Evropy, severní Afriky a Blízkého východu během spodního miocénu před
cca 17 mil. let (svrchní burdigal, otnang): (1) moře ve východním Středomoří se uzavřelo a tím se vytvořil
pevninský most umožňující migrační výměnu savců mezi Afrikou a Asií (B). (2) na západě byl Atlantský
oceán spojen se Středozemním mořem (Tethydou) a dále na severovýchod s Paratethydou (A). Severní
migrační výměna savčích faun (C) se svým složením lišila od jižní (B), a byla od ní zřejmě v prostoru
dnešního Balkánského poloostrova oddělena. Hvezdičkou je označena poloha Džebelu Zelten.
103
In any case, the particularly characteristic feature of Jebel Zelten small mammal assemblages is that nearly all forms composing them reveal apparent relations both to the clades
known from other African or Arabian Paleogene or Neogene sites and to those found in early
or middle Miocene of Pakistan. This fact corresponds well to the extensive rearrangements
of the paleogeographic situation in the zone of the eastern Tethyan seaway during the Early
Miocene which promoted multiple faunal interchanges between Eurasia and Africa. Two main
migration waves have been recognized until now. The first, dated approximately to 18–19 Ma,
and the second, dated to around 16–17 Ma (Thomas 1985, Rögl 1998). Ochotonidae, primitive
cricetids, sciurids and rhizomyines invaded Africa during the first period of faunal interchange
while the anthracothere Brachyodus dispersed into Europe and Pakistan. The range expansion
of Myocricetodontinae and Ctenodactylidae fall most probably in the stage of the second migration wave but it was limited onto North Africa. If the age determination of Potwarmus is
correct (younger than 16 Ma), then Potwarmus migrated into Africa during the Middle Miocene,
perhaps during the period when Griphopothecus, Alloptox and Heterosminthus migrated into
Anatolia and Central Europe (Rögl 1998).
Unfortunately, the available fossil record is still too scarce to enable a detailed paleobiogeographic reconstructions of the Early to Middle Miocene Afro-Asiatic faunal interchange. In
any case, the situation found in Jebel Zelten demonstrate as well the vivid dispersal dynamics
in the respective taxa as the fact that the range expansions were most probably followed by
well pronounced subsequent divergences in marginal populations. In result, not only the geographically distant populations diverged but apparent anagenetic shifts can be observed during
the Early and Middle Miocene also on a local scale. Worth mentioning is that such a dynamics
most probably characterized not only the Asiatic invaders in Africa but, as the above discussed
situations in Myocricetodontidae, Phiomyidae-Thryonomyidae, and Phylisinae-Scotophilus
suggest, the clades whose ranges were centered in the North Africa and/or in Arabia in the late
Paleogene and which might contributed the respective interchange in an essential way too.
Souhrn
Práce podává přehled společenstev drobných savců získaných v průběhu let 1982, 1983 a 1997 ze spodněmiocenních uloženin Džebelu Zelten v Libyi. Celkem zde bylo nalezeno 10 druhů hlodavců, dva druhy
pišťuchovitých zajícovců a jeden druh netopýra. Ve všech případech vykazují nalezené formy zřetelné
vztahy jak k pozdně paleogenním či miocenním formám africkým tak k formám doloženým se spodního
či středního miocenu Pakistanu. Zvláště důležitým je nález myšovitého hlodavce rodu Potwarmus, který
dokládá jeden z prvních výskytů této čeledi v Africe a současně datuje nejmladší ze zkouamných faun do
úseku 15–16 miliónů let. Zkoumané fauny dokládají výmluvně velmi dynamickou tzv. “jižní” faunovou
výměnu mezi Afrikou a Asií související se změnami paleogeografické situace ve spodním miocenu. Současná “severní” výměna faun mezi Evropou a Asií měla odlišný charakter.
References
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De Bruijn H., Hussain S. T. & Leinders J. J. M., 1981: Fossil Rodents from the Murree Formation near
Banda daud Shah, Kohat, Pakis­tan. Proc. Konink. Nederl. Akad. Wetensch. B, 84: 71–99.
Horáček I., Fejfar O. & Hulva P., 2006: A new genus of vespertilionid bat from Early Miocene of Jebel
Zelten, Libya, with comments on Scotophilus and early history of vespertilionid bats (Chiroptera).
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104
Jacobs L. L., 1978: Fossil rodents (Rhizomyidae and Muridae) from Neogene Siwalik deposits, Pakistan.
Mus. North Arizona Press, 52: 1–95.
Jaeger J.-J., 1971: Un cténodactiylidé (Mammalia, Rodentia) nouveau, Irhoudia bohlini n. g. n. sp. du
Pléistocène inférieur du Maroc, rapports avec les formes actuelles et fossiles. Notes Serv. Géol. Maroc.,
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Jaeger J.-J., 1977. Rongeurs (Mammalia, Rodentia) du Miocene de Beni-Mellal. Palaeovertebrata, 7(4):
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Lavocat R., 1961: Le gisement de vertébrés Miocènes de Beni-Mellal (Maroc). Etude systématique
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109–144.
Lavocat R., 1973: Les Rongeurs du Miocène d’Afrique Orientale. Ecole Pract. Hautes Etudes (3ème
Sect.) Mém. Trav. Inst. Montpellier, 1: 1–248.
Lindsay E. H., 1988: Cricetid rodents from Siwalik deposits near Chinji Village. Part I: Megacricetodontinae, Myocricetodon­tinae and Dendromurinae. Palaeovertebrata, 18: 95–154.
Qiu Z. D., 1996: Middle Miocene Micromammalian Fauna from Tunggur, Nei Mongol. Science Press,
Beijing, 216 pp.
Rögl F., 1998: Palaeogeographic Considerations for Mediterranean and Paratethys Seaways (Oligocene
to Miocene). Ann. Naturhist. Mus. Wien 99A: 279–310.
Savage R. J. G. & Hamilton W. R., 1973: Introduction to the Miocene mammal faunas of Gebel Zelten,
Libya. Bull. Brit. Mus. Natur. Hist., Geol., 22: 515–527.
Thomas H., 1985: The Early and Middle Miocene land connec­tion of the Afro-Arabian plateau and Asia:
a major event of hominoid dispersal? Pp.: 42–50. In: Delson E. (ed.): Ancestors: the Hard Evidence.
Alan R. Liss, Inc., New York, x–xii+1–366 pp.
Tong H. & Jaeger J.-J., 1993: Muroid rodents from the middle Miocene Fort Ternan locality (Kenya) and
their contribution to the phylogeny of muroids. Palaeontographica A, 229: 51–73.
Wessels W., 1996: Myocricetodontinae from the Miocene of Pakistan. Proc. Konink. Nederl. Aka­d.
Wetensch, 99: 253–312.
Wessels W. & De Bruijn H., 2001: Rhizomyidae from the lower Manchar Formation (Miocene, Pakistan).
Ann. Carnegie Mus., 70: 143–168.
Wessels W., De Bruijn H., Hussain S. T. & Leinders J. J. M., 1982: Fossil rodents from the Chinji Formation, Banda Daud Shah, Kohat, Pakistan. Proc. Konink. Nederl. Akad. Wetensch. B, 85: 337–364.
Wessels W., Fejfar O., Peláez-Campomanes P., van der Meulen A. & De Bruijn H., 2003: Miocene small
mammals from Djebel Zelten, Libya. Pp.: 699–715. In: López-Martínez N., Peláez-Campomanes P.
& Hernández Fernández M. (eds.): Coloquios de Paleontología. En Honor al Dr. Remmert Daams.
Volumen Extraordinario 1. Universitad Complutense de Madrid, 715 pp.
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Lynx (Praha), n. s., 37: 107–110 (2006).
ISSN 0024–7774
Pipistrellus pygmaeus and two more species of bats recorded on the Island
of Kefalonia, Greece (Chiroptera: Vespertilionidae)
Zjištění Pipistrellus pygmaeus a dalších dvou druhů netopýrů na ostrově Kefalonie,
Řecko (Chiroptera: Vespertilionidae)
Jiří GAISLER
Institute of Botany and Zoology, Faculty of Science, Masaryk University, Kotlářská 2, CZ–611 37 Brno,
Czech Republic; [email protected]
received on 21 July 2006
Abstract. At a small harbour town of Poros, SE Kefalonia, numerous foraging Pipistrellus kuhlii and
P. pygmaeus were acoustically recorded (Petersson D200 bat detector) on 22 and 24 June, 2006. On
22 June, a high flying noctule, probably Nyctalus lasiopterus, was also heard. These records complete the
list of bats, known to occur on the island, to a total of five.
INTRODUCTION AND METHODS
Kefalonia (also Cephalonia, Kefaloniá, Kefallinia) is the largest of Ionian Islands. Contrary to the
more northerly Corfu (Kérkira), the bat fauna of the island has been little studied. In Argostoli,
the capital of the island, 10 individuals of Pipistrellus kuhlii (Kuhl, 1817) (3 ma­les and 1 female
identified) were recorded in 1908. Fitídi Cave, Karavómilos, was the only other known locality
where bats were observed. In 1970, one male of Rhinolophus hip­po­si­de­ros (Bechstein, 1800)
and in 1971, two males and one female of R. euryale Blasius, 1853 were collected in the cave.
For the localities and details of the records, see Hanák et al. (2001).
During a touristic sojourn at a small town of Poros in June 2006, the author and his wife
recorded the echolocation signals of bats with the aid of a Petersson D200 heterodyne bat detector. Foraging bats were seen and heard under a canopy of trees in a dry bed of a brook. In
addition to very numerous P. kuhlii (peak energy frequency 40 kHz), foraging individuals of
P. pygmaeus (Leach, 1825) were also detected (53–55 kHz). The main goal of the paper is to
con­tri­bu­te to the knowledge of the distribution of the latter species. The raison d’etre of this
taxon, originally separated from P. pipistrellus on the basis of its relatively high echolocation
signals, has been generally accepted (Simmons 2005). Its occurrence, however, is far from being
suf­fi­ci­ent­ly known.
107
RESULTS AND DISCUSSION
A locality where bats were expected to fly was discovered in the centre of Poros at a bridge over
a brook which was dry at this time of year. The only water existing during our observation of
bats was a small pool ca. 5 by 0.5–1.0 m, at the left bank upstream of the brook. The lo­ca­li­ty is
situated only about 100 m away from the estuary, its elevation therefore is roughly 0 m. While
the bed downstream of the bridge is bare, the banks of that lying upstream are grown with de­
ci­du­ous trees forming a dense canopy. On 26 June, we found that the bed was bull­do­zed and
the pool destroyed, probably to prevent the development of mosquito larvae. This situation is
shown in Fig. 1.
Acoustic bat detectoring was performed on 22 June from 21:20 to 21:45 hours and on 24 June
from 21:00 to 21:30 hours local (eastern European summer) time. In both cases, the greatest
activity was recorded at the beginning when the bats were also detected visually. No bats were
detected during the last five minutes of each observation. The bats were foraging and their feeding buzzes were frequently heard. The greatest density of foraging bats was recorded above
and close to the pool but commuting and foraging bats were also recorded upstream to about
50 m from the bridge. Within the same time span no activity was recorded downstream between
Fig. 1. Dry bed of a brook at Poros, Kefalonia, where foraging Pipistrellus kuhlii and P. pygmaeus were
recorded by a bat detector on 22 and 24 June 2006. Photo by the author.
Obr. 1. Vyschlé řečiště potoka v městečku Poros na Kefalonii, kde byli 22. a 24. 6. 2006 detektorem ul­tra­
zvu­ku zjištěni lovící netopýři druhů Pipistrellus kuhlii a P. pygmaeus. Snímek autora.
108
the bridge and the estuary. It was difficult to estimate the number of flying bats but there were
certainly more than 10 individuals flying at the pool at one time. Judging from the density of
40 kHz and 55 kHz signals, there were ca. three times more P. kuhlii than P. pygmaeus.
After the end of the first observation in the dry bed of the brook, we continued the bat detectoring walking slowly through Poros. At 21:50 h we recorded the typical loud, low fre­quen­cy
signals of a noctule. The bat (bats?) was heard for five seconds but was not seen, probably
performing a commuting flight at a high elevation. Kefalonia is a green island and the slopes
bordering Poros are grown by shrubs and some trees. Noctules, therefore, can be expected to
occur in that habitat. Since the signals we recorded had the peak frequency of 18 kHz, Nyctalus
leisleri (Kuhl, 1817) can be excluded and, of the two remaining species, N. lasiopterus (Schreber,
1780) is more likely than N. noctula (Schreber, 1774).
According to the present evidence, P. pygmaeus is distributed from British Isles, S Scan­di­na­
via, Ukraine and W Russia south to the Mediterranean (Mayer & von Helversen 2001, Hulva
et al. 2004, Simmons 2005). Molecular and phylogeographic analysis demonstrated that the
di­ver­gen­ce of P. pipistrellus and P. pygmaeus probably took place in the Mediterranean region
(Hulva et al. 2004). Mediterranean islands, therefore, can be expected to harbour one of the two
or both Pipistrellus species, depending on habitats. While P. pipistrellus is rather a generalist,
P. pygmaeus prefers lowland, in particular riparian habitats (Mayer & von Hel­ver­sen 2001).
In addition to Kefalonia, we recorded P. pygmaeus on Rhodes, Mallorca and Cyprus, of which
the Rhodes and Cypriot records were published (Hanák et al. 2001). On Mallorca (Balearic
Isles), P. pygmaeus was acoustically detected at Playa d’Alcúdia, between Port d’Alcúdia, the
sea and the hills Puig de Sant Martí, on 3 and 5 September, 2002. As in Kefalonia, P. kuhlii
were more frequently detected than P. pygmaeus. In the same habitat, Eptesicus serotinus was
recorded on 4 September. The bats were only flying where there were trees growing (Gaisler,
unpublished).
According to Hanák et al. (2001), the presence of P. pygmaeus has been demonstrated by
genetical analyses from four localities in the mainland Greece and with the aid of bat de­tec­tors
from another 11 localities in both the mainland and two Greek islands. Very likely, the species
is more common than shown by the hitherto evidence. Concerning Kefalonia, our records increase the number of bat species known to live there to five but the species identity of the noctule
remains uncertain. Considering that 12 species of bats have been known from Corfu (Hanák et
al. 2001), the list of the Kefalonian species is certainly incomplete. There are various habitats,
from lowlands up to 1620 m a. s. l., and the island is also rich in natural caves. The situation is
promising for a more systematic study the Kefalonian bat fauna.
SOUHRN
Početně létající a lovící netopýři druhů Pipistrellus kuhlii a P. pygmaeus byli zjištěni ultrazvukovým de­
tek­to­rem (Petersson D200) ve dnech 22. a 24. 6. 2006 v přístavním městečku Poros na ostrově Ke­fa­lo­nia,
Řecko. Dne 22. 6. byly zachyceny také signály vysoko letícího netopýra rodu Nyctalus, prav­dě­po­dob­ně N.
lasiopterus. Včetně námi zjištěných je v současnosti z ostrova Kefalonia známo pět druhů netopýrů.
LITERATURA
Hanák V., Benda P., Ruedi M., Horáček I. & Sofianidou S., 2001: Bats (Mammalia: Chiroptera) of the
Eastern Mediterranean. Part 2. New records and review of distribution of bats in Greece. Acta Soc.
Zool. Bohem., 53: 279–346.
109
Hulva P., Horacek I., Strelkov P. P. & Benda P., 2004: Molecular architecture of Pipistrellus pipistrellus
/ Pipistrellus pygmaeus complex (Chiroptera: Vespertilionidae): further cryptic species and Me­di­ter­ra­
ne­an origin of the divergence. Mol. Phylogenet. Evol., 32: 1023–1035.
Mayer F. & von Helversen O., 2001: Sympatric distribution of two cryptic bat species across Europe.
Biol. J. Linn. Soc., 74: 365–374.
Simmons N. B., 2005: Order Chiroptera. Pp.: 312–529. In: Wilson D. E. & Reeder D. M. (eds.): Mammal
Species of the World. Third Edition. The John Hopkins University Press, Baltimore, xxxviii+2142 pp.
110
Lynx (Praha), n. s., 37: 111–121 (2006).
ISSN 0024–7774
On some “forgotten” samples of small mammals from the Western Tatra
Mts – Roháče Mts (Slovakia) (Insectivora, Rodentia, Carnivora)
O několika “zapomenutých” vzorcích drobných savců ze Západních Tater – Roháčů,
Slovensko (Insectivora, Rodentia, Carnivora)
Jiří GAISLER1 & Jan ZEJDA2
Institute of Botany and Zoology, Masaryk University, Kotlářská 2, CZ–611 37 Brno, Czech Republic;
[email protected]
2
State Plant Protection Administration, Trnkova 103, CZ–628 00 Brno, Czech Republic; [email protected]
1
received on 22 February 2006
Abstract. In 1964 to 1974, 511 specimens of 14 mammal species were collected in the Roháče Mts, northern Slovakia, mostly at elevations of 1,000–2,000 m. The following mammalian families and species are
re­pre­sen­ted in the samples, viz, Soricidae: Sorex araneus, S. alpinus, S. minutus, Neomys fodiens; Talpidae:
Talpa europaea; Arvicolidae: Clethrionomys glareolus, Chionomys nivalis, Microtus sub­ter­ra­ne­us, M.
tatricus, M. agrestis; Muridae: Apodemus flavicollis; Gliridae: Muscardinus avellanarius, Dry­o­mys ni­te­
d­u­la; Mustelidae: Mustela nivalis. In addition, Eliomys quercinus was observed on 22 May 1964 in the
valley Spálená dolina, 1,520 m a. s. l. Besides its description, an exact drawing of the animal supports
the record. Details are given on the sample of 170 specimens obtained in September 1964 by 4,730 trap-nights at elevations of 1,500–2,100 m. Mammals were trapped at the forest upper edge (ecotone) and at
higher situated habitats up to the mountain ridge. Alpine vole species, M. tatricus and C. nivalis, clearly
dominate this sample (Table 1). Additional eight samples were obtained mostly within the forest zone and
their species diversity and numbers of records per species corroborate previous data on the development
of small mammal communities with respect to the ecosystem succession (Kratochvíl & Gaisler 1967).
Cranial me­a­su­re­ments were recorded of all adult and grown-up individuals of M. tatricus and C. nivalis
trapped in 1964–1974 in the Roháče Mts and available as museum specimens with un­da­ma­ged skulls.
The data are given in Tabs. 2 and 3, together with external measurements and body weights of the same
animals as recorded in the field. Although certain biometrics of the specimens may have been included in
earlier statistical evaluations (Kratochvíl 1970, 1981), individual measurements of particular specimens
have never been published. The purpose of their presentation in this paper is to submit data on rare and
not readily obtainable material for future examination.
INTRODUCTION
In the 1960’s, Professor J. Kratochvíl, director of the then Institute of Vertebrate Research,
CSAS, Brno, built a team to study small terrestrial mammals on the territory of the valley
Ro­háč­ská dolina in the Western Tatra Mts. In addition to problems of sampling methods dealt
with in two short papers (Kratochvíl & Gaisler 1964, Gaisler & Kratochvíl 1966), the main
goal of the study was to follow natural succession of small mammal communities. Kra­to­chvíl
was inspired by the paper of Grodziński (1959), who investigated the succession by comparing
mammal communities on an overgrown clearing and subsequent stages of a moun­tain forest
111
existing simultaneously and close to each other. Accordingly, six plots of a Sor­be­to-Piceetum
forest were selected in the Roháče Mts, representing stands of various ages from a clearing grown
with trees one to 10 years old up to a forest 100–110 years of age, considered a climax stage. In
1963, small mammal communities inhabiting the plots were sampled by a standardized method
and the results were published by Kratochvíl & Gaisler (1967). In addition to the material
obtained in 1963, results of a preliminary sampling in 1959–1962 and a sample obtained in 1964
were included as well. The latter sample, however, was only used to complete the knowledge
of the spectrum of small mammal species living on the territory in question.
The original goal of the present paper was to publish both qualitative and quantitative data
on a sample of small terrestrial mammals obtained in September 1964 in the Roháče Mts by
trapping above the upper edge of a continuous forest, at the elevation of ca. 1,500–2,100 m.
Kratochvíl himself did not participate in this trapping campaign but delegated the first author
of this paper (J. Gaisler) to form a team and organize the field work. Six persons, including
the second author (J. Zejda), participated in the work. The results obtained were entered into
a field notebook which is still deposited with the collection of the present Institute of Ver­tebra­
Fig. 1. Professor Kratochvíl and his team in front of a log cabin near Zverovka, Roháče, September 1963.
From left to right: Z. Machař (technician), F. Tenora (helminthologist), M. Lichard (microbiologist),
M. Klíma (zoologist), J. Kratochvíl (zoologist), J. Šmarda (botanist), I. Toušková (zoologist), and M.
Hejnolová (technician).
Obr. 1. Profesor Kratochvíl a jeho tým před srubem nedaleko turistické chaty Zverovka v Roháčích, září
1963. Zleva doprava (bez titulů): Z. Machař (preparátor), F. Tenora (helmintolog), M. Lichard (mi­k­ro­
bi­o­log), M. Klíma (zoolog), J. Kratochvíl (zoolog), J. Šmarda (botanik), I. Toušková (zooložka) a M.
Hejnolová (technička).
112
te Biology, AS CR, Brno. After having analysed this material, we realized that further samples
of small mammals collected in the Roháče Mts in 1964–1974 were also worth con­si­de­ring,
although their do­cu­men­ta­ti­on in field notebooks was less complete. This was pro­bab­ly due to
other than faunistical or ecological aims of the field works, such as obtaining material for anatomical, embryological, or caryological studies. In this paper, we describe those samples briefly.
Identification of the re­spe­cti­ve notebooks and names of the organizers of research are given in
the next chapter (Material etc.). All persons who took part in the fieldwork, as far as recorded
in the notebooks and excepting ourselves, are mentioned in Acknowledgments.
In addition to the basic data of the 1964–1974 samples, we aimed at the biometrics of alpine
species Chionomys nivalis and Microtus tatricus, which can still be regarded as re­la­ti­ve­ly little
known. Kratochvíl published three monographs focussed on these vole species, yet the first
one could not concern that material because of the publication date (Kratochvíl 1956). The
material collected in 1964–1974, or a portion of it, could have been included in further two
monographs (Kratochvíl 1970, 1981) but only within the statistical evaluation of samples, not
as measurements of particular individuals. In contrast, we give biometrical data of each complete specimen to enable their future use in both ecological and taxonomical studies. Numerous
papers by Slovak mammalogists (Kocian et al. 1985, Kocian & Ko­cia­no­vá 2002, Kocianová
et al. 2002, Martínková et al. 2004, Žiak et al. 2004, Adamcová 2004, Žiak 2005) are based
on recent research of C. nivalis, M. tatricus and other small mammals in the Western Tatra Mts.
Our Slovak colleagues suggested that historical data be presented on small mammal populations
comparable with the recent data (Kocian in litt.).
MATERIAL, LOCALITIES, METHODS
The samples comprise 15 mammal species the full generic names of which are given when mentioned for
the first time, henceforward by their first letter only. All taxa follow the present nomenclature, regardless of
how they were entered into the respective field notebook, e.g. Microtus tatricus, not Pitymys tatricus. The
notebook KR contains data on specimens collected in 1964–1966, other notebooks are specified below.
The first sampling on 16 to 24 June 1964 was done by J. Kratochvíl and his collaborators in spruce
forest and dwarf pine habitats in the valley Roháčská dolina at elevations ca. 1,000–1,700 m. Both standard (small) and large snap traps were used, 6,400 trap nights in total, and an unspecified number of live
traps. The traps were baited alternatively with a piece of wick soaked in fat and root vegetables, and this
method was applied in the second sampling as well. The total number of snap-trapped mammals was 123
(1.8% of trap nights), that of live-trapped 10, with the following distribution among the species: Micro­
tus subterraneus 39, Micro­tus tatricus 31, Clethrionomys glareolus 21, Sorex araneus 18, Chi­o­no­mys
nivalis 11, Microtus agrestis 8, Dryomys nitedula 2, Muscardinus avellanarius 1, Apodemus fla­vi­col­lis
1 and Neomys fodiens 1. Two species, D. nitedula and N. fodiens, were recorded by only this sampling.
One D. nitedula was trapped in a building, probably in the log cabin near Zverovka where the researchers
were accommodated, the other in a forest 1,000 m a. s. l. N. fodiens was trapped at a brook in the valley
Salatínská dolina, 1,400 m a. s. l.
The next sample, most important from the point of view of this study (see Introduction), was obtained
by J. Gaisler and J. Zejda plus four researchers on 14 to 23 September 1964. Line trapping with standard
snap traps (4,736 trap nights in total) yielded the species Sorex minutus, Sorex alpinus, S. araneus, C.
nivalis, M. tatricus, M. subterraneus, M. agrestis, C. glareolus, A. flavicollis and M. avellanarius (see
Table 1 for details). Except inside a hunting lodge just beneath the forest upper edge, probably in the
valley Spálený žlab (Kocian in litt.), all trappings were carried out in various habitats above the tree
line in the valleys Spálená dolina and Zadná Spálená dolina, around the tarn Štvrté Roháčské pleso and
on the slopes and ridges of Mts Salatín, Rákoň, Zelené and Tri Kopy. The highest situated trap line (75
traps) was set at an elevation of ca. 2050–2100 m but there was a heavy snow-fall the next night. Owing
113
to that, most traps were lost, yet a female C. nivalis was excavated from under the snow close to the first
summit of Mt Tri Kopy.
The third sample was obtained on 30 August to 3 September 1966 by J. Kratochvíl and his col­la­bo­
ra­tors, incl. J. Gaisler. Unspecified numbers of live traps were set in the valleys Roháčská and Spálená
dolina ca. 1000–1500 m a. s. l. Only specimens that died in traps were entered into the field notebook:
S. araneus 16, S. minutus 1, C. glareolus 8, C. nivalis 3, M. tatricus 2, M. subterraneus 1, A. flavicollis
2, and M. avellanarius 2.
The next sampling was focussed on the beginning of the growing season and was done by J. Kra­to­
chvíl, O. Štěrba and V. Hrabě (assisted by technicians?). Trapping, probably using snap traps, was carried
out on 24 to 27 April 1968. The results concerning reproduction of small mammals were published by
Kra­to­chvíl (1968), other data were either not utilized or incorporated in the data sets for the monographs on Pitymys species (Kratochvíl 1970) and C. nivalis (Kratochvíl 1981). The bag was obtained in
the valley Roháčská dolina in plots II and IV (see Results), ca. 1200–1300 m a. s. l., and consisted of S.
araneus 9, S. alpinus 1, Talpa europaea l, C. glareolus 21, C. nivalis 2, M. tatricus 16, M. subterraneus
11 and M. agrestis 1 (field notebook R).
The following sampling was done by B. Král and V. Králová in October 1968, the exact date not spe­
ci­fied. According to the field notebook RO, S. araneus 21, C. glareolus 20, C. nivalis 1, M. sub­ter­ra­ne­us
1, and A. flavicollis 4 were snap or live trapped in plot VI in the valley Roháčská dolina and on the bank
of the river Orava near Podbiel. Only the name of B. Král is connected with the next three samples from
the Roháče Mts (notebook CH). The first trapping was done on 28 April to 5 May 1969, the bag consisted
of C. nivalis 2, M. subterraneus 2 and M. agrestis 2, and neither traps nor localities were specified. The
second trapping, 9 to 16 June1969, concerned plot VI and a locality on the river Orava near Podbiel. The
bag of 24 specimens included M. subterraneus 19, M. agrestis 3 and M . avellanarius 2. The third sampling by B. Král, with both snap and live traps (numbers unspecified), was done in the valley Roháčská
dolina and near Oravský Podzámok on 9 to 15 June 1971. In the field notebook, the following localities
are specified: environs of the chalet Ťatliakova chata, plots II, III and VI, at a brook (probably Studený
potok) and above a waterfall. The bag of 30 specimens consisted of C. glareolus 5, C. nivalis 2, M. sub­
terraneus 8, M. tatricus 7, M. agrestis 3, A. flavicollis 4. A weasel, Mustela nivalis, was live trapped. The
last sampling in the Roháče Mts was carried out by O. Štěrba, R. Obrtel and V. Hrabě in September
1974. We found neither an exact date nor information about the localities in the respective notebook (CH).
The bag consisted of C. glareolus 3, M. tatricus 1, M. agrestis l and Microtus arvalis 1. Since the latter
species was not recorded in any other sample and the respective specimen was not preserved, we consider
the record of M. arvalis questionable and exclude it from the evaluation.
Museum specimens of M. tatricus and C. nivalis collected in 1963–1974 in the Roháče Mts and de­po­si­
ted in the collection of the Institute of Vertebrate Biology were examined. Only adult and fully grown-up
individuals were selected and their skulls checked for completeness. Undamaged or nearly undamaged
skulls were then measured under a stereo microscope with a vernier calliper (J. Zejda). Data on external
me­a­su­re­ments and weights of the same specimens were assumed from field notebooks. All me­a­su­re­ments
are specified in Table 2. To determine male maturity and sexual activity, a testovesicular index (TVI) was
cal­cu­la­ted as a multiple of testicle and vesicular gland lengths (Hrabě 1972).
RESULTS AND DISCUSSION
During eleven years 1964 to 1974, 87 insectivores, 423 rodents and 1 carnivore, totalling 511
small mammals, were trapped in the Roháče Mts by Brno zoologists. The species are re­pre­sen­ted
by the following number of individuals: S. araneus 81, S. alpinus 3, S. minutus 2, N. fodiens 1,
T. europaea 1, C. glareolus 107, C. nivalis 58, M. subterraneus 114, M. tatricus 102, M. agrestis
20, A. flavicollis 14, M. avellanarius 6, D. nitedula 2, and M. nivalis 1.
The results of the September 1964 sampling are summarized in Table 1. The total number of
trapped individuals represents 3.6% of the number of trap nights. Relatively highest (18.8%)
114
Tab. 1. The main sample according to species and habitats, TN = trap nights, n = individuals, % = individuals in percent of trap nights. Habitats: A – hunting lodge; B – upper forest edge; C – dwarf pine stands;
D – brook; E – tarns; F – meadows up to 2,000 m a. s. l.; G – block fields up to 2,000 m; H – ridge above
2,000 m
Tab. 1. Hlavní vzorek podle druhů a prostředí, TN = počet pastí krát nocí, n = počet jedinců, % = n jako
% TN. Prostředí: A – lovecká chata; B – horní hranice lesa; C – kosodřevina; D – potok; E – plesa; F
– louky do 2000 m n. m.; G – balvanitá pole do 2000 m; H – hřeben nad 2000 m
habitat / prostředí TN A
16
B
1200
C
925
D
620
E
300
F
680
G
920
H
75
S. araneus n
%
S. minutus n
%
S. alpinus n
%
C. nivalis n
%
M. tatricus n
%
M. subterraneus n
%
M. agrestis n
%
C. glareolus n
%
A. flavicollis n
%
M. avellanarius n
%
0
0
0
0
0
0
0
0
0
0
0
0
0
0
2
12.5
1
6.3
0
0
11
0.9
1
0.1
1
0.1
8
0.7
14
1.2
4
0.3
1
0.1
9
0.8
0
0
1
0.1
1
0.1
0
0
0
0
8
0.9
10
1.1
1
0.1
0
0
6
0.7
0
0
0
0
2
0.3
0
0
1
0.2
2
0.3
7
1.1
11
1.8
1
0.2
6
1.0
2
0.3
0
0
1
0.3 0
0
0
0
4
1.3
8
2.7
0
0
0
0
3
1.0
0
0
0
0
1
0.2
0
0
0
0
0
0
2
0.3
16
2.4
0
0
0
0
0
0
0
0
1
0.1
0
0
0
0
14
1.5
4
0.4
1
0.1
0
0
3
0.3
0
0
0
0
0
0
0
0
0
0
1
1.3
0
0
0
0
0
0
0
0
0
0
0
0
total / celkem 3
18.8
50
4.2
26
2.8
32
5.1
16
5.3
19
2.8
23
2.5
1
1.3
n
%
was the bag from inside a hunting lodge where two common forest species, C. glareolus and A.
flavicollis, were captured. In contrast, relatively lowest (1.3%) was the bag from a moun­tain ridge
above 2,000 m where only one C. nivalis was found in a trap due to unfavourable weather (see
Material). For various reasons the two samples A and H are incomparable with the re­ma­i­ning
samples and are excluded from further evaluation. Among the samples B to G, that from the
environs of the tarn Štvrté Roháčské pleso (E, 5.3%) and from the bank of an unspecified brook
(D, 5.1%) show relatively high percentages of trapped mammals. The sam­p­le from the upper
border of the forest stand (ecotone) in the valley Zadná Spálená dolina (B, 4.2%) ranks third,
comprising nine species and thus showing the highest species diversity. In samples B to G, M.
tatricus is represented by 45 specimens, thus being the most often trap­ped species. Its relative
abundance ranges from 0.3% (meadows) to 2.7% (tarn environs). The second most common
species, C. nivalis, is represented by 36 specimens with relative abun­dan­ce ranging from 0.3%
(brook) to 1.5% (rock fields), the species was not recorded in alpine meadows. In contrast, M.
subterraneus, represented by the total of 33 specimens, was most often trapped in meadows
115
(2.4%). Next comes C. glareolus with 27 specimens evenly distributed among various habitats
(0.3–1.0%), missing in the habitat meadows only. S. ara­ne­us was the most common shrew
species. The 17 specimens of this shrew are evenly dis­tri­bu­ted over all habitats though in low
numbers (0.1–0.9%). The remaining species are only represented by 1–2 specimens each.
The qualitative (species) and quantitative composition of further eight samples has already
been specified under Material etc. We are unable to evaluate this material according to ha­bi­tats
Fig. 2. Habitat of the snow vole (Chionomys nivalis) in the Spálená dolina Valley, with Mt. Tri Kopy in
the background, September 1966. Both photos by J. Gaisler.
Obr. 2. Stanoviště hraboše sněžného (Chionomys nivalis) na konci Spálené doliny, v pozadí v mracích
vrcholky Trech kop, září 1966. Oba snímky J. Gaislera.
116
Tab. 2. External and cranial measurements of Microtus tatricus. Explanations: No. – logo of the notebook
and excursion number; G – weight; H+B – head and body length; T – tail length; HF – hind foot length; E
– ear length; M – male; TVI – testovesicular index (Hrabě 1972); F – female; gr – pregnant; MC – placental scars; Cb – condylobasal length; LSp – length of splanchnocranium (measured from prosthion to the
aboral edge of M3); DMx – length of the maxillar diastema; Zyg – zygomatic breadth; LMd – maximum
length of mandible (measured to the aboral edge of processus angularis); DMd – length of the mandibular
diastema. Weight in g, measurements in mm, cranial ones to the nearest 0.1 mm
Tab. 2. Tělesné a lebeční rozměry Microtus tatricus. Vysvětlivky: No. – označení protokolu a exkursní
číslo; G – hmotnost; H+B – délka těla; T – délka ocasu; HF – délka zadní tlapky; E – výška ušního boltce;
M – samec; TVI – testo-vesikulární index (Hrabě 1972); F – samice; gr – březí; MC – děložní skvrny;
Cb – kondylobazální délka lebky; LSp – délka splanchnokrania (měřená od prosthionu k zadnímu okraji
M3), DMx – délka horní diastemy; Zyg – zygomatická šířka; LMd – největší délka mandibuly (měřená
po zadní okraj processus angularis); DMd – délka dolní diastemy. Hmotnost v gramech, rozměry v mm,
u lebečních s přesností 0,1 mm
No.
KR 13
KR 38
KR 46
KR 47
KR 48
KR 63
KR 92
KR 97
KR 98
KR 100
KR 130
KR 134
KR 162
KR 214
KR 234
KR 258
RY 31
RY 33
G
31.5
25.5
31.5
28.0
48.0
29.5
26.0
24.0
30.5
29.5
26.0
29.5
26.0
23.5
25.0
24.0
31.0
25.0
H+B
T
HF
113
40
114
41
111
44
109
41
104
43
116
48
106
103
37
101
112
41
103
43
107
43
112
40
109
39
102
42
100
40
111
44
104
40
17.5
17.0
18.0
18.0
17.5
18.0
17.0
17.0
17.5
18.0
18.0
17.0
18.0
17.0
17.0
17.0
18.5
18.0
E
M TVI
F
Cb
11.5
168
12.0
80
24.6
11.0 grMC 25.1
10.5
121
24.8
10.5 MC 24.1
12.0
121
25.3
12.0
110
24.7
13.0 grMC 24.1
11.5 gr 23.8
11.5
108
24.6
12.0 gr 24.2
11.0 72 R
24.6
11.0 gr 24.1
11.0 MC 24.2
11.0 MC 24.3
11.2
70
24.3
11.5
120
25.1
11.5
110
LSp DMx Zyg LMd DMd
15.3
15.1
15.3
14.9
14.7
15.4
14.8
14.8
14.9
15.5
15.2
14.9
15.3
14.8
15.1
15.1
15.4
14.3
7.7 14.5
7.7 14.4
7.8 14.8
7.7 14.6
7.5 14.3
8.1 14.7
7.5 14.4
7.2 13.8
7.8 13.7
7.5 14.5
7.4 14.1
7.9
7.5 14.5
7.4 14.2
7.5
7.8 14.2
8.1 15.2
7.5 14.3
14.5
14.4
14.3
14.3
14.2
15.2
14.2
14.1
14.6
14.2
14.5
14.5
14.2
13.9
14.5
13.8
14.7
14.1
4.2
4.1
4.2
3.6
4.1
4.7
3.8
4.1
4.3
4.1
4.1
3.9
3.8
3.8
4.0
4.1
4.3
4.2
due to the lack of pertaining data contained in the respective field notebooks. In these no­te­books,
trapping localities are often denoted by Roman numbers of study plots. They co­r­re­spond to the
plot labelling in Kratochvíl & Gaisler (1967) where plots I to VI are descri­bed in detail and
entered in a map. Most often, plot VI was mentioned as the locality trapped in 1964 and later.
It was situated at the elevation of 1,150–1,250 m and grown with a stand resembling an old
growth forest. The large number of species and the relatively large number of trapped animals
correspond to the data in Kratochvíl & Gaisler (l. c.) who recorded the largest number of species
and the second largest number of individuals on plot VI. The same number of species (nine)
and a still larger number of individuals was recorded on plot I, but that plot was not trapped in
1964 and later. Instead, two other plots were selected. Plot II was grown by patches of P. abies
10–20 years old at the elevation of 1,950–1,130 m and there was a permanent brook. Plot III
was grown by a dense spruce forest 20–30 years old at the same elevation as plot II. Again, the
samples taken in 1964 and later on correspond to the earlier data on the large number of species
117
(13 on plot II and 11 on plot III) and on the average or low number of individuals (the lowest
on plot III from all plots studied) (Kratochvíl & Gaisler l. c.).
Considering all samples obtained in 1964 and later on in the Roháče Mts, the high re­pre­sen­
ta­ti­on of alpine vole species M. tatricus a C. nivalis is obvious. This can be due to the fre­quent
trappings at high elevations. In the total material, only C. glareolus and M. sub­ter­ra­ne­us were
still more numerous. The former species was recorded in a variety of forest or fo­rest-like ha­bi­
tats, which corresponds to its nature as a forest generalist. The latter was most often recorded in
habitats corresponding to early stages of forest succession, as far as it can be deduced from the
data in the field notebooks. In general, the data underlying this paper correspond to the earlier
ones presented by Kratochvíl & Gaisler (1967) and Kratochvíl (1956, 1970, 1981) and to the
recent ones obtained by Slovak authors (Kocian et al. 1985, Kocianová et al. 2002, Mar­tín­ko­vá
et al. 2004, Žiak et al. 2004, Adamcová 2004, Žiak 2005). The only major difference seems to
be in the extremely low representation of A. flavicollis in the 1964–1974 material, compared to
both earlier and later samples from the Roháče Mts, the causes of which are unclear.
In addition to specimens collected, the field notebook KR contains an observation and drawing
concerning a species not recorded by trapping. On 22 May 1964 at 11:30 a.m., M. Klíma (Brno)
and M. C. Saint-Girons (Paris) observed a dormouse, Eliomys quercinus, in the valley Spálená
dolina near a small tarn at the elevation of 1,520 m, where the yellow and blue marked tourist
pathways meet (or did so at that time). That tarn was probably Zelené pleso below Mt Predné
Zelené (Kocian in litt.). The dormouse was observed from a distance of two metres as it climbed
down a shrub (dwarf pine?), ran slowly across an open space and climbed up another shrub. Its
distinct species-specific characters were recorded, namely the grey brownish upper parts, large
eye, black stripe from eye to ear, and the enlarged black and white tail tip. Its tail appeared to be
longer than head and body. We consider the description and species determination by two experienced zoologists reliable. Furthermore, M. Klíma, now professor emeritus of human ana­to­my,
is a well-known artist and illustrator. To our knowledge, this observation has not been published
so far. According to Anděra & Horáček (1982), the species was mentioned “in earlier reports
from Orava“. The authors do not quote the source of that information. Very likely, however, it
had been adopted from Kocyan, a well known naturalist who worked in the region of Orava in
the second half of the 19th century. The following is the statement by Kocyan (1888) about the
garden dormouse, referred to as Myo­xus quercinus: “Sehr selten. Ich habe während 24 Jahren
nur zwei Exemplare aus dem Hochgebirge bekommen”. Although no particular locality can be
inferred from the term “Ho­ch­ge­bir­ge” (high mountains), it implies occurrence at high elevations
(Kocian in litt. – by the way, Kocian is the grandson of Kocyan). Therefore, the observation by
Klíma and Saint-Girons is not as unlikely as it could appear at first sight.
The external and cranial measurements of adult and grown up individuals of M. tatricus and
C. nivalis available as museum specimens are given in Tables 2 and 3. In two M. tatricus and
three C. nivalis, the zygomatic breadth could not be reliably determined and in one C. nivalis
the maximum length of mandible was undeterminable. The skulls of other specimens were
complete. Of the total number of animals trapped, only a small percentage could be used in this
respect, viz., 17.6% of M. tatricus and 37.9% of C. nivalis. On the one hand, this reflects the
low re­pre­sen­ta­ti­on of fully adult individuals in the sample and, on the other, the un­su­i­ta­bi­li­ty
of com­mer­cial snap traps that damage skulls of captured animals. All measurements pre­sen­ted
in Tabs. 2 and 3 lie within the minimum-maximum range known in adults of the two species
(cf. Kratochvíl 1970, 1981) and they corroborate the species-specific status of the two taxa.
The data are de­si­gned for future analyses and the specification of each specimen has to assist
118
Tab. 3. External and cranial measurements of Chionomys nivalis. For explanations see Tab. 2
Tab. 3. Tělesné a lebeční rozměry Chionomys nivalis. Vysvětlivky u tabulky 2
No.
G
H+B
T
HF
E
M TVI
F
Cb
KR 13 31.5 113
40 17.5 11.5
168
KR 119 52.0 123
61 20.5 18.0 grMC
KR 120 115
61 21.0 MC 28.8
KR 121 53.0 123
56 21.0 16.0 gr 28.6
KR 122 58.5 126
54 21.0 16.0 MC 28.7
KR 124 49.5 125
54 21.5 16.5
255
28.7
KR 125 58.5 130
61 21.0 16.0
224
29.1
KR 132 49.0 126
64 21.0 16.0
182
29.1
KR 145 53.5 125
61 21.0 16.0 75 R
30.2
KR 157 43.0 119
53 21.0 16.0 MC 27.9
KR 173 42.0 117
55 21.0 17.0 MC 28.2
KR 190 46.0 124
62 21.0 16.5 MC 29.1
KR 199 48.0 123
60 20.4 15.8 MC 29.1
KR 217 41.5 119
54 21.0 18.0
6 R
27.9
KR 262 46.5 115
62 20.0 16.0 MC 28.1
KR 299 56.5 122
57 21.0 19.0 MC 30.5
CH 286 42.5 123.5
58 20.5 16.8 MC 29.1
CH 293 43.0 128.5 59,5 20.8 16.8 gr 29.2
RY 89 54.0 115
54 19.0 18.0
120
RY 122 58.0 126 19.0 15.0
162
30.1
RY 125 54.0 116
57 21.0 15.0
224
28.5
RY 167 54.0 117
60 20.0 16.0 MC 28.2
RY 174 60.0 121
67 20.0 17.5
gr 30.1
LSp DMx Zyg LMd DMd
15.3
17.7
17.3
17.4
16.9
17.1
17.2
17.2
17.7
16.8
16.9
17.3
17.5
16.4
17.2
18.4
17.1
17.5
17.1
17.7
16.8
16.9
17.5
7.7 14.5 14.5
9.5 17.1 17.9
9.0 17.1 18.1
9.1 16.7 16.5
9.1 17.1
8.3 17.1 17.1
9.2 17.2 17.1
9.3 16.5 16.7
9.3 16.9 17.6
9.1 16.1 17.2
9.1 16.6 16.8
9.1 16.7 16.5
9.3 17.1
8.4 16.1 16.2
8.8 16.2 16.8
9.3 17.5 17.7
9.4 16.4 17.8
9.4 16.8 16.6
9.1 16.7
9.8 16.7 16.7
9.3 16.7 16.7
8.5 16.4
9.4 16.7 17.1
4.2
5.7
5.7
4.5
5.2
4.7
4.9
4.5
4.9
4.7
4.7
4.8
5.1
4.7
4.8
4.7
5.1
5.2
4.7
5.7
4.7
3.5
5.1
researchers who wish to use more sophisticated methods (epigenetic, molecular, etc.) to locate
the material in the collection.
SOUHRN
V letech 1964 až 1974 bylo v Roháčích na severním Slovensku, většinou v nadmořských výškách
1000–2000 m, uloveno 511 jedinců 14 druhů savců. Ve vzorcích jsou zastoupeny tyto čeledě a druhy:
Soricidae: Sorex araneus, S. alpinus, S. minutus, Neomys fodiens; Talpidae: Talpa europaea; Arvicolidae:
Cle­th­ri­o­no­mys glareolus, Chionomys nivalis, Microtus subterraneus, M. tatricus, M. agrestis; Muridae:
Apodemus flavicollis; Gliridae: Muscardinus avellanarius, Dryomys nitedula; Mustelidae: Mustela nivalis.
Kromě toho byl 22. 5. 1964 pozorován ve Spálené dolině, 1520 m n. m., živý jedinec Eliomys quercinus.
Pro vě­ro­hod­nost pozorování svědčí nejen popis, ale i zdařilá kresba zvířete. Podrobně je zhodnocen vzorek 170 jedinců drobných savců získaný v září 1964 v nadmořských výškách 1500–2100 m odchytem do
sklapovacích pastí (4730 pastí × nocí). Pasti byly kladeny při horním okraji smrkového lesa (ekotonu) a ve
vyšších polohách až po hřebeny. Ve vzorku zřetelně dominují vysokohorské druhy hrabošů M. tatricus
a C. nivalis (tab. 1). Dále jsou podány základní informace o složení jiných osmi vzorků z Roháčů. Tyto
odchyty do sklapovacích nebo živolovných pastí byly většinou provedeny v pásmu lesa Sorbeto-Picee­
tum a jejich složení potvrzuje dřívější poznatky o vývoji populací drobných savců během sukcese tohoto
ekosystému (Kratochvíl & Gaisler 1967). Ze sbírkového materiál druhů M. tatricus a C. nivalis z téže
oblasti a období byly vybrány nepoškozené lebky dospělých a plně dorostlých kusů. Zjištěná kra­ni­o­me­t­
ric­ká data spolu s tělesnými rozměry a údaji o váze přejatými z terénních protokolů jsou uvedena v tab. 2
119
a 3. Tento materiál mohl být částečně či zcela zahrnut v dřívějších publikacích (Kratochvíl 1970, 1981),
ale jen v rámci statistického hodnocení souborů, rozměry jednotlivých kusů publikovány nebyly. Cílem
jejich zveřejnění je poskytnout data o vzácném a ne­snad­no získatelném materiálu pro pozdější analýzu.
ACKNOWLEDGMENTS
The authors wish to thank Ľ. Kocian (Bratislava) for stimulus to compile the paper, various advices con­cer­
ning literature and geographic names, and readiness to consult any problems connected with the Western
Tatra Mts and their small mammals. The authors are also indebted to all who participated in field trips to
obtain small mammal specimens in the Roháče Mts in 1964–1974, some of whom are no more among us.
They were, in alphabetic order: M. Hejnolová, V. Hrabě, J. Kocian, B. Král, V. Králová, J. Kratochvíl,
L. Kratochvílová, R. Obrtel, O. Štěrba, F. Tenora, and M. Zapletal (Brno). Finally, the authors wish
to thank J. Zima and P. Koubek for loan of field notebooks and authorization to study museum specimens
in the collection of the Institute of Vertebrate Biology, AS CR in Brno.
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ta­t­ran­ské­ho (Chionomys nivalis mirhanreini Schaefer, 1935) v Západných Tatrách – Roháčoch [Home
range characteristics of Tatra snow vole (Chionomys nivalis mirhanreini Schaefer, 1935) in the West
Tatra mountains-Roháče]. Pp.: 121–130. In: Urban P. (ed.): Výskum a ochrana civcavcov na Slovensku
V. Zborník referátov z konferencie (Zvolen 12.–13. 10. 2001) [Research and Protection of Mammals in
Slovakia V. Proceedings from the conference (Zvolen 12–13 October 2001)]. Štátna ochrana prírody
Slovenskej republiky, Banská Bystrica, 174 pp (in Slovak, with an abstract in English).
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Kratochvíl J., 1956: Hraboš sněžný tatranský Microtus (Chionomys) nivalis mirhanreini Schaefer, 1935
[Tatra snow vole Microtus (Chionomys) nivalis mirhanreini Schaefer, 1935]. Acta Acad. Sci. Čechoslov.
Brno, 28(1): 1–39 (in Czech).
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von Kleinsäugern bei ökologischen und populationsdynamischen Arbeiten. Zool. Listy, 13: 289–294.
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-Piceetum. Zool. Listy, 16: 301–324.
Martínková N., Žiak D. & Kocian Ľ., 2004: Habitatová selekcia drobných cicavcov v heterogénnom
prostredí subalpínského stupňa Západných Tatier [Habitat selection of small mammals in heterogeneous landscape of subalpine zone in the Western Tatra Mountains]. Pp.: 167–181. In: Adamec M. &
Urban P. (ed.): Výskum a ochrana cicavcov na Slovensku VI. Zborník referátov z konferencie (Zvolen
10.–11. 10. 2003) [Research and Protection of Mammals in Slovakia VI. Proceedings of the Conference
(Zvolen 10–11 October 2003)]. Štátna ochrana prírody SR, Banská Bystrica, 189 pp (in Slovak, with
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in the Roháčske plesa National Nature Reserve]. Doktorandská práca, Bratislava, 64+34 pp.
Žiak D., Kocianová-Adamcová M., Kocian Ľ. & Martínková N., 2004: Vysoká diverzita drobných
zemných cicavcov v subalpínskom stupni Západných Tatier [High diversity of small mammals in the
subalpine zone of the Western Tatra Mts]. Pp.: 45–57. In: Adamec M. & Urban P. (ed.): Výskum a ochrana
cicavcov na Slovensku VI. Zborník referátov z konferencie (Zvolen 10.–11. 10. 2003) [Research and
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Štátna ochrana prírody SR, Banská Bystrica, 189 pp (in Slovak, with an abstract in English).
121
Lynx (Praha), n. s., 37: 123–130 (2006).
ISSN 0024–7774
Brown-nosed coati (Nasua nasua vittata) on the Roraima tepui
(Carnivora: Procyonidae)
Výskyt nosála červeného (Nasua nasua vittata) na stolové hoře Roraima
(Carnivora: Procyonidae)
Pavla HAVELKOVÁ 1, Jan ROBOVSKÝ 1, Marek AUDY2 & Amelia DÍAZ DE PASCUAL3
1
Department of Zoology, Faculty of Biological Sciences, University of South Bohemia, Branišovská 31,
CZ–370 05 České Budějovice, Czech Republic; [email protected]; [email protected]
2
Tyršova 332, CZ–679 06 Jedovnice, Czech Republic; [email protected]
3
Department of Biology, Faculty of Sciences, University of Los Andes, 5101 Mérida, Venezuela;
[email protected]
received on 12 June 2006
Abstract. Tepuis are geomorphologically (physically) isolated mountains in the northeastern area of South
America. Their long-time isolation has promoted the evolution of a highly endemic fauna at the genus and
species level. In general, the rare incidence is typical for all vertebrates, especially for mammals (there are no
fishes on the tops of the tepuis). Herein we report of two independent observations of coatis (genus Nasua)
in March 2002 and January 2003 on Roraima tepui (on the borders of Venezuela, Guyana and Brazil). The
observed animals possess evident distinctive black and olive-brown coloration. Based on its distribution
and published reports we suppose that these individuals belong to the subspecies of Brown-nosed coati
(Nasua nasua vittata). Its distribution on Roraima tepui could be interpreted as a case of habitat and food
source opportunism in coatis. The observation of January 2003 includes aggressive interaction between
two individuals. Our observations could be added to other examples of mammal distribution on tepuis.
The exact number of coatis on Roraima, their detailed feeding strategy and the potential dependence on
human activity in this area remain unclear.
INTRODUCTION
The Guayana highlands are a complex of igneous and sedimentary formations in the nor­the­as­tern
part of South America. The sedimentary formations take the form of many isolated sand­sto­ne
and quartzite mesas (tepuis). These tepuis are of archean and proterozoic age, making them
one of the oldest massifs on Earth (e. g. Rosales & Huber 1996, Givnish et al. 2000, Kelloff
& Funk 2004). Tepuis are typical with orthogonal hillsides that caused the long isolation of
their plateaus from the surrounding landscape. Consequently, very specific habitats developed
on top of tepuis with a high percentage of plant and animal endemism (e. g. Rosales & Hu­
ber 1996, Pérez-Zapata et al. 1992, Rull 2005). In spite of the high en­de­mis­m, tepuis do not
house some “Lost World fauna” sensu Sir A. I. Conan Doyle. In contrast, the Roraima tepui is
disproportionately much more barren than the belletristic au­thors supposed. All tepuis are, in
fact, a paradise for geologists (and recently for spe­le­o­lo­gists too) (e. g. Rosales & Huber 1996,
Audy 2003), botanists and algologists (e. g. Rull 1991, Pokorný 1996, Givnish et al. 2000,
123
Kelloff & Funk 2004, Rull 2004a, Rull 2005), and entomologists, but vertebrate zoologists
are partly unsatisfied (e. g. Rosales & Huber 1996). Vertebrates are poorly distributed on tepuis
– represented by some endemic frogs (e. g. Rivero 1961, Zweifel 1986, Myers & Donnelly
2001), reptiles (e. g. Myers & Donnelly 2001, Macculloch & Lathrop 2004), birds (e. g. Bar­
ro­wclou­gh et al. 1997, Barber & Robbins 2002, Braun et al. 2003, Hilty 2003) and mammals
(e. g. Pérez-Zapata et al. 1992). Mammals on tepuis are scarce and probably represent stray
animals (e.g., Panthera onca in Auyán tepui) or extremely rare endemic taxa (e.g. Podoxymys
roraimae) (Pérez-Zapata et al. 1992, Pokorný 1996). For a long time, the tepuis were considered as refugia for archaic forms (“living fossils”) (the Lost World hypothesis – for details see
Rull 2004a, b), but, in spite of high endemism new research shows that tepui biotas have many
relatively young con­necti­ons with other biogeographical areas (e.g., Andean region, Amazonia
etc.) (e. g. Rivero 1964, Pérez-Zapata et al. 1992, Da Silva & Patton 1998, Kelloff & Funk
2004, Steiner & Cat­ze­f­lis 2004) with vertical movements of biotas during the Pleistocene
glacial cycles (the Vertical Displacement hypothesis – for details see Rull 2004a, b). Nowadays, the dispersal scenario (i.e., repeated dispersion and local speciation) in the combination
with vicariant sce­na­rio is more considered than simply vicariant (refuge) scenario (for details
see e. g., Rull 2004a, b, c, 2005) – in this context, many so-called living fossils (e. g., the toad
Oreophrynella) are now unsupported illusions.
Mount Roraima (5° 12’ N, 60° 44’ W, 2810 m a. s. l.) represents one of the highest tepuis and
is situated on the secondary deforested savanna (called Gran Sabana) on the borders of Ve­ne­zu­
e­la, Brazil and Guyana. Roraima possesses a typical flora with high percentage of endemism
(Pokorný 1996). It is also occupied by some typical birds (Barber & Robbins 2002, Braun et
al. 2003, Hilty 2003), and other endemic herpetological fauna like e.g., Ro­rai­ma Bush Toad
(Ore­o­phry­nel­la quelchii) (Rivero 1961), Colostethus roraima (Barrio 2004) and the rattlesnake
Crotalus durissus ruruima (Hoge 1966) and very rare example of mammalian endemism of
tepuis – akodontine cricetid Podoxymys roraimae (Pérez-Zapata et al. 1992). We report here
on the occurrence of Brown-nosed coati (Nasua nasua) on the Roraima tepui. Our observations
are interesting for two main reasons: first, the occurrence of coatis has not been reported from
the top of tepuis yet (Mondolfi 1997) – coatis inhabit mainly woodland areas (Nowak 1991).
Se­cond, the observed coatis were of a relatively unu­sual appearance in comparison with in­di­
vi­du­als of normal-colored Nasua nasua with reddish brown to black general coloration and
yel­lowish to dark brown below (Nowak 1991).
RESULTS AND DISCUSSION
The first author (PH) observed and took a picture of one unusually colored adult specimen
of Brown-nosed coati, Nasua nasua (Linnaeus, 1766) on the top of Roraima tepui in March
2002 (Fig. 1). In January 2003, M.A. repeatedly observed and photographed two individuals
(Figs. 2, 3) of the same unusual coloration. During both these occasions (2002 and 2003), the
observed in­di­vi­du­als were not too shy. Good quality photographs were taken in January 2003,
due to repeated visits of the camp by one (?) male of coati. All observations were made during
daytime, which is congruent with the known diurnal activity of coatis (Nowak 1991).
The appearance of these coatis is relatively unusual in comparisons with the typical co­lo­ra­
ti­on of Brown-nosed coati with orange or reddish to dark brown pelage coloration and head
with white spots around the eyes (for more details see Gompper & Decker 1998). However
there is extensive variation known throughout the distribution range of Nasua nasua (Gomp­per
124
& Dec­ker 1998). The individual on the Figs. 2 and 3 is probably an adult male with re­la­ti­ve­ly
slender snout and a yellow-black pattern of pelage. Its head is black without distinct facial light
mar­kings, only one small yellow spot is situated on the right side of left eye (for left lateral
view see Fig. 2). The nose is suspiciously gray. Dorsal side of the ears and fur behind them
is orange or orange-brown. The black color of the head continues over the dorsal side of the
neck to the level of shoulders. Feet and distal parts of limbs are black. Dorsal side of the tail
Fig. 1. Our first observation of the distinct colored coati on the Roraima tepui (2002). Photo by Pavla
Havelková.
Obr. 1. Naše první pozorování odlišně zbarveného nosála na Roraimě (2002). Snímek Pavly Havelkové.
125
Fig. 2. The male of coati near the speleologists’ camp (2003). Photo by Marek Audy.
Obr. 2. Samec nosála při návštěvě speleologického tábora (2003). Snímek Marka Audyho.
is blackish melting to brown at the tip, ventral side is yellow. Approximately six black rings,
which are about twice as wide as the light ones, and a black tip are well re­co­gni­za­ble on the
tail. The rest of the body (throat, trunk, proximal parts of limbs) are light – yellow or beige, but
the underfur is probably black (Fig. 3). This individual was determined as Nasua nasua vittata
Tschudi, 1844, based on published data (Allen 1904, Gompper & Decker 1998, Linares 1998)
and consultations with the fourth author. Observed individuals correspond in almost all as­pects
with the description by J. A. Allen of Nasua phaeocephala (junior synonym of N. n. vittata).
This form was originally described by J. J. von Tschudi as Nasua vittata from the region near
Roraima tepui. In Venezuela, two subspecies of Nasua nasua occur – N. n. dorsalis Gray, 1866
in the Andean region and N. n. vittata that is distributed in the region south of the Orinoco River,
but it is minimally reported from tepuis, although it was recorded from Auyán tepui (Linares
1998). It would be interesting to investigate using molecular methods how old and how much
differentiated are the subspecies of Nasua nasua throughout their range.
The distribution of such a relatively big carnivore is quite unexpected in so barren habitat.
The third author (M.A.) had the opportunity of repeated and long-lasting observations of this
coati. His observations include an aggressive interaction (a tag) between two individuals. The
stron­ger individual drove its rival by hardly defined bark to the edge of the tepui. These re­pea­
ted observations were possible due to several visits of coatis to the expedition camp. It could
126
imply some exploitation of the human touristic activity by coatis at the top of Roraima tepui.
Diet of the Brown-nosed coati consists mainly of forest floor invertebrates, fruits and car­ri­ons
with a minor amount of vertebrates (fishes, snakes, infrequently chickens in urban en­vi­ron­ment,
small mam­mals or eggs of caiman) (Bisbal 1986, Gompper & Decker 1998). Its ecology (e.g.,
the presence in secondary forests, etc.) implies ecological plasticity (Eisenberg 1989, Gompper &
Decker 1998). In this point of view, the coati as an omnivorous opportunist is well predisposed
for its distribution on tepuis. We can only estimate the diet of the Roraima coati – it may consist
of rare fruits, ver­tebra­tes (e.g., the endemic frogs and akodontine rodents), invertebrates (e. g.,
endemic beetles and many other insects and spiders etc.), and the­o­re­ti­cal­ly birds or their eggs,
plus finally some human foodrests. Recently Beisiegel & Mantovani (2006) showed relatively
robust home range shifts of Nasua nasua in a period of three years in a pluvial tropical Atlantic
forest area. These shifts could be connected to resource avai­la­bi­li­ty. The same authors also
described coatis foraging for food (invertebrates and vertebrates) inside bromeliads as part of
Fig. 3. Portrait of the coati from Roraima tepui with its remarcable distinct coloration (2003). Photo by
Marek Audy.
Obr. 3. Portrét neobvykle zbarveného nosála z Roraimy (2003). Snímek Marka Audyho.
127
their typical behavior. The bromeliads are relatively abundant on top of the Roraima tepui, and
this behavior could thus facilitate the survival of coatis in this relatively inhospitable habitat.
There are two different points of view for this coati’s distribution. It could either be caused by
a stray of a few individuals, or be a regular distribution, either seasonal or permanent (stable).
The following conditions could be of importance: First, the slopes of Roraima and Kukenán
(the tepui in near neighborhood of Roraima) are covered by tropical rain forest which could
re­pre­sent a suitable and stable habitat for coatis, in contrast to the secondary deforested dry and
grassy savanna. Second, the top of Roraima is relatively well accessible – a touristic ac­ti­vi­ty
could facilitate the access for coatis and moreover, the climb on the top should be no problem
for such agile animals. It could be interesting that both our observations happened in the end of
the dry season. At least four other expeditions in different seasons did not observe any coatis.
It could imply some seasonal distribution of coatis on the top of Roraima (related to possible
seasonal specific behavior – e.g., higher male’s exploration activity for females, higher density
of individuals in forest, decrease of food sources in surrounding areas etc.).
In conclusion, we have reported a rare observation of the omnivorous opportunistic Brown-nosed coati on Roraima tepui. The exact number of coatis on Roraima, their detailed feeding
strategy and potential dependence on human visiting activity in such area remain unclear. We
will be deeply indebted for any additional information about coatis (or some other mammals)
on whichever tepui.
SOUHRN
Stolové hory venezuelské Guayany (tzv. tepui) jsou izolovanými horami či pohořími, u kterých jejich
dlou­ho­do­bá samostatnost snížila výskyt řady živočichů a rostlin, ale na druhou stranu přispěla k vy­so­ké­mu
podílu místního endemismu. K velmi vzácným zvířatům vrcholových partií stolových hor patří savci. Náš
příspěvek zmiňuje opakované pozorování poddruhu nosála červeného (Nasua nasua vittata) na stolové
hoře Roraima (hranice Venezuely, Guyany a Brazílie) v letech 2002 a 2003. Výskyt této středně velké
šelmy v tomto relativně nehostinném prostředí spojujeme s všeobecným oportunismem nosálů (ve vztahu
k potravě i bi­oto­pu). V roce 2003 se dokonce podařilo pozorovat agresivní střet dvou jedinců. Vzhledem
k tomuto sledování se zdá, že nejde pouze o zatoulané jednotlivce, ale možná o výskyt více zvířat trvalejšího rázu. Nelze také vyloučit, že přežívání nosála červeného na Roraimě je v určité míře umožněno
také přiživováním na turistech.
ACKNOWLEDGEMENT
Our thanks are due to Václav Mikeš (Faculty of Biological Sciences, University of South Bohemia, České
Budějovice, Czech Republic) for some ornithological information from Roraima tepui and Oldřich Říčan
(Faculty of Biological Sciences, University of South Bohemia) for valuable comments. This work was
partly supported by the grant No. MSMT 6007665801.
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Lynx (Praha), n. s., 37: 131–150 (2006).
ISSN 0024–7774
A new genus of vespertilionid bat from Early Miocene of Jebel Zelten,
Libya, with comments on Scotophilus and early history of vespertilionid
bats (Chiroptera)
Nový rod netopýra z basálního miocenu Džebelu Zelten, Libye, s poznámkami
k rodu Scotophilus a rané historii vespertilionidních netopýrů (Chiroptera)
Ivan Horáček1, Oldřich Fejfar2 & Pavel Hulva1
1
Department of Zoology, Charles University, Viničná 7, CZ–128 44 Praha, Czech Republic;
[email protected], [email protected]
2
Department of Geology and Palaeontology, Charles University, Albertov 6, CZ–128 43 Praha,
Czech Republic; [email protected]
received on 8 December 2006
Abstract. A well preserved mandible of a vespertilionid bat is described from the MN4–5 site Jebel Zelten
MS2, Libya. The bat shows a greatly derived state in most of dental characters, but it differs from the
Recent genera with corresponding degree of dental reduction (Eptesicus, Scotomanes, Hesperoptenus),
in shape of molars and symphyseal region. In certain respects it reminds the Recent Scotophilus and the
Late Paleogene African genus Philisis. A possibility that Philisis and the Jebel Zelten bat, described here
as Scotophilisis libycus gen. nov et sp. nov, form a stem line of Scotophilus is discussed in context with
recent molecular data on position of the genus.
Introduction
The fossil records from a series of sedimentary series in Jebel Zelten in northern Libya (Savage
& Hammilton 1973) represent quite important source of information on development of mammal fauna of the eastern part of North Africa in the Early and Middle Pleistocene, i.e. after the
stage well documented in classical Oligocene sites in Fayum and prior the aridisation during
the Messinian Crisis and the extensive rearrangements related to it. Structure of the mammalian
assemblages from Jebel Zelten and details on particular finding sites were recently surveyed by
Wessels et al. (2003), further data and their palaeobiogeographic implications are in another
paper of this volume (Fejfar & Horáček 2006).
The present contribution is confined to just one group that is particularly rare in the North
African fossil records: bats. Unfortunately it is represented with just a single specimen in Jebel
Zelten collections. Nevertheless, it is for more respect quite curious and intersting. The specimen
belongs undoubtedly to a vespertilionid bat, which, at first sight, must be closely related to extant
genera bearing a greatly advanced dental pattern (such as Scotophilus Leach, 1821 to which
it was tentatively assigned in previous reports by Horáček 2001 and Wessels et al. 2003). Its
actual species and even generic identity is in no way a simple and transparent, of course. We
will demonstrate that in fine details the specimen differs markedly from any yet known genus
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both extant and fossil. This fact and its possible consequences present a non-trivial problem
that is worth a separate analysis.
A bat of Jebel Zelten: results of a study
The specimen. The specimen was found in site MS2 of Jebel Zelten, northern Libya (see Wessels
et al. 2003, Fejfar & Horáček 2006 for details) which fauna dates it to MN4–5, i.e. ca. 14–16
My. The specimen is a well preserved right mandible with M1 and M2 and alveoli of all remaing
teeth. The mandible is figured in Fig. 1, dimensions are (in mm, alveolar in parantheses): (I1–M3):
7.8, (C–M3): 7.2, (P4–M3): 5.8, (M1–M3): 4.6, M1–M3): 5.2, M2–M3): 3.5, M1–M2: 3.55, (I1–I3):
0.81, (I1–P4): 3.1, (C–P4): 2.4, heigth of mandible below M1: 2.4, symphysis 2.7×1.5, M1: length
1.76, trigonid length 0.85, talonid length 0.91, trigonid width 1.06, talonid width 1.20, M2: length
1.85, trigonid length 0.88, talonid length 0.97, trigonid width 1.20, talonid width 1.22.
Description. (1) Dental formula is 3123, (2) unicuspid section is compressed to about 37% of
tooth row and 60% of the molar row length: (3) 3 incisor alveoli minute, all of roughly the same
size and their position suggest considerable overlap between crowns of the respective teeth
(type C in sense of Menu 1985), (4) C alveolus large, nearly rounded (0.86 mm), not axially
compressed, also (5) P3 alveolus is rounded and not compressed or displaced of the row, (6) two
P4 alveoli are of the same size as that of P3 but compressed mesiodistally, (7) alveolar lenght
of P4 (0.93) is almost the same as that of C. (8) molars are myotodont in sense of Menu & Sigé
(1971), i.e. postcristids connect hypoconids and entoconids, hypoconulids stay appart, (9) both
molars are robust with well developed talonids, (10) conspicuously thick wall of trigonids and
shallow trigonid fossids (protofossids), (11) high standing lingual base of protofossid that is at
the same level as the deepest inflexion of entoconid crest, (12) longitudinal distance between
para- and metaconid is equal to the distance between meta- and entoconid or slightly larger (on
M1), (13) hypoconulid is low standing without connection with the cingular band, (14) paraconid
of M1 is somewhat tappered mesially and relatively low in comparison with metaconids, (15)
there is an indistinct but clear undulation of crown base at level of metaconid, (16) alveoli of
M3 are roughly of the same size as in M1 and M2, including the distance between the alveoli, the
distal alveolus of M3 is not reduced, (17) the mandible body is robust and haevily build, (18)
symphysis is conspicuously broad and symetrical, oval-shaped, without a chin thickenning or
mesial tappering, (19) inflexion point of the line of symphysal distal margin is situated below
the centre of P3 and at nearly at a half of height of the mandible body, (20) foramen mentale
is situated below P3, (21) angulus mandibulae is indistinct, the base of ramal section diverges
from the line of of the mandible base by 13° only.
Comparisons. Most of the above surveyed characters are just the apomorphies of vespertilionid
dental organisation, hence, the specimen under study belongs apparently to a clade characterized
by quite advanced degree of dental specialisation. The considerable degree of premolar reduction
excludes the genera bearing the less advanced pattern, first of all Myotis Kaup, 1829 or Keri­
voula Gray, 1842 and ancient vespertilionine genera like Karstala Czaplewski et Morgan, 2000

Fig. 1. The bat of Jebel Zelten: Scotophilisis libycus gen. nov. et sp. nov. – right mandible, lateral, lingual
and occlusal view.
Obr. 1. Netopýr z Džebelu Zelten: Scotophilisis libycus gen. nov. et sp. nov. – pravá mandibula, laterální,
linguální a oklusální pohled.
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133
or Hanakia Horáček, 2001 that in Miocene may come in account, eventually. Also the genera
bearing the plesiomorphic nyctalodont molar pattern (cf. Menu & Sigé 1971) can be excluded:
this concerns among other those to which the specimen in study approach in dental formula and
general morphology, e.g. Scotoecus Thomas, 1901, Scotozous Dobson, 1875, Nyctalus Bowditch,
1825. What remains are the genera characterized by myotodont molars and considerbale degree of
premolar reduction. Excluding those which differ in their small body sizes and/or in their sleder
or less specialized dentitions (Hypsugo Kolenati, 1856, Neoromicia Roberts, 1926, Nycticeinops
Hill et Harrison, 1987, Laephotis Thomas, 1901, Glauconycteris Dobson, 1875, Arielulus Hill et
Harrison, 1987), or those improbable for geographic reasons (Australian Scotorepens Troughton,
1943, Nyctophilus Leach, 1821, Falsistrellus Troughton, 1943, Vespadellus Troughton, 1943,
Neotropic Rhogeessa Allen, 1866, Baeodon Miller, 1906), the list of the extant genera which
come in acount is further reduced onto: Eptesicus Rafinesque, 1820, Scotomanes Dobson, 1875,
Hesperoptenus Peters, 1868, Vespertilio Linnaeus, 1758, Otonycteris Peters, 1859, Antrozous
Allen, 1862, and Scotophilus Leach, 1821. None of them, fit in all the characters, of course. The
fossil form exceeds the limits of variation in most genera particularly in the characters 2, 10, 11,
18, 19, 20, and 22. Considerable differences are in architecture of molars which are distinctly
haevily build. Spacious and deep fossids, narrow and vertically oriented para- and metaconids in
Eptesicus, Hesperoptenus, Scotomanes, Vespertilio, Antrozous, or Otonycteris, clearly contrast
with high massive lingual wall and shallow bases of fossids in M1 and M2 of the fossil form.
Even the most robust forms of these genera differ also in shape and position of symphysis and
degree of reduction of incisor row. We stress these facts especially because just these genera,
especially Eptesicus, were the first hot candidates for defaut identification of the fossil specimen at least for clear resemblance in structure of dentition, shape of mandible and last but not
least for biogeographic reasons. Eptesicus is the genus recently wide-spred and taxonomically
diversified in the respective region. We compared directly the fossil specimen with Eptesicus
serotinus (Schreber, 1774), E. (s.) turcomanus (Eversmann, 1840), E. isabellinus (Temminck,
1840), E. anatolicus Felten, 1971, E. bottae (Peters, 1869), E. nasutus (Dobson, 1877) and E.
nilssonii (Keyserling et Blasius, 1839) (besides of further taxa) and constantly confirmed the
above mentioned differences. Vespertilio exhibits correspondingly high degree of reduction in
incisor row but markedly differ in molar design and the same hold true for Scotomanes ornatus
(Blyth, 1851) and Hesperoptenus tickelli (Blyth, 1851) that well approach the fossil specimen
also in overall size. Otonycteris hemprichii (Peters, 1859) and Samonycteris majori Revilliod,
1922 which would come in account also for biogeographic reasons differ in molar design even
more (comp. Figs. 3, 4).
Compared to the primitive condition in vespertilionids (e.g. in Stehlinia Revilliod, 1919,
Myotis, Hanakia) the fossil form exhibits an apparent increase in relative mass of trigonids. In
contrast to the above mentioned extant genera, all these trends are particularly pronounced in

Fig. 2. Right mandibles of Scotophilisis libycus gen. nov. et sp. nov., Jebel Zelten, leg. O. Fejfar (1ab),
Scotophilus cf. viridis, Khartoum, Sudan, leg. P. Štys (2ab), Scotophilus kuhlii, CAM 40, Pakse, Laos, leg.
I. Horáček (3ab), Eptesicus serotinus, Q2 Chlum4, Bohemia, leg. I. Horáček (4a), and Eptesicus (serotinus)
turcomanus, CT84/19 Frunze, Kirghizia, leg. I. Horáček (5b). a – lateral view, b – occlusal view.
Obr. 2. Pravé mandibuly Scotophilisis libycus gen. nov. et sp. nov., Džebel Zelten, leg. O. Fejfar (1ab),
Scotophilus cf. viridis, Chartúm, Sudan, leg. P. Štys (2ab), Scotophilus kuhlii, CAM 40, Pakse, Laos, leg.
I. Horáček (3ab), Eptesicus serotinus, Q2 Chlum4, Čechy, leg. I. Horáček (4a), a Eptesicus (serotinus)
turcomanus, CT84/19 Frunze, Kirgisie, leg. I. Horáček (5b). a – laterální pohled, b – oklusální pohled.
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135
Scotophilus, the genus which bears the most advanced state of the respective dental characters
of all vespertilionid bats. There is a broad measure of agreement between the Jebel Zelten bat
and smaller species of Scotophilus [viridis (Peters, 1852), kuhlii Leach, 1821, and partly dingani
(Smith, 1833) and leucogaster (Cretzschmar, 1830)] in overal size and proportions of mandible
(including relative height of mandible body, shape and form of symphysis and/or in indistinct
angulus mandibulae) as well as in proportions of molar to unicuspid row or a degree of reduction of incisors and premolars contrasting to quite a large canine. In general dimensions, the
fossil form falls in variation range of S. viridis. Nevertheless, all the examined extant species
of Scotophilus clearly differ in the diagnostic character of the genus: considerable degree of
reduction of talonids, which are compressed mesio-distally and low staying in respect to the
lingual basis of a tooth. The larger forms of the genus, e.g. S. heathi (Horsfield, 1831), reach
even higher degree of that trend. These differences seem to exclude a direct coidentification
of the Jebel Zelten bat with Scotophilus, though it is the extant genus which shows the most
similarities with it in comparison to other extant genera.
Genus Scotophilus is also reported from middle Miocene of Central Europe by Engesser
(1972): a well preserved mandible from MN7 Steinheim and 8 isolated teeth from MN8 Anwil
(including 3 M1 with length: 2.08–2.12, and 3 M2: 2.20–2.24) were found to correspond both
in size and in shape of the particular teeth (C1, M1, M2) to the situation in extant Scotophilus
temminckii (Horsfield, 1824) (= S. kuhlii Leach, 1821 – comp. Simmons 2005). Unfortunately,
the respective items were not figured in details and until now, we did not succeed either to
examine them or to compare them and the specimen in question directly. In any case, similarly
as with extant Scotophilus kuhlii, their measurements are clearly larger than those in the Jebel
Zelten bat. In comparison to the extant species, the Steinheim specimen (judging from Engesser
1972: 129, Fig. 37) exhibits a lesser degree of canine and premolar mesio-distal compression
and fairly less reduced talonids of M1 and M2. In these respects it might remind the Jebel Zelten
bat eventually – a direct comparison is neccessary, of course.
The other group that is to be taken in account is a mysterious African clade of early vespertilionids, family Philisidae Sigé, 1985. The family and its type species, Philisis sphingis
Sigé, 1985, were described based on two jaw fragments (P4–M3, (P2–)P4–M2) obtained during
1961–1962 excavation in famous early Oligocene site of Jebel El Quatrani in Fayum, Egypt.
The detailed analysis by Sigé (1985) can be for purpose of the present comparisons tentatively
summarized as follows: the remains belonged to a very large bat (P4–M3 9.29, M1–M2 6.08
mm) exhibiting a striking differences from any genus then available in its combination of the
following characters: (a) a broad basis of orbit with lateral extension of zygomata at level of
M1–M2, (b) oval shaped and vertically oriented foramen infraorbitale at level of P4, (c) robust

Fig. 3. M1,2 dental morphology in Scotophilisis libycus gen. nov. et sp. nov., Jebel Zelten, leg. O. Fejfar
(1ab), Scotophilus cf. viridis, Khartoum, Sudan, leg. P. Štys (2b), Scotophilus kuhlii, CAM 40, Pakse,
Laos, leg. I. Horáček (3ab), Eptesicus anatolicus, NMP 48192, Bavineh, Iran, leg. P. Benda (4a), Eptesicus
(serotinus) turcomanus, CT84/19, Frunze, Kirghizia, leg. I. Horáček (5ab), and Otonycteris hemprichii,
E40, Egypt. a – lateral view, b – occlusal view. Not in a scale.
Obr. 3. Morfologie M1 a M2 u Scotophilisis libycus gen. nov. et sp. nov., Džebel Zelten, leg. O. Fejfar
(1ab), Scotophilus cf. viridis, Chartúm, Sudan, leg. P. Štys (2b), Scotophilus kuhlii, CAM 40, Pakse, Laos,
leg. I. Horáček (3ab), Eptesicus anatolicus, NMP 48192, Bavineh, Iran, leg. P. Benda (4a), Eptesicus
(serotinus) turcomanus, CT84/19, Frunze, Kirgisie, leg. I. Horáček (5ab), and Otonycteris hemprichii,
E40, Egypt. a – lateralní pohled, b – oklusální pohled. Upraveno na stejnou velikost.
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but mesiodistally shortened upper molariforms (P4–M3), (d) very high inner wall of upper molars with postprotocrista terminating with a sharp sweep at a hypocone region, (e) shalow fossa
without para- or metalophi, (f) split of mesostyle into mutually separated lateral terminations of
inner crests of para- and metacone at M1 and M2 but (g) retaining complete plagiocrista at M3
that connects parastyle, paracone, mesostyle and metacone that is preserved at M3 despite the
tooth is moderately reduced, (h) a short but broad P4 with robust mesiopalatal thickening and
distinct inflexion of the distal crown margin, (i) a robust mandible body, (j) three premolars with
the middle one reduced and displaced lingually, (k) haevily build lower molars with (l) high
lingual walls, (m) relatively low paraconid and shalow protofossid, (n) indistinct undulation of
lingual base of the teeth at level of paraconid, (o) a distinct but low standing hypoconulide, (p)
hypocone is relatively lower in M2 than in M1, (r) distal alveolus of M3 not reduced, alveolar
length of M3 seems to be nearly equal to that of M1. Except for (h), the mandibular characters
of Philisis sphingis resemble clearly the situation demonstrated in the Jebel Zelten bat except
for its size which is nearly twice larger that in the latter form.
Further data concerning Philisidae arose of the analyses of bat remains from the Tunisian
early Eocene fauna in Chambi (for its stratigraphical setting see Hartenberger et al. 1997):
Sigé (1991) demonstrated that at least two of four bats remains found there (M1, M1–M2) fit
well the diagnostic characters of the family (in general architecture of molars, particularly at
M1) but differ from Philisis sphingis in its very small size (length of M1: 1.23, width of M1:
1.71, length of M2: 1.12 ), and, in particular, nyctalodont condition of the lower molars. The
form described as Dizzya exsultans Sigé, 1991 further differs in surprisingly high degree of
talonid reduction at M2 contrasting to broad and shalow talonid at M1. Alveoli of M3 suggests
a considereble degree of reduction, too. Thus, in comparison to Philisis sphingis, Dizzya exsul­
tans shows a combination of both primitive characters (nyctalodoncy, small size) and greatly
advanced characters (reduction od M2 and M3) for which it cannot be regarded a direct ancestor
of the former one.
In respect to the present comparisons, the most pertinent data are those by Sigé et al. (1994)
reporting further two forms of Philisidae from the early Oligocene site Taqah from Oman.
Philisis sevketi Sigé, 1994 described from there based on isolated M1 and seven other teeth
resembles our specimen in available characters (comp. Fig 18 in Sigé et al. 1994) but differs
in comparativelly larger size (M1 length: 2.28, talonid width: 1.91). Besides it, of course, Sigé
et al. (1994) reports from the site also two incomplete fragments of upper molars (M3 and M1)
identified as “cf. Philisis sp.” which are of a smaller size roughly corresponding to that of our
specimen (length of M3 ca. 1.14, length of M1 ca. 1.48).
Summing up the above comparisons we can conclude: (i) the Jebel Zelten bat differs clearly from
the extant genera which come here in account mostly for specificities in its molar architecture
and for its robust mandible, its symphyseal region, and a considerable degre of reduction of
premolar and incisor rows. (ii) To a considerable degree these specificities are shared with extant
genus Scotophilus. Nevertheless, all of the six species examined essentially differ in degree
of reduction and positional rearrangements of talonids in which they exceed corresponding
tendencies in any other vespertilionid genus including Philisis and/or the Jebel Zelten bat. (iii)
The specimen in study shows a broad measure of agreement with the Early Eocene to Early
Oligocene Afro-Arabian genus Philisis, particularly in shape and general architecture of M1
and M2. It markedly differ from Philisis by its smaller size and a reduced premolar row (P3
absents at all, P4 compressed).
138
Fig. 4. Type specimen of Samonycteris majori Revilliod, 1922 (Mytilini, Samos, Greece), Mus. Lausanne
945(S)/21908. Above – lateral view of the specimen, middle left – cochlear region (comp. also Horáček
1991), others – details of dentition.
Obr. 4. Typový exemplář Samonycteris majori Revilliod, 1922 (Mytilini, ostrov Samos, Řecko), Mus.
Lausanne 945(S)/21908. Nahoře – laterální pohled na exemplář, střed vlevo – kochleární oblast (viz také
Horáček 1991), ostatní – detaily dentice.
139
Taxonomy
For the above mentioned reasons, the Jebel Zelten bat cannot be coidentified with any of the
yet described genera both extant and fossil. At the same time, it is clear, of course, that a more
robust conclusions on its actual affinities could be drawn only after a more complete material
will be available, particularly after it would support the above predictions with at least some
of the upper jaw characters. Thus, it would be perhaps the most correct to stop our treatment at
this point and wait for supplementing of the material with further items. Nevertheless, finally
we decided to express the results of the above comparisons in form of a taxonomic opinion not
only for apparent improbability of the expected record, but also as a way to emphasize the non
trivial aspects of the matter in order to stimulate its further reexamination. Since, the specimen
cannot be arranged under any yet described genus, its proper taxonomic treatment necessitates
to propose a new genus for it.
Scotophilisis gen. nov.
Type Species. Scotophilisis libycus sp. nov.
Derivatio nominis. Supposedely related to stem line of the extant genus Scotophilus Leach,
1821, and at the same time to a Palaeogene genus Phylisis.
Diagnosis. A derived member of Phylisidae Sigé, 1985, sharing the characters of M1 and M2
architecture with Philisis sphingis Sigé, 1985, particularly the heavily build protoconide complex,
high lingual crown wall and shalow protofossid of the respective teeth. It differs from Philisis
by a reduced premolar row consisting of P2 and P4 only. It is further characterized by three small
incisors, large canine, robust and broadly oval symphysis not extended behind a level of P2,
and indistinct angulus mandibulae. It resembles Scotophilus in the latter characters but differs
from it by a lesser degree of reduction and specific rearrangement of talonids, diagnostic for
the extant genus (Miller 1907, Koopman 1994). From eptesicoid genera Eptesicus, Hespero­
ptenus or Scotomanes which share the corresponding degree of reduction of unicuspid row, it
differs mainly in the molar characters shared with Philisis, in shape of symphyseal region, more
advanced degree of compresion of incisor row, and smaller angle of angulus mandibulae.
Scotophilisis libycus sp. nov.
Holotype. Right mandible with M1 and M2, alveoli of I1 to P4 and of M3, well preserved mandible body including the symphyseal region and anterior part of ramus mandibulae, deposited
in collections of National Museum Praha under number NMPC/OF.J.Zel./Chi1.
Diagnosis. Same as diagnosis of the genus.
Type locality and stratum. Jebel Zelten, MS2, Northern Libya, 28° 28’ N, 20° 00’E, Early
Miocene, MN4–5 (see Wessels et al. 2003 and Fejfar & Horáček 2006 for details).
Details. For measurements, description and comparisons see above.
Discussion
Vespertilionid bats represent a group characterised by a generalised state of dental and cranial
characters and a prolonged maintenance of the ancestral states in most of them. The largest
genus of the family, Myotis, and several other genera bear the complete dentition with unreduced number of particular tooth types while only few genera reach the level of advanced
dental rearrangements at the extent common in other chiropteran families. Simply said, dental
140
evolution in vespertilionid bats was mostly restricted to clade-specific variation in relative size
of particular dental elements, typically including enlargements of the molariform sector and/or
canines, reductions of relative size of individual incisors or premolars, their displacements from a
toothrow up to their disapperance in several clades, eventually, often accompanied with reduction
of distal elements at M3/3 and relative enlargements of rostral part of skull. Worth mentioning is
that until middle Miocene a vast majority of vespertilionid fossil record consisted of the forms
exhibiting the ancestral dental pattern corresponding to the state in extant genus Myotis though
– as demonstrated elsewhere (Horáček 2001) – they most probably belonged to the clades which
extant members are characterized by rather derived dental pattern (Eptesicus, Vespertilio, a. o.).
Parallel appearance of the same trends in dental rearrangements (reduction and mesio-distal
compression of premolar and incisor rows, reduction of distal elements of M3/3) seems to be
perhaps the most characteristic phenomenon in Late Caenozoic history of the family. Despite such
characters are undoubtedly relevant (and in most instances reliable) for distinguishing of extant
vespertilinid genera, for phylogenetic interpretations of the Tertiary record of the family are of
a limited significance only. Nevertheless, under presumption that the respective morphoclines
are irreversible, an appearance of a derived state exceeding the maximum degree observed in
a Recent taxon, to which the fossil item could be assigned, can be used as an indirect argument
against such decision. Such a sylogism was applied also in the above comparisons.
In contrast to the above mentioned traits, the form of molars and molar design (except for
M3/3, of course) represent in vespertilionid bats an extremelly conservative character which
variation operates within very narrow limits characterizing particular genera and/or suprageneric clades. It provides perhaps the most reliable information for generic assignment and
morphological comparisons at suprageneric level (Menu 1985). As pointed out by Menu (o.c.),
in vespertilionids, the most primitive conditions in molar characters are in Myotis (which at
the same time exhibits relativelly large intrageneric variation), while far the most derived state
appears in Scotophilus.
Scotophilus: a neontologic perspective
The genus Scotophilus Leach, 1821 is distributed throughout sub-Saharan Africa, in southern
Arabia, Madagascar and Reunion (Koopman 1984, 1994, Robbins et al. 1985, Gaucher 1993,
Simmons 2005), and throughout considerable part of southern and southeast Asia including
Phillipines, Borneo and Celebes (Corbet & Hill 1992, Bates & Harrison 1997, Kitchener
at al. 1997).
All species included in the genus are nearly uniform in their cranial, dental and external characters while, at the same time, in almost all dental and cranial characters they exhibit the most
derived state of all vespertilionid bats (Menu 1987). This separates the genus quite distinctly
from the other clades but, simultaneously, it provides almost no chance to comprehend its true
affinities. The genus represents a taxonomic puzzle in all classifiations and numerous confusions
accompanied also further aspects of its study.
The genus Scotophilus Leach, 1821 was described based on a juvenile individual bearing
milk teeth what subsequently caused numerous confusions on actual meaning and content of
genus. The name was later applied nearly to all vespertilionids with a derived dental pattern
(comp. Hoofer et al. 2006) until its current meaning was established by Miller (1907), who,
at the same time, of course, suggested a homonymy of the name Scotophilus Leach, 1821 and
Scotophila Hübner, 1816 (Lepidoptera) and denoted the genus with the name Pachyotus Gray,
141
1831. Tate (1942) proposed to include Scotophilus together with Scotomanes, Stotoecus, Scotei­
nus, Nycticeius, Rogeessa, Baeodon and Otonycteris (i.e. those sharing the most derived dental
characters) into a separate tribe Nycticeini diagnosed among other by apomorphic condition in
upper incisors with (“the outer incisor is obsolete, the inner one usually lacks the supplementary
cusp and is simply conical”).
Hill & Harrison (1987) suggested that Nycticeini as assembled by Tate (1942) is not
a natural group and reports that Scotophilus and Scotomanes have bacula reminiscent of the
flattened triangular structure of Eptesicus and its immediate associates. Because of derived
dental and cranial characters they placed Scotophilus and Scotomanes in a separate tribe Scotophilini, while Otonycteris, Nycticeius, Rhogeessa and Baedon placed in a tribe Plecotini.
Despite that Koopman (1994) in regard to his former proposals on reality of the tribe Nycticeini retained the traditional concept and stressed relations of Scotophilus to Scotomanes and
Otonycteris.
Results of molecular analyses by Hoofer & Van Den Bussche (2003) strongly contradicts
any close association between Scotomanes and Scotophilus. The former genus was found to be
closely related to Eptesicus, while Scotophilus is, in its mtDNA, the most derived genus within
Vespertilioninae (Hoofer & Van Den Bussche 2003: 28). Our morphological comparisons
supports this conclusion quite a well: Scotomanes lacks the derived characters of Scotophilus
on upper molars (C type molars in sense of Menu 1985) and also its lower molars exhibit the
characters of Eptesicus (low lingual wall of trigonid, deep and sharply bordered protofossid).
In respect to their results, Hoofer & Van Den Bussche (2003) assigned Scotophilus to its own
tribe Scotophilini and expressed serious doubt on its relationship to other vespertilionid clades
(commented as “sedis mutabilis”). The detailed karyological comparisons undertaken by Vol­
leth et al. (2006) convincingly support the same view, too.
The phylogenetic analyses perfomed with cyt b mtDNA sequences of major vespertilinid
genera (Fig. 5, Appendix) suggests for Scotophilus a position at basal split of the vespertilionid
stem group (worth mentioning that it immediately follows that of Miniopterus Bonaparte, 1837
and Cistugo Thomas, 1912, whose African origin is quite probable too.
The genus is uniform in apomorphic condition of its dental and cranial specificities but greatly
variegated in other respects (body size, pelage colouration, age and sex variation etc.). Its members can be quite easily distinguished from other vespertilionid bats but taxonomic structure of
the genus, status of particular forms and their variation and phylogenetic relationships present
traditionally quite confused, controversial and largerly unresolved topics.
Koopman (1984, 1994) who first revised the topics in detail suggested an extensive synonymization over the many named forms and reduced the content of the genus onto 5 phenotypically
well defined species (three African: borbonicus (Geoffroy, 1803), leucogaster, nigritta (Schreber,
1774), and two Asian: kuhlii, heathii).

Fig. 5. Bayesian tree based on complete 1140 bp sequence of cytochrome b showing relationships among
extant vespertilionid genera. Note basal position of Scotophilus. For details see Appendix.
Obr. 5. Fylogenetický strom vybraných zástupců čeledi Vespertilionidae spočtený na základě sekvencí
kompletního genu pro cytochrom b (technika Bayesianské analysy). Povšimněte si basální posice rodu
Scotophilus. Detaily analysy jsou v appendixu.
142
143
The most detailed morphometric analysis on the African Scotophilus was undertaken by Robbins
et al. (1985), who examined over 2000 specimens from throughout sub-Saharan Africa including
11 type specimens. They recognized six species of Scotophilus occurring in sub-Saharan Africa: S.
dinganii, S. leucogaster, S. nigrita, S. nucella Robbins, 1983, S. nux Thomas, 1904, and S. viridis.
Kitchener et al. (1997) revised the taxon in Indo-Malayan region and demonstrated a distinct
species status of S. celebensis Sody, 1928 and S. collinus Sody, 1936 apart from widespread
Oriental species S. kuhlii. With these additions, the results by Robbins et al. (1985) have been
accepted in the most recent account of the genus by Simmons (2005) who recognized 12 species
within it: S. borbonicus (Geoffroy, 1803); S. celebensis Sody, 1928; S. collinus Sody, 1936; S.
dinganii (Smith, 1833); S. heathi (Horsfield, 1831); S. kuhlii Leach, 1821; S. leucogaster (Cretzschmar, 1830); S. nigrita (Schreber, 1774); S. nucella Robbins, 1983; S. nux Thomas, 1904;
S. robustus Milne-Edwards, 1881; and S. viridis (Peters, 1852). The list that should be further
completed with S. tanrefana Goodman, Jenkins et Ratrimomanarivo, 2005.
The phylogenetic relations among particular species of the genus present another topic quite
confused. Koopman (1984) and Robbins et al. (1985) proposed different phenetic groupings for
the African taxa while their relations to the Asian species has not been discussed.
Hoofer & Van Den Bussche (2003) based on molecular analysis of three mtDNA genes
suggested a close relationship among the Ethiopian species but a distant relationship between
the two Oriental species (S. heathi and S. kuhlii).
The topics was recently reexamined into great details by Trujillo (2005) who analyzed 138
specimens of 10 spp. with aid od large segments of mtDNA and the Y-chromosome zfy gene.
His results represent undoubtedly the most comprehensive recent contribution to taxonomy and
phylogeography of the genus. He confirmed the above mentioned conclusions by Hoofer & Van
Den Bussche (2003), provided evidence for two additional cryptic species within S. viridis and
S.dinganii (by which the number of species incereased to 16 species) and further draw a detailed
view on phylogenetic structure of the genus and its phylogeographic specificities. The study
demonstrated S. kuhlii as the most basal taxon, monophyly of African Scotophilus with S. nux
as the most basal African taxon, and a considerable distance between the two Indomalayan
species suggesting multiple origins of Asian Scotophilus (S. heathi represents an inner clade of
the African radiation related to viridis-dinganii group). It was also demonstrated that Malagasy
Scotophilus appear to have affinities to different African species and likely originated from
multiple colonization events from Africa. Further cryptic species was recently described from
Madagascar (Goodman et al. 2006) and also the most recent studies on African Scotophilus (Ja­
cobs et al. 2006) demonstrated appearance of further cryptic species within the south African S.
dinganii group. All that suggest that the genus may produce a considerable cladogenetic activity
without marked effects on the morphometric characters of the once stabilized phenotypes.
Summing up the neontologic data on the genus, it seems that: Scotophilus is a product of a very
early divergence from ancestor stock of other vespertilionid clades (tribes), it is apparently of
the Palaeotropic origin and its major radiation occured in Africa from where it colonized either
Madagascar and neighbouring islands (Reunion with S. bourbonicus) and the Indian region (the
most recently with S. heathii). In these connection it should be reminded that Scotophilus bats
rank among the top vespertilionid specialist in aerial fast hawking what dispose them to distant
migrations quite a well. Despite that they respresent a very common element of bat communities
both in Sub-Saharan Africa and in Oriental region, they nowhere (except for the southernmost
Arabia, e.g. Gaucher 1993) crossed the southern border of the Palaearctic region (Horáček et
al. 2000) and entered the temperate zone.
144
Scotophilus and Philisidae: a stem line?
The above surveyed neontologic evidences suggest that Scotophilus originated by early divergence from ancestral vespertilionids and its major radiation is confined to Africa. Unfortunately,
as to our knowledge, any fossil record which would support these statements absents except
for those from the middle Miocene of central Europe which are extralimital and should be
reexamined with particular care. Nevertheless, the mandible from MN6 Steinheim figured by
Engesser (1972: 129) resembles the extant forms of the genus quite a much, perhaps except
for less reduced talonids. In contrast, M2 from MN8 Anwil, figured by Engesser (1972: 130)
exhibits a well developed mesostyle at a buccal position instead of the deep inflexion characteristic for extant Scotophilus. If the generic affiliation of these items, suggested by Engesser
(o.c.) is correct then they would suggest that as late as in the late middle Miocene the dental
specificities of the genus were not attained completely, what, at the first sight, contradicts the
conclusion on early divergence of the genus. The Jebel Zelten bat supports that view too (at
least if the prediction of its relations to Scotophilus is actually correct, of course). It shares with
the extant genus derived characters of incisor and premolar reduction, robustness of the mandible and symphyseal region and massive body of molars with high lingual walls and shallow
fossids. Nevertheless, the major dental apomorphy of the extant genus – extensive reduction
of talonids and their shift to the base of a tooth is here apparently not attained. Alternatively,
this fact migh fit to a theoretical expectation that the most derived character states appear just
at the latest stage of phylogenetic morphocline of the respective characters, at least in a case
of the characters with conservative variation dynamics as in the molar shape in bats. Further
extension of the same sylogism would result in a prediction that the ancestor of Scotophilisis­
-Scotophilus clade should share with it the essential specificities such as robustness of molars
and mandible body but exhibit an ancestral stage of the common morphocline, i.e. unreduced
number of premolars, incompressed incisors etc. Exactly this characterize the mandibular pattern
in Philisis in which the robustness of the respective structures is moreover related to extremely
large overall size. Philisidae reported as endemic African Late Paleogene group (Sigé 1985,
1991, 1994) would perfectly fit both the predictions on ancestry of the clade in question: an
early divergence and the African origin.
It seems that also the other specificities of Philisis can well be explained in the same way: split
of the mesostyle of upper molars can well be looked upon as the state preceding the complete
reduction of that structure and deep undulation of buccal crown wall at its position, characteristic
of Scotophilus, similarly as the broad base of orbit with conspicuous zygomatic extension of
maxilla, reported for Philisis by Sigé (1985) is partly retained in Scotophilus where it is restricted
from rostral side by excessive enlargement of infraorbital region, apparently connected with
conspicuous enlargement of dorsal structures of skull, the another character in which Scoto­
philus reached the most derived state among all vespertilionid bats. The respective concept of
phylogenetic morphocline of the infraorbital region is further supported by a specific shape of
infraorbital foramen in Scotophilus which is large, fissure-like and vertically tappered.
In conclusion: despite the actual evidence is largerly incomplete and no direct support for the
following statemetnts is available, we hypothesize that Philisidae in sense of Sigé (1985, 1991,
1994) and the newly described genus represent the stem line of Scotophilus and as suggested
by distant position of the extant genus in molecular analyses, the group should be classified as
separate subfamily of Vespertilionidae for which the prior available name is Philisinae Sigé,
1985. Under such a view, proposed content of the clade would be then:
145
Vespertilionidae Gray, 1821
Philisinae Sigé, 1985 stat. nov.
†Dizzya Sigé, 1991 (Eocene, Africa)
†Philisis Sigé, 1985 (Oligocene, Africa)
†Scotophilisis gen. nov. (Miocene, Africa)
Scotophilus Leach, 1821 (middle Miocene, Europe – Recent, Africa, S and SE Asia, Madagascar,
Reunion).
Palaeobiogeographic notes
The above surveyed alternative view of relationship among the genera in question suggests
also several non-trivial palaeobiogeographic hypotheses that are worth of a brief comment. To
give a “safe” background to them first we have to repeat the facts which seem to be invariant
in the present respects: (a) the deepest divergence within the Recent Scotophilus is between the
Asian S. kuhlii group and diversified groups of the African clade including the Asian S. heathi,
(b) the former Asian taxon (S. kuhlii group) seems to be more primitive both in morphological
and molecular respects, (c) extensive radiation of the genus in Africa may suggest the African
origin of the genus, (d) the only records beyond limits of its Recent range (i.e. Palaeotropics)
are from MN7–8 in central Europe, i.e. from a retreat period of the Miocene Climatic Optimum
(Böhme 2003), some 12–13 My, (e) the mtDNA distance between S. kuhlii and the African
species is 16–22% (Trujillo 2005) what indicates the divergence time of some 10–12 My,
according to the most widely used calibration for animal mtDNA (Brown et al. 1979) – about
two percent sequence divegence between pairs of lineages per million years or one percent
divergence per lineage per million years, (f) Philisis and related taxa exhibit even at the early
Oligocene much higher degree of dental specialisation than any vespertilionid clade known
from the Early and Middle Miocene. This holds true also for the Early Miocene Jebel Zelten
bat. (g) The Asian and/or North Tethyan elements in Jebel Zelten fauna (Muridae a.o.) suggest
closing of the Tethyan seaway in the eastern Mediterranean and Iranian region (comp. Rögl
1999, Hrbek & Mayer 2003) as well as expansion of open habitats and onset of aridisation of
the Eastern Mediterranean in the early Miocene (Tschernov 1992).
It cannot be excluded that both these factors (immigration of North Tethyan elements and
spread of open ground habitats) contributed to extensive phenotypical rearrangement in the
above discussed African vespertilionid clade and finally established the apomorphic design of
Scotophilus and its subsequent spread to the North Tethyan and Asian range. Both the molecular
clock estimate and the above mentioned records from Central Europe suggest dating of that
event to about 12 My, i.e. some 4 My after the situation illustrated by the Jebel Zelten bat. Worth
mentioning is that the supposed transition from the stem line to the crown taxon (Scotophilus)
and spread of its range is situated in the period when this most probably occurred also in other
vespertilionid clades such as Eptesicus, Hypsugo etc. (Sigé & Legendre 1983, Ziegler 2000,
Horáček 2001), i.e. in the peak of the Miocene Climatic Optimum (Böhme 2003, Zachos et
al. 2001).
Souhrn
Dobře zachovaná spodní čelist vespertilioidního letouna je popsána z naleziště MS2 z Džebelu Zelten v Libyi. Nález vykazuje vysoce odvozenou vývojovou úroveň ve většině dentálních znaků, ale liší se zároveň
od současných rodů s odpovídajícím stupněm redukce zubů (Eptesicus, Scotomanes, Hesperoptenus), a to
v utváření molárů a symfysy. V určitém ohledu připomíná současný rod Scotophilus a svrchnopaleogénní
146
africký rod Philisis. V kontextu výkladů molekulárních analys je v článku diskutována možnost, že formy
Scotophilus, Phylisis a letoun z Džebelu Zelten, popsaný zde jako Scotophilisis libycus gen. nov. et sp.
nov. vytvářejí spoečnou vývojovou linii.
Acknowledgements
We wish to thank to all collegues who kindly enabled us to study the comparative materials under their
care: Petr Benda & Vladimír Hanák (Praha), Jiří Gaisler & Jan Zima (Brno), John Edwards Hill (London),
Dieter Kock & Gerhard Storch (Frankfurt am Main), Renate Angermann and Robert Asher (Berlin), Kurt
Bauer & Friederike Spizenberger (Wien), and Burkhard Engesser (Basel). Petr Benda kindly discussed
previous version of the manuscript. Special thanks go to Bernard Sigé, a leading personality in study of
fossil bats, who kindly discussed many details of the topics though he is in no way responsible for the
opinions presented here. The study was supported by grant GAČR 206/05/2334 and COST B23.3 (IH).
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Appendix
Molecular phylogenetics
Sequences were aligned by eye since there were no ambiguities in coding sequences of cytochrome b.
Phylogenetic analyses were performed in PAUP* 4.0b2 (Swofford 1998). The model of sequence evolution was inferred using Modeltest 3.06 (Posada & Crandall 1998) and used for computing of Bayesian
tree. Bayesian analysis was chosen because of its ability of processing large number of taxa with complex
model of sequence evolution, effective exploration of posterior probability landscape and straightforward
interpretation of Bayesian probabilities (see e.g. Murphy et al. 2001). Bayesian analysis was performed in
MrBayes (Huelsenbeck & Ronquist 2001) using MCMC analysis with one cold and three incrementally
heated chains with length of 1.000,000 replicates and under the GTR+I+G model of DNA substitution
with R-matrix = (0.2397, 13.7245, 0.3104, 0.5302, 9.9049, 1.0000), base frequencies = (0.3922, 0.3018,
0.0562, 0.2498), gamma shape parameter = 0.484 and burnin = 10,000 based on empirical evaluation.
List of taxa included in the analysis and GenBank accession numbers of respective sequences (in alphabetical order): Chalinolobus tuberculatus (Forster, 1844): AF321051; Cistugo seabrae (Thomas, 1912)
1: AJ841962; Cistugo seabrae 2: AY485685; Cistugo lesueuri Roberts, 1919: AY485687; Eptesicus
diminutus Osgood, 1915: AF376833; Eptesicus fuscus (Beauvois, 1796): AF376835; Eptesicus hotten­
totus (Smith, 1833): AJ841963; Eptesicus nilssonii (Keyserling et Blasius, 1839): AF376836; Eptesicus
serotinus (Schreber, 1774): AF376837; Harpiocephalus mordax Thomas, 1923: AJ841971; Hypsugo savii
(Bonaparte, 1837): AJ504450; Kerivoula papillosa (Temminck, 1840) 1: AJ841969; Kerivoula papillosa 2:
AJ841970; Laephotis wintoni Thomas, 1901: AJ841964; Lasiurus Gray, 1831 sp.: AF376838; Miniopterus
fraterculus Thomas et Schwann, 1906: AJ841975; Miniopterus fuliginosus Hodgson, 1835: AB085735;
Miniopterus natalensis (Smith, 1834): AJ841977; Miniopterus schreibersii (Kuhl, 1817): AY208139;
Murina cyclotis Dobson, 1872: AJ841972; Murina leucogaster Milne-Edwards, 1872: AB085733; Myotis
albescens (Geoffroy, 1806): AF376839; Myotis annectans (Dobson, 1871): AJ841956; Myotis bechsteinii
(Kuhl, 1817): AF376843; Myotis blythii (Tomes, 1857): AF376842; Myotis brandtii (Eversmann, 1845):
AF376844; Myotis capaccinii (Bonaparte, 1837): AF376845; Myotis chinensis (Tomes, 1857): AB106588;
Myotis dasycneme (Boie, 1825): AF376846; Myotis dominicensis Miller, 1902: AF376848; Myotis emar­
ginatus (Geoffroy, 1806): AF376849; Myotis formosus (Hodgson, 1835): AB106592; Myotis frater Allen,
1923: AB106593; Myotis hasseltii (Temminck, 1840): AF376850; Myotis horsfieldii (Temminck, 1840):
149
AF376851; Myotis ikonnikovi Ognev, 1912: AB106603; Myotis keaysi Allen, 1914: AF376852; Myotis
levis (Geoffroy, 1824): AF376853; Myotis lucifugus (Le Conte, 1831): AF376854; Myotis macrodactylus
(Temminck, 1840): AB085736; Myotis macrotarsus (Waterhouse, 1845): AF376856; Myotis montivagus
(Dobson, 1874): AF376858; Myotis muricola (Gray, 1846): AF376859; Myotis myotis (Borkhausen,
1797): AF376860; Myotis mystacinus (Kuhl, 1817): AF376861; Myotis nattereri (Kuhl, 1817): AF376863;
Myotis nigricans (Schinz, 1821): AF376864; Myotis oxyotus (Peters, 1867): AF376865; Myotis petax Hollister, 1912: AB106590; Myotis pruinosus Yoshiyuki, 1971: AB085737; Myotis ricketti (Thomas, 1894):
AB106608; Myotis riparius Handley, 1960: AF376866; Myotis ruber (Geoffroy, 1806): AF376867; Myotis
sicarius Thomas, 1915: AJ841951; Myotis thysanodes Miller, 1897: AF376869; Myotis velifer (Allen,
1890): AF376870; Myotis volans (Allen, 1866): AF376872; Myotis welwitschii (Gray, 1866): AF376874;
Myotis yanbarensis Maeda et Matsumura, 1998: AB106610; Myotis yumanensis (Allen, 1864): AF376875;
Neoromicia capensis (Smith, 1829): AJ841965; Nyctalus leisleri (Kuhl, 1817): AF376832; Nyctalus noctula
(Schreber, 1774): AJ841967; Perimyotis subflavus (Cuvier, 1832): AJ504449; Pipistrellus abramus (Temminck, 1838): AB085739; Pipistrellus hesperidus (Temminck, 1840): AJ841968; Pipistrellus kuhlii (Kuhl,
1817): AJ504444; Pipistrellus nathusii (Keyserling et Blasius, 1839): AJ504446; Pipistrellus pipistrellus
(Schreber, 1774): AJ504443; Pipistrellus pygmaeus (Leach, 1825): AJ504441; Plecotus auritus (Linnaeus,
1758): AB085734; Scotophilus heathi (Horsfield, 1831): AF376831; Vespertilio murinus (Linnaeus, 1758):
AF376834; Vespertilio sinensis (Peters, 1880): AB085738.
150
Lynx (Praha), n. s., 37: 151–159 (2006).
ISSN 0024–7774
Bats of the middle and upper Kızılırmak regions, Central Anatolia, Turkey
(Chiroptera)
Netopýři oblasti středního a horního Kizilirmaku, střední Anatolie, Turecko
(Chiroptera)
Ahmet Karataş1 & Mustafa Sözen2
1
Department of Biology, Faculty of Science-Arts, Niğde University, TR–51100 Niğde, Turkey;
[email protected]
2
Department of Biology, Faculty of Science-Arts, Zonguldak Karaelmas Uni­ver­si­ty;
TR–67100 Incivez-Zonguldak, Turkey; [email protected]
received on 11 July 2006
Abstract. In this study, we found 15 bat species in the middle and upper Kızılırmak regions of central
Anatolia. Four of these species were new for this region, while four previously recorded species were not
found. Our new results raise the total number of species recorded in these subregions to 19. The taxonomic
position of Plecotus bats, which is under discussion recently, was evaluated and P. teneriffae was recorded
under its correct name for the first time in this region. Among the provinces in the area, Niğde and Kayseri
have more bat species than the others by 11 species.
Introduction
The Central Anatolian region, one of seven geographic regions of Turkey, is divided into four
subregions. Two of these, the middle and upper Kızılırmak subregions are situated in the eas­
tern part of the region. There are the following provinces in these subregions: Çankırı, Kay­se­ri,
Kırıkkale, Kırşehir, Nevşehir, Niğde, Sivas, and Yozgat (Fig. 1). The area has a xeric con­ti­nen­tal
climate. Bat species recorded in this region have been reviewed by Benda & Horáček (1998),
who also added some new species for this region. According to Benda & Horáček (1998),
15 bat species have been found in this region. However, individuals of Plecotus au­ri­tus and
P. aus­tri­a­cus recorded in these areas are under discussion and may actually belong to other
species of the genus (see Spitzenberger et al. 2006). This study is a part of a large project on
bat fauna of Turkey.
Material and Methods
In total, 102 specimens were captured using mist net or hand net in the middle and upper Kızılırmak
sub­re­gi­ons of central Anatolia in 1994–2006. 38 individuals were released after measuring. The voucher
samples were prepared in a standard way, the skins, skulls and bacula were deposited in the Mammal
Collection of the Department of Zoology, Niğde University, Turkey (ZDNU).
151
ReCORDS
We recorded altogether 15 bat species in the middle and upper Kızılırmak subregions of central
Anatolia (Table 1).
Rhinolophus ferrumequinum (Schreber, 1774)
Kayseri: İncesu, Kırkinler Cave, 22 May 2005: 1 female (ZDNU 2005/35) (leg. O. Kizilcik); Sarnıcın
Dere vicinity, 20 May 2005: 1 male (ZDNU 2005/36) (leg. O. Kizilcik). – Nevşehir: Avanos, Karakaya
vicinity (ancient rocky settlement), 5 August 1997: 1 sad. female (ZDNU 1997/01), Avanos, between
Sarılar and Özkonak, Hemdionun Cave, 19 September 2001: 3 ad. females (ZDNU 2001/224–226).
– Niğde: Gümüşler, Epçik Cave, 13 October 2001: 1 ad. female (ZDNU 2002/174); near Gebere Dam
Lakelet (1411 m), 21–22 July 2005: 1 ad. female (lact.) (released); 23–24 July 2005: 1 ad. female (lact.)
(ZDNU 2005/66), 1 ad. female (lact.) (released); Ulukışla, Çiftehan, Maden Village, mines, 17 June 2002:
1 ad. male (ZDNU 2002/41). – Sivas: Akıncılar, Deliklitaş Cave, 9 September 2004: 1 ad. male (leg. R.
Bİlgİn & A. Karataş) (ZDNU 2004/346), 1 ind. (obs.); Hâfik, Koşutdere Village (mines), 31 July 2003:
1 ad. ind. (mummy) (ZDNU 2003/52). – Yozgat: Hacıbekir Farm, ruins of Çeşka Castle, 12 April 2002:
1 ad. female (ZDNU 2002/22) (leg. H. C. Öztekİn); Darıcı Village, 17 July 2001: 1 ad. male (ZDNU
2001/81) (leg. H. C. Öztekİn).
Rhinolophus hipposideros (Bechstein, 1800)
Kayseri: Yeşilhisar, Soğanlı, 7 July 2001: 4 ad. females (ZDNU 2001/39–42) (leg. Y. Karakaya). – Niğde: Gümüşler, Epcik Cave, 13 October 2001: 1 ad. male (ZDNU 2002/175); Çamardı, Akpınar Alabalık
Fishfarm, 10 September 2006: 1 sad. male (ZDNU 2006/93), ca. 10 sad. ind. (obs.); Ulukışla, Gümüş
Village (cave), May 1996: 1 ind. (obs.); – Sivas: Hâfik, 3–4 km NW, Yıkılgan vicinity, 10 August 2003:
1 ad. male (ZDNU 2003/51).
Myotis myotis (Borkhausen, 1797)
Kayseri: Kocasinan, Kuşçu, Sarıağıl Cave, 12 July 2001: 1 ad. female (ZDNU 2001/52) (cf. Karataş
et al. 2003); Melikgazi, 2 km NW of Gürpınar, 13 July 2001: 1 ad. male (ZDNU 2002/81); Bünyan,
Karadayı Village, Karatay Hanı Caravanserai, 19 September 2002: 1 ad. male (ZDNU 2002/132); Talas,
Başakpınar Örenyeri vicinity, 13 July 2001: 1 ad. male (ZDNU 2001/65). – Nevşehir: Avanos, between
Sarılar and Özkonak, Hemdionun Cave, 19 September 2001: 1 ad. male, 1 ad. female (ZDNU 2001/227,
228); Gülşe­hir, Açıksaray, Açıksaray Ruins (ca. 1150 m), 28 August 2001: 1 ad. female (ZDNU 2001/147);
Ha­cıbek­taş, Kütükçü Village, 28 August 2001: 1 ad. female (ZDNU 2001/146); Kozaklı, Kaşkışla Village
(cave), 28 Au­gust 2001: 1 ad. male, 1 ad. female (ZDNU 2001/148, 152). – Niğde: Gümüşler, Epcik Cave,
28 May 2001: 1 pregnant female (measured); 29 July 2002: 2 ad. males (ZDNU 2002/103, 110); Uluağaç
Village, Uluağaç Dam Lakelet (artificial cave), 2 May 1996: 1 ad. female (ZDNU 1996/40); Ulukışla,
Öküz Mehmet Paşa Caravanserai, 15 August 1999: 2 ad. males (ZDNU 1999/32–33). – Yoz­gat: Yozgat
(center), 26 May 2002: 1 ad. male (ZDNU 2002/31).
Myotis blythii (Tomes, 1857)
Kayseri: Kocasinan, Kuşçu, Sarıağıl Cave, 12 July 2001: 1 sad. male, 1 ad. female (ZDNU 2001/53,
60) (cf. Karataş et al. 2003). – Niğde: near Gebere Dam Lakelet (1411 m), 21–22 July 2005: 1 ad. male
(ZDNU 2005/61); Aşlama (artificial cave), 21 April 2001: 1 ad. male (ZDNU 2001/17); Çamardı, Çukurbağ
Village, Ece­miş Stream (ca. 1445 m), 14–15 August 2005: 1 male (net. Ş. Özkurt & A. Karataş) (ZDNU
2005/89); Ulukışla, Öküz Mehmet Paşa Caravanserai, 15 August 1999: 1 ad. male (ZDNU 1999/31).
–Yozgat: Şefaatli, Armağan Village, 15 July 2001: 1 ad. male (ZDNU 2001/82) (leg. H. C. Öztekİn).
Myotis aurascens Kusjakin, 1935
Niğde: N. U. Campus, Milli Piyango Hostel, 6 May 2003: 1 ad. female (ZDNU 2003/06).
152
Myotis brandtii (Eversmann, 1845)
Yozgat: Hacıbekir Farm, Çeşka Castle ruins, 12 April 2002: 1 ad. female (ZDNU 2002/23) (leg. H. C.
Öztekİn) (cf. Benda & Karataş 2005).
Eptesicus serotinus (Schreber, 1774)
Niğde: near Gebere Dam Lakelet (1411 m), 23–24 July 2005: 2 ad. males (ZDNU 2005/70, 71); 1 ad.
male, 1 sad. female (released); 12–13 August 2006: 1 ad. male (released).
Fig. 1. Map of the area under study.
Obr. 1. Mapa studovaného území.
Legend / legenda: Kayseri: 1. Bünyan, Karadayı, 2. İncesu, Kırkinler Cave, 3. Sarnıcın­de­re, 4. Kocasinan, Kuşçu, 5. Melikgazi, Gürpınar, 6. Talas, Başakpınar, 7. Yahyalı, 8. Yeşilhisar, Soğanlı; Kı­rık­ka­le:
9. Kırıkkale (centrum); Kırşehir: 10. Kırşehir (centrum); Nevşehir: 11. Nevşehir (centrum), 12. Avanos,
Hemdionun Cave, 13. Derinkuyu, Suvermez, 14. Gülşehir, Açıksaray, 15. Tozköy, 16. Hacıbektaş, Kütükçü,
17. Kozaklı, Kaşkışla, 18. Ürgüp, Sarıhıdır; Niğde: 19. Niğde (centrum), Kayabaşı, 20. Niğde University
Campus, 21. Gebere Dam, 22. Aşlama, 23. Gümüşler Monastery, 24. Gümüşler, Eski Gümüş, 25. Gümüşler,
Epcik Cave, 26. Sazlıca, Akkaya Dam, 27. Uluağaç, 28a. Çamardı, Çukurbağ, 28b. Akpınar Fishfarm, 29.
Ulukışla, 30. Gümüş, 31. Maden, 32. Bolkar mines; Sivas: 33. Akıncılar, 34. Hâfik, Koşutdere, 35. Hâfik,
Yıkılgan, 36. Zâra; Yozgat: 37. Yozgat (centrum), 38. Çeşka Castle, 39. Darıcı, 40. Şefaatli, Armağan.
153
Myotis capaccinii (Bonaparte, 1837)
Kayseri: Kocasinan, Kuşçu, Sarıağıl Cave, 12 July 2001: a maternity colony of ca. 200 ind. (obs.); 3 ad.
males (ZDNU 2001/61–63) (cf. Karataş et al. 2003). – Nevşehir: Gülşehir, Tozköy Airport (leg. H. Bİl­
gen), 22 September 2003: 1 ad. male (ZDNU 2003/108); Ürgüp, Sarıhıdır Village, Armutludelik (a tunnel
near Kızılırmak River) (890 m), 6 September 2004: 1 ad. male (ZDNU 2004/344), 2 ind. from a colony
of ca. 20–30 ind. (released).
Pipistrellus pipistrellus (Schreber, 1774) s. l.
Kırıkkale: Kırıkkale (centrum), Gürler Quarter (house), 7 November 2003: 1 ad. male (ZDNU 2003/138)
(leg. H. Yurtseven). – Kırşehir: Kırşehir (centrum), 9 October 2004: 1 male (ZDNU 2004/363); 1 male,
2 females (leg. Ş. Özkurt) (released). – Nevşehir: Nevşehir (centrum) (house), 12 November 2003: 1 ad.
female (leg. H. Bİlgen) (ZDNU 2003/137). – Niğde: Kayabaşı Quarter, 1996: 1 ad. cf. female (ZDNU
55/1996); June 2001: 1 ind.; near Gebere Dam Lakelet (1411 m), 21–22 July 2005: 1 ad. male (ZDNU
2005/62), 1 ad. male (released); 23–24 July 2005: 2 ad. males (released); Aşlama (house), 21 April 2001:
1 ad. female (ZDNU 2001/17); 2 ind. (deposited in Ankara Univ.); Sazlıca, near Akkaya Dam Lakelet;
spring 2002: net. 2 ad. males (released); Ulukışla, Çiftehan, Maden Village (house), 23 May 2001: 1 sex
? (ZDNU 2001/17), 1 male (deposited in Ankara Univ.), a colony of 20–30 ind. (obs.).
Pipistrellus kuhlii (Kuhl, 1817)
Kayseri: ca. 2 km N of Yahyalı, a stream along road to Develi, 14–15 July 2001: net 2 ad. males.
Hypsugo savii (Bonaparte, 1837)
Niğde: near Gebere Dam Lakelet (1411 m), 23–24 July 2005: 3 ad. males (ZDNU 2005/67–69); 12–13 Au­
gust 2005: 2 ad. females (net. A. Karataş & Ş. Özkurt) (ZNDU 2005/83–84); 13–14 August 2005: 1 ad.
female (net. A. Karataş & Ş. Özkurt) (ZDNU 2005/87).
Plecotus macrobullaris Kuzjakin, 1965
Kayseri: Bünyan, Karadayı Village, Karatay Hanı Caravanserai, 19 September 2002: 1 ad. female (ZDNU
2002/133); Talas, Başakpınar, Örenyeri, 13 July 2001: 1 ad. male (ZDNU 2001/64; as P. auritus in Karataş
et al. 2003). – Nevşehir: De­rin­kuyu, Suvermez, 15 June 2002: 1 ad. male (ZDNU 2002/40); – Niğde:
Gümüşler, Eski Gümüş (house under con­struc­ti­on), August 1999: a colony of 10–15 ind. (obs.); Gümüşler
Monastery, 20 July 1996: 1 ad. male, 2 ad. females (ZDNU 42–44/1996; as P. auritus in Karataş et al.
2003); Ulukışla, Çiftehan, Maden Village, Bolkar mines, snow tunnels, 27 June 2001: 1 ad. male (coll. A.
Karataş & K. Ünsal; as P. auritus in Karataş et al. 2003). – Sivas: Hâfik, 3–4 km NW, Yıkılgan vicinity,
10 August 2003: 1 ad. female, 1 sad. male (ZDNU 2003/53–54), 1 sad. female (released).
Plecotus teneriffae Barrett-Hamilton, 1907
Kayseri: Melikgazi, 2 km NW of Gürpınar, 13 July 2001: 1 sad. male, 1 sad. female (ZDNU 2001/69,
70; as P. austriacus in Karataş et al. 2003); Yeşilhisar, Soğanlı, Eski Soğanlı, Ballık vicinity, 06 July
2001: 3 juv. males, 2 ad. females (lact.) (ZDNU 2001/49–51, 72, 80; leg. Y. Karakaya; as P. austriacus
in Karataş et al. 2003). – Niğde: N. U. Campus, Milli Piyango Hostel (leg. N. İpek), 28 September 2003:
1 ad. female (ZDNU 2003/101).
Miniopterus schreibersii (Kuhl, 1817)
Kayseri: Kocasinan, Kuşçu, Sarıağıl Cave, 12 July 2001: a maternity colony of ca. 300 ind. (obs.); 2 males, 4 females (ZDNU 2001/54–59) (cf. Karataş & Sözen 2004). – Nevşehir: Ürgüp, Sarıhıdır Village,
Ar­mutlu­de­lik (tunnel) (890 m), 6 September 2004: 1 ad. male (ZDNU 2004/341), 9 ind. from a colony
of ca. 200 ind. (released) (cf. Karataş & Sözen 2004). – Niğde: N. U. Campus, 22 September 2003: 1
male (found dead) (ZDNU) (cf. Karataş & Sözen 2004); near Gebere Dam Lakelet (1411 m), 23–24 July
154
Table 1. List of bat species recorded in the study area; + literature records (see Benda & Horáček 1998,
Spitzenberger et al. 2006); * first recorded; ** previously recorded under different names
Tab. 1. Přehled druhů netopýrů nalezených ve studovaném území: + publikované nálezy (viz Benda &
Horáček 1998, Spitzenberger et al. 2006); * nalezen poprvé; ** původně hlášen pod odlišným jménem
Sivas
Yozgat
Rhinolophus ferrumequinum
Rhinolophus hipposideros
Rhinolophus euryale
Rhi­no­lo­phus mehelyi
Myotis bly­thii
Myotis myotis
Myotis capaccinii
Myotis aurascens
Myotis brandtii
Myotis mystacinus s. l.
Ep­te­si­cus serotinus
Hypsugo savii
Pipistrellus kuhlii
Pipistrellus pipistrellus s. l.
Plecotus macrobullaris
Plecotus teneriffae
Barbastella cf. barbastellus
Miniopterus schreibersii
Tadarida teniotis
species \ district
Çankırı KayseriKırıkkaleKırşehirNevşehir Niğde
+
+
–
–
+
–
–
–
–
+
+
–
–
+
–
–
–
–
–
*
*
–
–
*
*
*
–
–
–
+
–
+
–
**
**
–
*
+
+
–
–
+
–
–
–
–
–
–
–
–
–
*
–
–
–
–
–
+
–
–
+
+
+
–
–
–
–
–
–
–
*
–
–
–
–
–
*
–
–
–
+
+
*
–
–
–
–
–
–
+
**
–
+
*
–
*
+
–
–
+
+
–
*
–
–
*
*
–
*
**
*
–
+
–
+
*
+
–
+
–
–
–
–
–
+
–
–
–
*
–
–
–
+
*
–
–
–
+
*
–
–
*
–
+
–
–
–
–
–
–
–
–
total
6
11
3
5
8
11
7
5
2005: 1 ad. female (lact.) (ZDNU 2005/43); Gümüşler, Epcik Cave, 12 August 1999: 1 sad. male, 1 sad.
& 1 ad. females (ZDNU 1999/28–30); 29 July 2002: 2 ad. males, 4 ad. females (ZDNU 2002/104–109),
3 males, 6 females (released) (cf. Karataş & Sözen 2004).
Tadarida teniotis (Rafinesque, 1814)
Sivas: Zâra, 9–10 August 2003: min. 20–50 ind. (obs.).
RESULTS AND Discussion
Rhinolophus ferrumequinum is the most frequently recorded species in Turkey and has been
recorded in all eight provinces in the study area. Another abundant species is Myotis blythii
which has been recorded in seven provinces in the study area but not in the Kırşehir province
(cf. Benda & Horáček 1998).
We did not find Myotis mystacinus s. l. which was recorded at the Devrez River (Çankırı: Ilgaz)
by von Helversen (1989). Benda & Tsytsulina (2000) and Benda & Karataş (2005) revised
the mystacinus group and stated that this record might be incorrect and most likely pertained
to M. aurascens. According to these studies, M. mystacinus s. str. is very rare in Anatolia. The
closest record to the study area was made near Kızılcahamam which is situated close to the
northeastern part of the area (Benda & Karataş 2005).
155
Table 2. Comparison of occurrence of bat species in Turkey with respect to geographic regions (Central
Anatolia is divided into two subregions, the eastern part [= the area under study] and the western parts)
Tab. 2. Srovnání výskytu netopýřích druhů v Turecku podle zeměpisných oblastí (střední Anatolie je
rozdělena na dvě podoblasti, východní [= studované území] a západní)
species \ region
Mediter- Aegean Marmara Black
E
SE
CW
CE
ranean Sea AnatoliaAnatolia AnatoliaAnatolia
Rousettus aegyptiacus
Taphozous nudiventris
Rhinolophus ferrumequinum
Rhinolophus hipposideros
Rhinolophus euryale
Rhinolophus blasii
Rhinolophus mehelyi
Myotis bechsteinii
Myotis myotis
Myotis blythii
Myotis nattereri
Myotis emarginatus
Myotis mystacinus
Myotis aurascens
Myotis nipalensis
Myotis brandtii
Myotis capaccinii
Myotis daubentonii
Nyctalus noctula
Nyctalus leisleri
Nyctalus lasiopterus
Eptesicus serotinus
Eptesicus bottae
Vespertilio murinus
Pipistrellus pipistrellus
Pipistrellus pygmaeus
Pipistrellus kuhlii
Pipistrellus nathusii
Hypsugo savii
Plecotus auritus
Plecotus macrobullaris
Plecotus austriacus
Plecotus teneriffae
Barbastella barbastellus
Barbastella cf. barbastellus
Otonycteris hemprichii
Miniopterus schreibersii
Tadarida teniotis
1
0
1
1
1
1
1
1
1
1
1
1
1?
1
0
0
1
0
1
1
0
1
1
0
1
0
1
1
1
0
0
0
1
0
0
0
1
1
0
0
1
1
1
1
1
0
1
1
1
1
1?
1
0
0
1
0
0
0
0
1
1
0
1
0
?
0
1
0
0
0
1
0
0
0
1
1
0
0
1
1
1
1
1
1
1
1
1
1
1
0
0
0
1
1
1
1
1
1
0
0
1
1
1
1
1
1
0
1
0
1
0
0
1
1
0
0
1
1
1
1
1
1
1
1
1
1
1
1
0
1
1
1
0
1
1
1
0
1
1
0
1
0
1
1
1
0
0
1
0
0
1
1
0
0
1
1
1?
0
1
0
1
1
1
0
1?
1
1
0
1
0
0
0
0
1
0
1
1
0
1
1
1
?
1
0
0
0
0
0
1
1
0
1
1
1
1
0
1
0
1
1
1
1
1?
0
0
0
1
0
1
0
0
1
1
1
1
0
1
0
1
0
0
0
0
0
1
1
1
1
0
0
1
1
1?
0
1
0
1
1
0
0
1
1
0
0
0
0
0
0
0
1
0
0
1
0
0
0
1
0
1
0
1
0
0
0
1
1
0
0
1
1
0
0
1
0
1
1
0
0
1?
1
0
1
1
0
0
0
0
1
0
0
1
0
1
0
1
0
1
0
1
0
1
0
1
1
Benda & Horáček (1998) recorded Barbastella cf. barbastellus near Ürgüp (Nevşehir), howe­
ver, they released the specimen without examining it in detail. It is not clear whether this bat
was the boreal B. barbastellus or the steppe B. leucomelas. The Ürgüp area is a steppe region
and so it is most likely that it was B. leucomelas.
156
Moreover, Rhinolophus euryale recorded at Zâra (Sivas), and R. mehelyi from Keskin (Kırıkkale) and from the Seyfe Lake (Kırşehir) (see Benda & Horáček 1998) were not found by
us in these regions.
On the other hand, Myotis brandtii, which had been recorded only in Rize in Turkey, was
recorded surprisingly in the steppe Yozgat province (cf. Benda & Karataş 2005). Hypsugo
savii, Myotis aurascens, M. capaccinii and Plecotus teneriffae were also recorded for the first
time in the study area.
Additional specimens of M. capaccinii collected in the study area clearly show that this species
is present not only in the Mediterranean (Akdeniz) and Marmara regions but also in suitable
habitats along the Kızılırmak River in central Anatolia.
Previously, western Palaearctic bats of the genus Plecotus had been included in P. auritus
and P. austriacus. Similarly, Plecotus in central Anatolia had been considered to be P. auritus
and P. austriacus (see Benda & Horáček 1998, Karataş et al. 2003). The genus Plecotus has
been recently re­vi­sed by Spitzenberger et al. (2006) and some new species have been described
(see Spit­zen­ber­ger et al. 2001, 2002, 2006, Benda et al. 2004, Juste et al. 2004). After that,
the ta­xo­no­mic position of Plecotus bats in central Anatolia became controversial. According to
Spit­zen­ber­ger et al. (2006), the species present in central Anatolia is P. macrobullaris (Fig. 2),
while P. auritus may have a limited distribution area in the eastern Black Sea (Karadeniz) region
and P. austriacus in Turkish Thrace. Moreover, according to DNA analysis of the specimens
from Karaman, which is situated in the sou­the­as­tern part of the study area, P. teneriffae is also
found in the area (Juste et al. 2004).
Fig. 2. Plecotus macrobullaris in a artificial cave at Talas, Başakpınar (Kayseri Prov.) .
Obr. 2. Podobnost východní části střední Anatolie podle fauny netopýrů.
157
Fig. 3. Relationship of the eastern part of Central Anatolia with respect to bat fauna.
Obr. 3. Podobnost východní části střední Anatolie podle fauny netopýrů.
This study shows that 3 of 5 Turkish bat families (60%), 9 of 14 genera (64%), and 19 of 37
species (51.4%) are present in this area (Table 2). According to the analysis using the MVSP 3.1
software (Multivariate Statistical Package, Kovach Computing Services), the species di­ver­si­ty
of the eastern part of Central Anatolia mostly resembles that of the western part of the region.
It seems that the area shows zoogeographic affinity with Eastern Anatolia (Fig. 3). The results
of the present study are consistent with those given by Benda & Horáček (1998).
Based on the data on distribution of bats and the above mentioned analysis, the following
four zoogeographic zones can be distinguished in Turkey.
1. Mediterranean (Akdeniz) zone: Akdeniz, Ege regions (except for the inner side), western
part of the Marmara region with Mediterranean climate;
2. Central Anatolian steppe zone: eastern part of Ege, Central Anatolia, Eastern and Sou­the­as­tern Anatolia (except for lowlands near the Syrian border) with continental climate;
3. Black Sea (Karadeniz) boreal zone: east of Marmara, Karadeniz, Erzurum and Kars plateau
of Eastern Anatolia with very humid climate;
4. Southeastern Anatolian eremial zone: lowlands along the Syrian border.
SOUHRN
V uvedeném přehledu předkládáme nálezy 15 druhů netopýrů v oblasti středního a horního toku řeky
Kizilirmak ve střední Anatolii (Turecko). Čtyři z těchto druhů (Myotis capaccinii, M. aurascens, M.
brandtii, Hypsugo savii) byly nalezeny v uvedeném regionu poprvé, zatímco čtyři další druhy netopýrů
zaznamenané dřívě nalezeny nebyly. Předložené výsledky zvyšily známý počet netopýrů oblasti na 19.
Taxonomická posice netopýrů rodu Plecotus, která je v současné době šířeji diskutována, byla vyhodnocena
158
i ve studovaném regionu; nalezen byl druh P. teneriffae a poprvé tak zaznamenán z regionu pod tímto
jménem. Z osmi provincií (vilájetů) střední Anatolie dvě vykazují bohatší faunu netopýrů (11 druhů),
Niğde a Kayseri.
Acknowledgements
We wish to thank M. Ak, H. Baştürk, S. Baştürk, H. Bİlgen, Dr. R. Bİlgİn, K. Doğan, N. İpek, H. Ka­
rakaya, Y. Karakaya, Dr. Ay. Karataş, O. Kizilcik, Ş. Özkurt, H. C. Öztekİn, F. Telli, K. Ünsal, M. A.
Yİğİt, and H. Yurtseven for their help with collecting data and material. A part of this study was supported
by the Research Fund of Niğde University (Nr. 01/FEB/023). We dedicate this study to the memory of
the Late Res. Assist. Kuddusi Ünsal.
References
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Benda P. & Karataş A., 2005: On some Mediterranean populations of bats of the Myotis mystacinus
morpho-group (Chiroptera: Vespertilionidae). Lynx, n. s., 36: 9–38.
Benda P. & Tsytsulina K., 2000: Taxonomic revision of Myotis mystacinus group (Mammalia: Chiroptera)
in the Western Palearctic. Acta Soc. Zool. Bohem., 64: 331–398.
Benda P., Kiefer A., Hanák V. & Veith M., 2004: Systematic status of African populations of long-eared
bats, genus Plecotus (Mammalia: Chiroptera). Folia Zool., 53, Monograph 1: 1–48.
von Helversen O., 1989: New records of bats (Chiroptera) from Turkey. Zool. Middle East, 3: 5–18.
Juste J., Ibáñez C., Muñoz J., Trujillo D., Benda P., Karataş A. & Ruedi M., 2004: Mitochondrial phy­
lo­ge­o­gra­phy of the Long-eared bats (Plecotus) in the Mediterranean Palaearctic and Atlantic Islands.
Mol. Phylogenet. Evol., 31: 1114–1126.
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Long-winged bat, Miniopterus schreibersii (Chiroptera: Vespertilionidae), in Turkey. Zool. Middle
East, 33: 51–64.
Karataş A., Benda P., Toprak, F. & Karakaya H., 2003: New and significant records of Myotis capaccinii
(Chiroptera: Vespertilionidae) from Turkey, with some data on its biology. Lynx, n. s., 34: 39–46.
Spitzenberger F., Piálek J. & Haring E., 2001: Systematics of the genus Plecotus (Mammalia, Ve­sper­ti­li­
o­ni­dae) in Austria based on morphometric and molecular investigations. Folia Zool., 50: 161–172.
Spitzenberger F., Haring E. & Tvrtković N., 2002: Plecotus microdontus (Mammalia, Vespertilionidae),
a new bat species from Austria. Natura Croatica, 11: 1–18.
Spitzenberger F., Strelkov P. P., Winkler H. & Haring E., 2006: A preliminary revision of the genus
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159
Lynx (Praha), n. s., 37: 161–172 (2006).
ISSN 0024–7774
Jan Svatopluk Presl’s (1821) family-group names of mammals
Jména savců skupiny čeledi vytvořená Janem Svatoplukem Preslem (1823)
Jiří Mlíkovský
Department of Zoology, National Museum (Natural History), Václavské náměstí 68,
CZ–115 79 Praha 1, Czech Republic; [email protected]
received on 5 September 2006
Abstract. Jan Svatopluk Presl created in 1821 eight family-group names of mammals, the existence of
which has been overlooked by modern mammalogists. Three of them were not listed in subsequent ca­ta­lo­
gues (Catadontidae, Hypsiprymnidae and Lipuridae), while the remaining five were generally at­tri­bu­ted to
various junior authors in modern literature (Myrmecophagidae, Petauridae, Chironectidae, Ge­o­rychi­dae,
and Thylacidae). In addition, the family-group name Phyllostomatidae, also used by Presl (1821), was
found to be attributable to Goldfuss (1820), not to Gray (1825) as generally believed.
Introduction
Jan Svatopluk Presl (1791–1849) was a renowned Czech naturalist, whose scientific in­te­rests
ranged from botany to mammalogy (Hoffmannová 1973). In 1821–1823 he published a sys­te­
ma­tic synopsis of vertebrates (Presl 1821, 1822, 1823; see also Presl 1834). In this modernly
structured classification he was one of the first naturalists to recognize the family level (cf.
Mayr et al. 1953, Bock 1994). Prior to him, the family level was recognized and family-group
names were properly formed in mammalogy only by a few authors, such as Karl Illiger
(1775–1813) in 1811, Gotthelf Fischer von Waldheim (1771–1853) in 1817, Georg August
Goldfuss (1782–1848) in 1820, Wilhelm Friedrich Hemprich (1798–1825) in 1820, and John
Edward Gray (1800–1875) in 1821 (Illiger 1811, Fischer von Waldheim 1817, Goldfuss
1820, Hemprich 1820, Gray 1821).
Presl’s classification of mammals (Presl 1821) was published in Czech language in the then
newly founded, prestigious, Czech scientific journal ‘Krok’ (see Laiske 1959), which might
have been the reason, why it has been overlooked by Palmer (1904) and subsequent students
of mammalian family-group names (Simpson 1945, Wilson & Reeder 2005). Below I present
an account of mammalian family-group names used by Presl (1821), with special respect to
no­menc­la­tu­ral issues, which arose from their re-discovery. Family-group names are arranged
al­pha­be­ti­cal­ly in the systematic part. Presl’s (1821) classification of mammals is given in full
in the Appendix.
161
Systematic part
Catodontidae Presl
Presl (1821: 84) created family Catodonta for Catodon Linnaeus, 1761, which is a junior sub­
je­cti­ve synonym of Physeter Linnaeus, 1758 (Mead & Brownell 2005). Catodonta Pre­sl, 1821
is thus a junior subjective synonym of Physeteridae Gray, 1821 (I assume here that the paper
by Gray was published earlier than that by Presl).
Chironectidae Presl
Palmer (1904: 734; see also Simpson 1935: 135; 1945: 41) attributed the family-group name
Chi­ro­ne­cti­dae to Anonymous (1897). The name was used already by Presl (1821: 80), who
spelled it Cheironectina and based it on Chironectes Illiger, 1811. Cheironectes is a sub­se­quent
incorrect spelling of Chironectes Illiger, hence the family-group name should be co­r­rec­ted to
Chi­ro­ne­cti­na (ICZN 1999: Art. 35.4.).
Water opossums of the genus Chironectes Illiger, 1811 are usually included in the sub­fa­mi­ly
Didelphinae of the family Didelphidae Gray, 1821 (Gardner 2005). Chironectina Presl, 1823
is thus a junior subjective synonym of Didelphidae Gray, 1821, but it is available if Chironectes
opossums are separated at the subfamily level, as recently suggested by Hershko­vitz (1997).
Georychidae Presl
Mole-rats of the genus Georychus Illiger, 1811 are usually included in the family Ba­thy­er­gi­dae
Waterhouse, 1841 (Simpson 1945, Woods & Kilpatrick 2005). Woods & Kilpatrick (2005: 1538)
attributed the family-group name Georychidae to Roberts (1951), but already Simpson (1945:
99) traced the name back to Gravenhorst (1841: facing p. 502), who spelled it Georychina.
The name should be attributed to Presl (1821: 81), who spelled it Georychina and based it on
the genus Georychus Illiger, 1811.
Georychina Presl, 1821 antedates Bathyergidae Waterhouse, 1843, which is the next oldest
family-group name available for the family. Georychina Presl was not replaced by Ba­thy­er­gi­
dae Waterhouse because it was based on a junior synonym (cf. ICZN 1999: Art. 40). However,
Georychina Presl should be set aside, because Bathyergidae Waterhouse is in prevailing use and
both conditions of Art. 23.9.1. (ICZN 1999) are met: the name has not been used as valid after
1899 (Art. 23.9.1.1.) and at least 10 authors used Bathyergidae Waterhouse as valid taxon in at
least 25 works in immediately preceding 50 years in a period that encompasses over 10 years
(Art. 23.9.1.2.). Required citations are as follows: Wood 1958, 1965, de Graff 1975, Jarvis
1978, Nevo 1979, Harvey et al. 1980, Nevo et al. 1987, Honeycutt et al. 1987, 1991, Bennett
& Jarvis 1988, Denys 1989, Lovegrove 1989, 1991, Jarvis & Bennett 1990, 1991, Allard &
Honeycutt 1992, Janeck et al. 1992, Burda & Kawalika 1993, Filipucci et al. 1994, Buffen­
stein 1996, Faul­kes et al. 1997, McKenna & Bell 1997, Bennett & Faul­kes 2000, Spinks et
al. 2000, Walton et al. 2000, Oosthuizen et al. 2003.
Hypsiprymnidae Presl
Presl (1821: 80) based his family Hypsiprimnea [sic!] on the genus Hypsiprimnus, which is
a subsequent incorrect spelling of Hypsiprymnus Illiger, 1811, The family-group name thus
should be corrected to Hypsiprymnidae (ICZN 1999, Art. 35.4.). Hypsiprymnus Illiger, 1811 is
162
a junior objective synonym of Potorous Desmarest, 1804 (Groves 2005b). Hence, Hyp­siprym­
ni­da Presl, 1821 is a junior objective synonym of Potoridae Gray, 1821 (I assume here that the
paper by Gray was published earlier than that by Presl).
Lipuridae Presl
Presl (1821: 80) based his family Lipurina on the genus Lipurus Goldfuss, 1817, which is a
junior subjective synonym of Phascolarctos Blainville, 1816 (Lee & Carrick 1989). Li­pu­ri­na
Presl, 1821 antedates Phascolarctidae created by Owen (1839), which is the next oldest family-group name available for the family. Lipurina Presl was not replaced by Phas­co­lar­ci­dae Owen
because it was based on a junior synonym (cf. ICZN 1999, Art. 40). However, Lipurina Presl
should be set aside, because Phascolarctidae Owen is in prevailing use and both conditions of
Art. 23.9.1. (ICZN 1999) are met: the name has not been used as valid after 1899 (Art. 23.9.1.1.)
and at least 10 authors used Phascolarctidae Owen as valid taxon in at least 25 works in immediately preceding 50 years in a period that encompasses over 10 years (Art. 23.9.1.2.). Required
citations are as follows: Turnbull & Lundelius 1970, Imai et al. 1983, Strahan 1983, Nagy
& Martin 1985, Aplin & Archer 1987, Haight & Nelson 1987, McKay 1988, Lee & Carrick
1989, Davidson & Young 1990, Harding & Aplin 1990, Gor­don 1991, Luckett 1994, Osawa
1993, Szalay 1994, Retief et al. 1996, Black & Archer 1997, Kirsch et al. 1997, Kemper
et al. 2000, Sherwin et al. 2000, Fisher et al. 2001, Grand & Barboza 2001, Cardillo et al.
2004, Kavanagh et al. 2004, Archer & Kirsch 2006, Mu­ne­ma­sa et al. 2006, Weisbecker &
Sánchez-Villagra 2006.
Myrmecophagidae Presl
The family-group name Myrmecophagidae is widely used and generally attributed to Gray
(1825b: 343) (e.g. Gardner 2005: 102). It should be attributed to Presl (1821: 82), who based
it on the genus Myrmecophaga Linnaeus, 1758.
Petauridae Presl
Groves (2005b: 53) attributed the family-group name Petauridae to Bonaparte (1838: 112).
It should be attributed to Presl (1821: 79), who spelled it Petaurina and based it on the genus
Petaurus Shaw, 1791.
Phyllostomatidae Goldfuss
Simmons (2005: 395) attributed the family-group name Phyllostomidae [sic!] to Gray (1825a:
242), although most authors attributed it to Gray (1825b: 338). Relative priority of these publications is irrelevant, however, because the name was used prior to Gray (1825a, b) al­rea­dy
by Presl (1821: 78) and Goldfuss (1820: 460), of whom the latter spelled it Phyl­los­to­ma­ta and
based it on the genus Phyllostoma “Geoffr[oy Saint-Hilaire]” = Cuvier, 1800, which is a junior
objective synonym of Phyllostomus Lacépède, 1799 (Hemming 1955). The family-group name
Phyl­los­to­ma­ti­dae thus should be attributed to Goldfuss (1820).
The family was generally spelled Phyllostomatidae until Kuzjakin (1974) and Handley (1980)
argued that the name should be spelled Phyllostomidae. However, this is linguistically incorrect.
Following Article 29.3.1. of the International Code of Zoological Nomenclature (ICZN 1999),
“the stem for the purposes of the Code is found by deleting the case ending of the appropriate
163
genitive singular”. Here, the genitive singular of ‘stoma’ is ‘stomatis’, and the family-group
name thus should be spelled Phyllostomatidae (see also Keržner 1974).
Thylacidae Presl
Presl (1821: 80) based his family Thylacina on the genus Thylacis Illiger, 1811. The latter genus
is generally included in the family Peramelidae Gray, 1825 (Simpson 1945, Grover 2005a).
Thy­la­ci­na Presl, 1821 antedates Peramelidae Gray, 1825, which is the next oldest family-group
name available for the family. Thylacina Presl was not replaced by Peramelidae Gray because
it was based on a junior synonym (cf. ICZN 1999, Art. 40). However, Thylacina Presl should
be set aside, because Peramelidae Gray is in prevailing use and both conditions of Art. 23.9.1.
(ICZN 1999) are met: the name has not been used as valid after 1899 (Art. 23.9.1.1.) and at least
10 authors used Peramelidae Gray as valid taxon in at least 25 works in immediately preceding
50 years in a period that encompasses over 10 years (Art. 23.9.1.2.). Required citations are as
follows: Lidicker & Follett 1968, Turnbull & Lundelius 1970, Archer & Kirsch 1977, 2006,
Vaughan 1978, Stoddart & Braithwaite 1979, Gemmell 1982, Szalay 1982, Strahan 1983,
Aplin & Archer 1987, Gordon & Hulbert 1989, Friend 1990, Groves & Flannery 1990, Kem­
per et al. 1990, Southgate 1990, Menzies 1991, Sher­win et al. 1991, Murphy & Serena 1993,
Retief et al. 1995, McKenna & Bell 1997, Short et al. 1998, Muirhead 2000, Westermann et
al. 2001, Brou­gh­ton & Dickman 2002, Cham­bers & Dickman 2002, Richards & Short 2003,
Price 2004, Wes­ter­man et al. 2004.
Souhrn
Jan Svatopluk Presl vytvořil roku 1821 ve své klasifikaci savců osm nových jmen skupiny čeledi. Z nich
tři jména vůbec nebyla zahrnuta do pozdějších katalogů (Catadontidae, Hypsiprymnidae a Lipuridae),
zatímco autorství ostatních pěti jmen bylo běžně připisováno různým mladším autorům (Myr­me­co­pha­
gi­dae, Pe­tau­ri­dae, Chironectidae, Georychidae a Thylacidae). Kromě toho bylo zjištěno, že jméno Phyl­
los­to­ma­ti­dae, zpra­vi­dla připisované Grayovi (1825a, b) a taktéž použité Preslem (1821), bylo vytvořeno
již Goldfussem (1820).
Acknowledgments
I am obliged to Petr Benda (Praha) for comments on the manuscript. The study was supported
in part by the grants of the Ministry of Culture of the Czech Republic No. DE06P04OMG008
and MK00002327201.
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Appendix
Presl’s (1821) classification of mammals. Czech names proposed by Presl (1821) for mam­ma­li­
an orders, families and genera are given in parentheses (most were not listed by Anděra 1999).
Note that Presl (1821) did not name families when an order included only a single family.
1. Bimana (dvaurucí)
1.1. [Family not given]: Homo (člowěk).
2. Quadrumana (čtwerorucí)
2.1. Simia (opicowití): Pithecus (op), Hylobates (ramenáč), Lasiopyga (dlakořit), Cercopithecus (koč­
ko­dán), Papio (martyška), Poago (mirkuwín), Cynocephalus (pso­hlaw), Colobus (kykatas), Ateles
(cha­pan), Cebus (malpa), Pithecia (chwostan), Aotus (nočák), Callitrix (pěknowlasec), Hapale
(kos­man).
2.2. Prosimia (munowití): Lemur (muna), Lichanotus (požast), Stenops (autloň).
2.3. Prehensilia (pal­cu­chowi­tí): Chirogeleus (palcucha).
2.4. Macrotarsia (nártaunowití): Tarsius (nár­taun), Otolicnus (uchoš).
3. Cheiroptera (letauni)
3.1. Galeopithecia (letuškowití): Galeopithecus (letuška).
3.2. Phyllostomata (řasonosowití): Phyl­los­to­ma (řasonos), Nycteris (šerowec), Rhinopoma (nosalec),
Rhinolophus (wrapenec), Megaderma (weloblanec).
3.3. Harpiae (upírowití): Pteropus (upír), Ce­pha­lo­tes (hlawan).
3.4. Noctiliones (nedopírowití): Ste­no­der­ma (auzkoblanec), Vespertilio (netopír), Plecotus (ušan), Mi­op­te­rus (spaka), Nyctionomus (pří­še­rec), Noctilio (mračník), Dysopes (psohubec), Thapazous (we­čer­
ník).
4. Ferae (šelmy)
4.1. Canina (psovití): Megalotis (welouch), Canis (pes), Hyena (hyena), Felis (kot), Rycaena (zeník).
4.2. Mustelina (kunowití): Herpestes (promyka), Mephitis (smrdoš), Mustela (kuna), Ichneumon (mun­
kos), Lutra (wydra).
4.3. Ursina (nedwědowití): Ursus (nedwěd), Meles (jezwec), Gulo (rosomak), Procyon (mýwal), Na­sua
(nosál), Cercoleptes (ohonowin).
4.4. Erinacea (ježovití): Centetes (bodlín), Erinaceus (jež).
4.5. Talpina (krtkowití): Condylura (uzloš), Chrysochloris (zlatokrt), Talpa (krt), Scalops (hrabuška).
4.6. Sorexina (reyskowití): Sorex (reysek), Mygale (wychuchol).
5. Marsupialia (waknatí)
5.1. Petaurina (letawcowití): Petaurus (létawec), Phalangista (parowník).
170
5.2.
5.3.
5.4.
5.5.
5.6.
Cheironectina (plawá­kowi­tí): Cheironectes (plawák).
Didelphina (wačicowití): Ambliotis (ru­hoš), Balantia (tokaun), Didelphis (wačice).
Thy­la­ci­na (torebníkowití): Dasyurus (srstaun), Thylacis (torebník).
Lipurina (kwíkolowití): Phascolomys (drapoš), Lipurus (kwíkol).
Hypsiprimnea (skokey­šowi­tí): Hypsiprimnus (skokeyš), Halmaturus (klokan).
6. Rosores (hlodawí)
6.1. Dipudina (tarbíkowití): Dipus (tarbík), Pedetes (nohas).
6.2. Sciurina (wewerowití): Myoxus (plch), Tamias (deňka), Pteromys (poletucha), Sciurus (wewer),
Che­i­ro­mis (letaha).
6.3. Leporina (zajícowití): Lepus (zajíc), Lagomys (pičuha).
6.4. Cavinida (morčowití): Co­e­lo­ge­nys (tlamák), Dasyproctes (nahoš), Cavia (morče, also wiska), Hyd­ro­chae­rus (plawaun).
6.5. Histricina (dikobrazowití): Histrix (dikobraz), Coëndu (ostnoš), Lon­che­res (ježowec).
6.6. Castorina (bobrowití): Hydromys (woduška), Guillino (dlakaun), Ondatra (ondatra), Castor
(bobr).
6.7. Georychina (hrabošowití): Georychus (hraboš), Hypudaerus (pestruška), Fiber (dlakoš).
6.8. Murina (myšowití): Mus (myš), Cricetus (křeček), Arctomys (swišť), Viscacia (wizkacha), Spalax
(slepec), Bathyergus (rypoš).
7. Cingulata (pásatí)
7.1. [Family not given]: Dasypus (pásowec), Tolipeutes (chaulan).
8. Pamphracta (Ssupani)
8.1. [Family not given]: Pamphractus (ssupan).
9. Ornithostomata (ptakohubí)
9.1. [Family not given]: Ornithorhynchus (ptakaun).
10. Tachiglossa (rychlozajiční)
10.1.[Family not given]: Tachyglossus (rychlojazan).
11. Vermilinguia (tenkojazyční)
11.1 Manisia (luskaunowití): Manis (luskaun).
11.2.Myrmecophagina (mrawenčíkowití): Myr­me­co­pha­ga (mrawenčík), Tamandua (tamandua), Ory­c­te­
ro­pus (kuťoš).
12. Tardigrada (lenochodi)
12.1.[Family not given]: Bradypus (lenochod), Choloepus (kulhoš).
13. Bisulca (dwaupaznehtní)
13.1.Taurina (beykowití): Bos (beyk), Ovibos (owoskot), Ovis (owce), Capra (kozel), Antilope (sajka).
13.2.Cervina (jelenowití): Cervus (jelen), Moschus (kabarha).
13.3.Girafina (girafowití): Ca­me­lo­par­da­lis (girafa).
13.4.Camelina (welblaudowití): Auchenia (wi­ku­ně), Camelus (welblaud).
14. Solidungula (jednopaznehtní)
14.1.[Family not given]: Equus (kůň).
15. Multungula (mnohopaznehtní)
15.1.Elephantina (slonowití): Elephas (slon).
15.2.Tapiracea (tapírowití): Tapirus (tapír).
15.3.Scrofina (wepřowití): Sus (wepř).
15.4.Hyracina (tlustošowití): Lipura (nehtaun), Hyrax (tlustoš).
15.5.Rhinocerina (rohošowití): Rhi­no­ce­ros (rohoš).
15.6.Hyppopotamea (hrochowití): Hyp­po­po­ta­mus (hroch).
171
16. Pinnipedia (ploskonozí)
16.1.[Family not given]: Phoca (teleň), Pusa (siwuč), Nepus (ťutě); also nerpa, lachták.
17. Syrt­o­ba­ti­ca (smeykali)
17.1.[Family not given]: Trichechus (morž).
18. Sirenia (ochechule)
18.1.[Family not given]: Manatus (kapustňak), Halicore (moroň), Ry­ti­na (koraun).
19. Cetacea (welrybi)
19.1.Balaenacea (kytowití): Balaena (kyt), Balanopter (pleytwák).
19.2.Catodonta (wor­wa­ňowi­tí): Physeter (perutoš), Oryx (zubaun), Cetus (olbroť), Catodon (worwaň),
Delphinus (pliskawice), Delphinapterus (běluha), Physalus (sykawice).
172
Lynx (Praha), n. s., 37: 173–177 (2006).
ISSN 0024–7774
Diet of the American mink (Mustela vison) in the Czech Republic
(Carnivora: Mustelidae)
Potrava norka amerického (Mustela vison) v České republice
(Carnivora: Mus­te­li­dae)
Michaela Nováková1,2 & Petr Koubek2
1
Institute of Botany and Zoology, Faculty of Science, Masaryk University, Kotlářská 2,
CZ–611 37 Brno, Czech Republic; [email protected]
2
Institute of Vertebrate Biology, Czech Academy of Science, Květná 8, CZ–603 65 Brno,
Czech Republic; [email protected]
received on 28 June 2006
Abstract. An analysis of 29 full stomachs of the American mink (Mustela vison Schreber, 1777), collected
in the Křivoklátsko Protected Landscape Area and in the Horažďovice region (Czech Republic) during
autumn and winter time in 2001–2006, showed a great diversity of the mink diet. Fish (Cyprinidae, Percidae, and Esocidae) were the dominant component of their diet (51.7% of occurrence, 41.0% of volume),
followed by mammals (Rodentia, Insectivora) (48.7% of occurrence, 31.1% of volume) and birds (20.7%
of occur­ren­ce, 13.2% of volume). Amphibians, insects, carrions and plants served only as sup­ple­men­ta­ry
food. Mink was found to be a typical generalist predator. Mink diet is, most probably, influenced by prey
availability, prey behaviour, season, prey abundance and habitat type.
INTRODUCTION
The American mink Mustela vison Schreber, 1777, is a North American mustelid predator that
was brought to Europe for its quality fur. The first wild living populations of American minks
appeared throughout Europe in the 1930s, as a large amount of American minks es­ca­ped from
captivity or were deliberately released into the wild (Sidorovich 1993). At the end of the 1990s,
American minks became a common European species living on the banks of rivers, lakes, ponds
and water reservoirs (Dunstone 1993, Bartoszewicz & Zalewski 2003). The first report on their
existence in the Czech Republic comes from the 1960s (Mazák 1964). The species started to
spread across Bohemia at the turn of the millennium. An eva­lua­ti­on of a national questionnaire
showed that in 2003 the American mink was present on 29.3% of the area of the Czech Republic
(Červený et al. 2005). The American mink is an un­spe­ci­a­li­sed predator (Wise et al. 1981, Br­
ze­ziński & Zurowski 1992, Jedrzejewska et al. 2001), hunting in water and also on land. Their
presence often leads to a steep decrease in their prey population (Bartoszewicz & Zalewski
2003). They hunt mostly in littoral vegetation and in water (Erlinge 1969). Water invertebrates
(crayfish), as well as various types of vertebrate species are their most common prey (Erlinge
1969, Sidorovich 2000, Jedrzejewska et al. 2001, Bartoszewicz & Zalewski 2003). Fish are
a dominant component of their diet (Akan­de 1972, Lodé 1993). The particular compositon of
173
their diet varies throughout the year and depends on the habitat the mink lives in (Gerell 1967,
Chanin & Linn 1980, Jenkins & Har­per 1980, Jedrzejewska et al. 2001).
MATERIAL AND METHODS
The diet of the American mink was studied using an analysis of stomach content of individuals coming
from Bohemia (Křivoklátsko Protected Landscape Area and Horažďovice region). In total, 51 stomachs
were analysed. The samples were continuously collected during autumn (September–November) and
winter sea­sons (December–February) in 2001–2006.
All minks were caught in accordance with nature conservation legislation, with the aim to protect po­pu­la­ti­
ons of indigenous animal species. All caught minks were processed using standard zoological methods.
Empty stomachs (22 samples, 43%) were not included in further analysis. Stomach contents were ana­
ly­sed using standard methods (Hammershøj et al. 2004). Each stomach content was mixed with water
on a Petri dish and individual components separated into groups. Various identification guides as well as
com­pa­ra­ti­ve material were used to determine the prey remains (bones, scales, feather, teeth, fur or hair
or other body parts) (Gaffrey 1961, Brom 1986, Teerink 1991). Results are showed as a percentage of
volume (V %), indicating the volume percentage of a certain food component in all of the analysed full
stomachs. In order to compare our results with those analysing American mink faeces, we also used
frequency of occurrence (F %), indicating the occurrence percentage of a certain food component in all
of the analysed full stomachs
RESULTS AND DISCUSSION
The analysis of 29 stomach contents showed that fish with the body length of up to 30 cm
(F=51.7%; V=41.0%) were the dominant component in the American mink diet. The most
frequently found species were from the families of Cyprinidae (similar to Erlinge 1969, Day
& Linn 1972), also Percidae (Erlinge 1969, Bartosewicz & Zalewski 2003) and Esocidae
(Erlinge 1969). It is clear that low activity of fish, caused by low water temperature during
winter months, was the main reason for their high abundance in the mink diet (Gerell 1967,
Erlinge 1969, Chanin & Linn 1980, Chanin 1981, Wise et al. 1981). The list of food com­po­
nents is shown in Table 1.
Mammals (Rodentia, Insectivora) were the second most important food component (F=48.7%;
V=31.1%). Considering that most predators avoid eating insectivorous mammals (Lockie 1961,
Macdonald 1977), occurrence of the lesser white-toothed shrew Crocidura suaveolens (F=6.9%;
V=3.8%) in mink stomachs was very interesting. Similar findings are very rare and if there are
some insectivores identified in the mink diet, it is usually only in small amounts (Wise et al.
1981, Brzeziński & Zurowski 1992, Maran et al. 1998, Jedr­ze­jewska et al. 2001). The presence
of a roe deer fur (Capreolus capreolus) in the stomachs proves the ability of minks to use large
mammal carcasses as a food source (F=3.5%; V=3.5%).
In comparison with other literature data, the richness of mammal species found in the diet of
minks in Bohemia is very low. This could be caused not only by the limited amount of analysed
samples and by the season of sample collection, but also, and mainly, by food avai­la­bi­li­ty in the
study area. For example, Lepus, Rattus or Apodemus species were totally absent from the diet of
minks in Bohemia, whereas they are very often hunted by mink populations living elsewhere
(Day & Linn 1972, Chanin & Linn 1980, Dunstone & Birks 1987, Bar­to­sewicz & Zalewski
2003). Exceptionally, small weasel predators such as the ermine Mustela erminea and weasel
Mustela nivalis (Akande 1972, Chanin & Linn 1980) can also be a prey of the American mink,
but neither these were recorded in our study. On the other hand, muskrats (Ondatra zibethicus)
174
Table 1. Diet of the American mink based on the analysis of stomach content. For explanations see Material and methods
Tab. 1. Potrava norka amerického podle analýzy obsahu žaludků. Vysvětlivky viz Material and methods
component / složka
F [%]
V [%]
Crocidura suaveolens
Talpa europea
Ondatra zibethicus
Microtus sp.
Glis glis
Rodentia sp. Capreolus capreolus
Mammalia unidentified
6.90
3.45
3.45
3.45
3.45
21.05
3.45
6.90
3.79
1.21
3.41
3.45
2.76
13.03
3.45
3.45
mammals birds
amphibians
52.10
20.69
3.45
27.55
13.24
1.72
Cyprinidae
Esocidae
Percidae
Pisces unidentified
27.59
6.90
10.34
10.34
19.81
4.13
9.01
6.86
fish invertebrates
plant material
unidentified
55.17
17.24
41.38
6.90
41.00
0.91
9.76
0.01
are known to be hunted more often than we recorded (Bartosewicz & Zalewski 2003). It is
obvious from the literature that mammals are a significant part of the American mink diet. The
frequency of mam­mal component in mink diet ranges from 28.0% (Akande 1972) up to 61.8%
(Brzeziński & Zu­rowski 1992).
Bird remains (Passeriformes, Galliformes) were recorded in 20.7% of cases (V=13.2%),
whe­re­as Bartosewicz & Zalewski (2003) found bird remains, during the same season, only
in 4–16% of cases. Birds happen to be the most common prey during breeding or migrating
seasons (Gerell 1967, Erlinge 1969, Akande 1972, Wise et al. 1981, Brzeziński & Zu­rowski
1992, Maran et al. 1998). Therefore, in spring and summer birds can become the dominant
component of the mink diet (Bartosewicz & Zalewski 2003).
Frogs are another significant component of the American mink diet (Gerell 1967, Br­ze­ziński
& Zurowski 1992, Hammershøj et al. 2004), especially bullfrogs. Minks avoid eating toads (Si­do­
ro­vich & Pikulik 1997). Despite the fact that minks often dig up wintering frogs (Jedrzejewska
et al. 2001), we found frog remains in our samples only once.
Remains of invertebrates were found in 17.2% of samples (V=0.9%). They were mostly water
arthropods and insects, although some of these could have already been present in the guts of
the consumed fish. Unlike water arthropods, especially crabs (Gerell 1967, Chanin & Linn
1980, Dunstone & Birks 1987, Brzeziński & Zurowski 1992, Sidorowich et al. 2001), insects
are con­su­med by minks very rarely (Erlinge 1969, Jedrzejewska et al. 2001).
175
Plant remains (grass, twigs, leaves, roots) were found in 41.38 % of samples (V=9.76%). In
six cases they formed the majority of the stomach content (V=20%=100%) and the­re­fo­re it is
highly unlikely that they were eaten by accident. A similar feeding behaviour was descri­bed
in the pine marten (Martes martes) by Lockie (1961) who found plant remains only in some
faeces samples during spring and autumn.
The season of sample collection and, in some cases, the used methods of stomach analysis did
not allow to prove the presence of bird eggs, earthworms or slugs in the American mink diet,
which are common food components of mink populations living elsewhere (Chanin & Linn
1980, Brzeziński & Zurowski 1992, Maran et al. 1998, Jedrzejewska et al. 2001).
The most significant factors influencing the American mink diet are season (Jedrzejewska et
al. 2001, Bartosewicz & Zalewski 2003) and the food availability in the given area (Cha­nin &
Linn 1980, Bartosewicz & Zalewski 2003). Other factors include prey behaviour, its abun­dan­
ce and the structure of mink habitat (Chanin & Linn 1980). For example, mammals, fish and
amphibians are the most important food sources in riverine habitats, whereas birds and fish are
dominating prey of minks living near lakes and ponds (Jedrzejewska et al. 2001, Bartosewicz
& Zalewski 2003).
Considering our small material of stomach samples collected mostly in autumn, it was not
possible to evaluate seasonal changes of the diet, sex differences in the diet nor differences
between sites. Our results show that food composition of American minks living in the Czech
Republic is almost identical with that of minks living in Scotland (all year round) (Akande
1972) and very similar to that of minks living in Poland (autumn and winter) (Bartosewicz &
Zalewski 2003).
SOUHRN
Potrava norka amerického, Mustela vison (Schreber, 1777), v České republice byla studována analýzou
žaludků pocházejících z podzimního a zimního období z let 2001–2006. Norci pocházeli z území Čech
(Ho­raž­ďo­vic­ko, CHKO Křivoklátsko). Celkem bylo získáno 51 žaludků, prázdné (43 %) byly z dalších
analýz vy­lou­če­ny. Ryby z čeledí Cyprinidae, Percidae a Esocidae tvořily nejvýznamnější část potravy
(frekvence výskytu [F] =51.7 %; objemové procento [V] =41.0 %), následované savci (Rodentia, Insectivora) (F=48.7 %; V=31.1 %) a ptáky (F=20.7 %; V=13.2 %). Obojživelníci, hmyz, kadávery velkých
savců a rostlinné zbytky doplňovaly potravu. Zajímavý je výskyt bělozubky Crocidura sua­veo­lens a naopak velmi vzácná přítomnost obojživelníků v potravě zkoumaných norků. Z našich výsledků vyplývá,
že norek americký se na území Čech živí poměrně širokým spektrem živočišných druhů (tab. 1). Potravní
složení je pravděpodobně ovlivněno ročním obdobím, potravní nabídkou, početností a chováním kořisti
a strukturou biotopu, který norek obývá.
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pla­ce­ment of Mustela lutreola by M. vision. J. Zool., London, 245: 218–222.
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ISSN 0024–7774
Distribution and status of Myotis bechsteinii in Bulgaria
(Chiroptera: Vespertilionidae)
Rozšíření a statut netopýra velkouchého (Myotis bechsteinii) v Bulharsku
(Chiroptera: Vespertilionidae)
Boyan P. Petrov
National Museum of Natural History, Tsar Osvoboditel blvd. 1, BG–1000 Sofia, Bulgaria;
[email protected]
received on 23 May 2006
Abstract. The first record of Myotis bechsteinii in Bulgaria dates from 1935. Since then, a total of 55 females, 141 males and two individuals of unknown sex have been recorded. Up to now only three breeding
colonies have been found in Bulgaria. At present, the Bechstein’s bat is known from 34 localities (33 UTM
squares) situated from sea level up to 1650 m. Since its first discovery, only two males have been found
hibernating in caves. Besides that no data are available on wintering sites of the species in Bulgaria. Although most localities were at altitudes below 300 m, the highest number of individuals during summer
was found in mountain beech and mixed coniferous woodlands at an elevation between 800 m and 1450 m
a. s. l. During the swarming period, some individuals were found to make vertical migrations of ca. 770 m
between their roosts and the site of the capture. Myotis bechsteinii was found in Pleistocene and Early
Holocene deposits of only two caves in Bulgaria. However, it was one of the most abundant and common
species during that time. At present, conservation of mature forests (i.e. sustainable forest management),
maintenance of their connectivity and further planting of new forest clearings are considered the most
important factors that could promote the occurrence of the species.
Introduction
There are comparatively less records of bats from the southern (both eastern and western)
parts of Europe than those from its central and western parts. This is especially true for the
Bechstein’s bat, Myotis bechsteinii (Kuhl, 1817). This species is considered to be distributed
continuously in most Central European countries, but has a scattered distribution in Southern
Europe and further to the east. Its occurrence in the south and east was reviewed for Portugal
and Spain (Benzal & de Paz 1991), Italy (Vergari et al. 1998), Croatia (Dulić 1959, Kovačić &
Dulić 1989), Slovenia (Kryštufek & Cerveny 1997), former Yugoslavia (Petrović et al. 1985),
Albania (Uhrin et al. 1996), Bulgaria (Benda et al. 2003, Petrov 1997), Romania (Nagy et al.
2005), Greece (Hanák et al. 2001, Helversen & Weid 1990), European Turkey (Kahmann 1962,
Furman & Özgül 2004), Asia Minor (Benda & Horáček 1998, Albayrak 2003), the Caucasus
region (Rakhmatulina 1990, Tsytsulina 1999) and by DeBlase (1980) who provided data
from the easternmost locality of the species in Iran. The Bechstein’s bat is considered a typical
species of the temperate humid zone whose range is centred mainly in the western part of the
Palaearctic region (Horáček et al. 2000). The distribution of the species is only well known in
179
the western and northern parts of its range. Southeastern records are often quite isolated from
the main distribution range. Therefore, additional information on the occurrence at the margins
of its range is needed to understand the recent scattered distribution of Myotis bechsteinii in
Europe. Furthermore, new records coming from the poorly studied regions, incl. Bulgaria, may
allow us to better understand what limits Bechstein’s bats in their occurrence. Besides depicting
precise maps of its distribution, chorological data are essential for the analysis of the species’
ecological requirements. This is an important prerequisite to design up-to-date national and
international conservation plans for the Bechstein’s bat, which has been listed as vulnerable in
the ‘Red List of Threatened Species’ since 1994 (IUCN 2004).
Material and Methods
This review is based mainly upon literature sources. However, new field records from the period 1997-2006
were added. Bechstein’s bats were caught at cave or mine entrances, above streams and in forest clearings using 3 m, 6 m or 12 m long mistnets. At four sites (7, 18, 24, and 33), radio-telemetry (Regal 2000
telemetry receiver, Titley electronics Ltd.; LB-2 transmitter, Holohil Systems Ltd.) was used to discover
the roosts of the tagged individuals. All captured bats were measured, aged, ringed and released afterwards.
Additionally, since 1999 a small piece of the bat’s wing membrane was collected, using a sterile biopsypunch for future DNA analyses. All bats were captured under the license of the Ministry of the Environment
and Waters (Bulgaria) and permissions from the local departments of forests (2001–2006).
Results
Review
of the records from Bulgaria
Heinrich (1936) was the first to record the Bechstein’s bat in the Balkans and particularly on
the Bulgarian Black Sea coast. Among other mammals, he reported six bat species, including
Myotis bechsteinii, for the first time for the Bulgarian fauna. Later on, Hanak & Josifov (1959)
added one locality, showing that the Bechstein’s bat also occurs in the mountains (Rila Mts.).
Beron (1961) found that this species hibernates in Bulgarian caves. After several field trips
to Bulgaria, Czech zoologists reported new localities thereby showing that the species is not
as rare as previously assumed (Horáček et al. 1974, Benda et al. 2003). Studies by Beshkov
(1993) raised the number of Bulgarian localities to eight. Petrov (1997) summarized all previous
records and added 4 new sites. Some recent regional bat surveys (Ivanova 1998, Pandurska &
Beshkov 1998b, Pandurska et al. 1999, Petrov 2001, Popov et al. 2006) reported Bechstein’s
bats from eight new localities. The most recent review of the distribution of bats in Bulgaria
(including M. bechsteinii) was published by Benda et al. (2003).
Only few papers deal with the historical evidence of this bat species in Bulgaria. Wołoszyn
(1982) first reported it in cave deposits from the Upper Pleistocene. Until today there has been
no contemporary record of the Bechstein’s bat from this locality (Bacho Kiro cave), although
environmental conditions are still suitable. Popov (2000) found abundant remains of this species in sediments dated to originate from the Upper Pleistocene and Holocene (cave 15 and 16,
Karlukovo village). He also suggested that the species was common and widely distributed in
periods with mild and humid climate, when forests still covered larger areas of the Balkans
and elsewhere. Recent occurrence of the Bechstein’s bat in the caves close to Karlukovo was
supported with field data by Benda et al. (2003).
In summary, the current knowledge of the distribution of Myotis bechsteinii shows that in
Bulgaria, the species inhabits diverse habitats, covering a broad altitudinal range. However, at
180
most places only single individuals were caught. A higher population density was estimated
for some of the localities (e. g., 7, 17, 18, and 33, see below) but only three maternity colonies
have been found so far in Bulgaria.
D i s t r i b u t i o n o f t h e B e c h s t e i n’ s b a t i n B u l g a r i a
The Bechstein’s bat is known to occur at 34 localities (33 UTM squares of 10×10 km) from
sea level up to 1650 m a. s. l. (Fig. 1, Table 1). All records before 1971 were more or less accidental findings. With few exceptions, density of the records from the mountainous regions
(e.g. Stara Planina, Rhodopes, Pirin) is higher compared to the scattered distribution in the
lowlands. There are no records of Myotis bechsteinii from the Bulgarian part of Dobrogea or
the upper parts of the Thracian plain where open agricultural landscape is prevailing. Occurrence of the species in the lowlands is mostly associated with larger forested areas (e.g. Strandja
Mts., 6, 7, 24, 26, and 34) or presence of a diverse mosaic of habitats (e.g. 10, 19, 21, 23, etc.).
In many regions, the species has not yet been found or has been only occasionally proved. For
example, no records are available from the northern slopes of the Rhodopes, eastern slopes of
the Pirin and Rila, Slavyanka, Belasitsa and other mountain ranges, which all offer suitable
environmental conditions.
Fig. 1. Records of the Bechstein’s bat (Myotis bechsteinii) in Bulgaria (1935–2006), see Table 1 for explanation of numbers. Shaded are mountains above 1200 m a. s. l.
Obr. 1. Nálezy netopýra velkouchého (Myotis bechsteinii) v Bulharsku (1935–2006), vysvětlení čísel viz
tab. 1. Stínována jsou horská území nad 1200 m n. m.
181
182
district
site
m a. s. l. method
m f
UTM
NH75
Heinrich 1936
Hanak & Josifov 1959
NH78
Hanak & Josifov 1959
GM18
Beron 1961
FP33
Horáček et al. 1974
KG71
Benda et al. 2003
KG71
B. Petrov, present paper
KG71
B. Petrov, G. Kerth & KG71
B. Koenig, present paper
B. Petrov, present paper
KG71
Benda et al. 2003
NG69
B. Petrov, G. Kerth &
B. Koenig, present paper
NG69
B. Petrov, G. Kerth & D. NG69
Dechmann, present paper
B. Petrov, present paper
NG69
B. Petrov & G. Kerth, NG69
present paper
B. Petrov & G. Kerth, NG69
present paper
B. Petrov & T. Stoyanov, NG69
present paper
B. Petrov & T. Stoyanov, NG69
present paper
B. Petrov, G. Kerth & T. NG69
Ivanova, present paper
B. Petrov, G. Kerth & T. NG69
Ivanova, present paper
Benda et al. 2003
KH68
Benda et al. 2003
KH78
Benda et al. 2003
KH68
Benda et al. 2003
KH68
Benda et al. 2003
KH68
date reference
1 Kamtchiya Dolni Chiflik lower course of the river
10 shot-gun 1 1 June 1935
2 Varna
Varna
pallace Evksinovgrad
10 hand
1 2 3. 8. 1935
3 Samokov
Sofia
Borovetz
1350 hand 18. 7. 1950
4 Belogradchik Montana
Gornata propast cave
600 hand
1
6. 2. 1960
5 Yagodina
Devin
Yagodinskata peshtera cave 1015 mist-net 1
2. 8. 1971
5 Yagodina
Devin
Yagodinskata peshtera cave 1015 mist-net 1 15. 8. 1978
5 Yagodina
Devin
Yagodinskata peshtera cave 1015 mist-net 1
4. 8. 1997
5 Yagodina
Devin
Yagodinskata peshtera cave 1015 mist-net 5 14. 9. 2001
5 Yagodina
Devin
Yagodinskata peshtera cave 1015 mist-net 2 17. 9. 2005
6 Primorsko
Bourgas
Arkutino swamp
10 mist-net 2 6. 6. 1972
7 Primorsko
Bourgas
Ropotamo Reserve, 10 mist-net 1 12. 9. 2001
Veliov vir
7 Primorsko
Bourgas
Ropotamo Reserve
10 mist-net 3 20. 4. 2002
7 Primorsko
Bourgas
Ropotamo Reserve
10 bat-box 2
4. 6. 2002
7 Primorsko
Bourgas
Ropotamo R., entrance gate
5 mist-net 1 1 25. 4. 2003
7 Primorsko
Bourgas
Ropotamo Reserve, 10 funnel trap 5 26. 4. 2003
Veliov vir
7 Primorsko
Bourgas
Ropotamo Reserve, 10 mist-net 1 25. 9. 2003
Veliov vir
7 Primorsko
Bourgas
Ropotamo Reserve, 5 mist-net 1
7. 5. 2004
entrance gate
7 Primorsko
Bourgas
Ropotamo Reserve, 5 mist-net 1 1 11. 6. 2004
entrance gate
7 Primorsko
Bourgas
Ropotamo Reserve, mouth
3 funnel trap 11 15. 6. 2004
of the Ropotamo river
8 Karlukovo Cherven bryagrocky amphitheatre
200 mist-net 1 15. 6. 1977
8 Karlukovo Cherven bryagrocky amphitheatre
200 mist-net 1
6. 8. 1978
8 Karlukovo Cherven bryagcave behind the monastery
200 mist-net 2
8. 8. 1978
8 Karlukovo Cherven bryagcave behind the monastery
200 mist-net 1
9. 8. 1978
8 Karlukovo Cherven bryagcave in the monastery
200 mist-net 4 9. 8. 1978
No.village
Table 1. List of record localities of the Bechstein’s bat (Myotis bechsteinii) in Bulgaria (1935–2006)
Tab. 1. Přehled lokalit nálezů netopýra velkouchého (Myotis bechsteinii) v Bulharsku (1935–2006)
183
9 Krivnya
Razgrad
Bozkova dupka cave
200 hand
1 15. 3. 1989
1 0 Buhovtsi
Targovishte Prilepnata dupka cave
200 hand
1 25. 4. 1989
11 Leshko
Blagoevgrad Leshtanskata peshtera cave 1000 mist-net 1
7. 6. 1996
12 Gorna Bela Varshets
mine
1000 mist-net 1
3. 4. 1996
13 Breze
Svoge
Travninata cave
1050 mist-net 1 15. 9. 1996
14 Gintzi
Godech
Dinevata peshtera cave
1200 mist-net 1
5. 9. 1994
15 Divchovoto Teteven
Grazdenitza cave
800 mist-net 1 28. 9. 1997
15 Divchovoto Teteven
Grazdenitza cave
800 mist-net 3 19. 9. 2001
16 Karnare
Karlovo
Mazata cave
1350 mist-net 1 25. 9. 1997
17 Vidima
Aprilitsi
Pleven hut, Vodnite dupki c. 1400 mist-net 9 15. 8. 1997
17 Vidima
Aprilitsi
Pleven hut, Vodnite dupki c. 1400 mist-net 101 28. 8. 2001
17 Vidima
Aprilitsi
Pleven hut, Vodnite dupki c. 1400 mist-net 4 16. 8. 2003
18 Bov
Svoge
Izdremets, mine 1450 mist-net 5
11. 9. 1998
18 Bov
Svoge
Izdremets, mine 1450 mist-net 7 14. 9. 1999
18 Bov
Svoge
Izdremets, mine 1450 mist-net 9 2 16. 9. 2001
18 Bov
Svoge
Izdremets, mine 1450 mist-net 2 2. 10. 2001
18 Bov
Svoge
Izdremets, mine 1450 mist-net 2
3. 8. 2002
18 Bov
Svoge
Izdremets, mine 1450 mist-net 2 2 2. 9. 2002
18 Bov
Svoge
Izdremets, mine 1450 mist-net 3 18. 9. 2002
18 Bov
Svoge
Izdremets, mine 1450 mist-net 10 22. 8. 2003
18 Bov
Svoge
Izdremets, mine 1450 mist-net 3 17. 9. 2003
18 Bov
Svoge
Izdremets, mine 1450 mist-net 5 19. 9. 2003
18 Bov
Svoge
Izdremets, mine 1450 mist-net 1 22. 10. 2003
19 Ribino
Kroumovgrad Samara cave
340 mist-net 1 20. 4. 1995
Beshkov 1993
MJ54
Beshkov 1993
MH89
Pandrurska & Beshkov FM64
1998a
Pandurska & Beshkov FN98
1998b
Pandurska & Beshkov FN86
1998b
Pandurska 1999
FN77
Ivanova 1998
KH64
B. Petrov, G. Kerth & KH64
B. Koenig, present paper
Ivanova 1998
LH04
Ivanova 1998
LH23
Schunger et. al. 2004
LH23
Schunger et. al. 2004
LH23
R. Pandurska & V. Beshkov FN97
present paper
B. Petrov, V. Beshkov & Y. FN97
Gorelov, present paper
B. Petrov, G. Kerth & FN97
B. Koenig, present paper
B. Petrov & S. Beshkov, FN97
present paper
B. Petrov & V. Beshkov, FN97
present paper
B. Petrov, present paper
FN97
B. Petrov, present paper
FN97
B. Petrov & V. Beshkov, FN97
present paper
B. Petrov & T. Stoyanov, FN97
present paper
B. Petrov & T. Stoyanov, FN97
present paper
B. Petrov & T. Stoyanov, FN97
present paper
Petrov 1997
LF78
184
district
site
m a. s. l. method
m f
UTM
LF78
C. Dietz & I. Schunger,
present paper
Petrov 1997
FN97
Petrov 1997
FM73
Petrov 2001
FM73
Petrov 1997
KH95
Benda et al. 2003
MG55
B. Petrov, present paper
NG78
B. Petrov, G. Kerth & NG69
T. Ivanova, present paper
Benda et al. 2003
KH58
B. Petrov & G. Stoyanov, KH58
present paper
Benda et al. 2003
NG26
C. Dietz & I. Schunger,
NG26
present paper
Benda et al. 2003
GM32
B. Petrov, present paper
FM92
B. Petrov, present paper
FM91
B. Petrov, present paper
FM91
C. Dietz & I. Schunger,
LJ23
present paper
Benda et al. 2003
LG90
E. Tilova, present paper
NH45
B. Petrov & G. Kerth,
GN06
B. Petrov & G. Kerth,
GN06
B. Petrov & G. Kerth,
GN06
B. Petrov & G. Kerth,
GN06
B. Petrov & G. Kerth, GN06
all five records by the present paper
Popov et al. 2006
NG85
date reference
1 9 Ribino
Kroumovgrad rocky bridge 320 hand
1 18. 9. 2002
20 Lakatnik
Svoge
Svinskata dupka cave
500 mist-net 1 24. 8. 1995
21 Kresna
Blagoevgrad Kresna gorge, on the road
200 hand
1 5. 10. 1995
21 Kresna
Blagoevgrad Kresna gorge, Sheitan dere 200 mist-net 1
2. 7. 1997
22 Golyama Jelyazna, Troyan Toplya cave
700 hand
1
2. 2. 1997
23 Ustrem
Topolovgrad Bozkite cave
200 mist-net 2 10. 4. 1998
24 Primorsko
Burgas
residence Perla
10 mist-net 1 18. 4. 1998
24 Primorsko
Burgas
oak forest 95 mist-net 1 14. 6. 2004
25 Brestnitsa
Yablanitsa
Seeva dupka cave
500 mist-net 1
1. 5. 1999
25 Brestnitsa
Yablanitsa
Seeva dupka cave
500 mist-net 4 15. 5. 2003
26 Mladezko
Malko TarnovoLeyarnitzata cave ? 4
160 mist-net 1 25. 8. 1999
26 Mladezko
Malko TarnovoLeyarnitzata cave ? 4
160 mist-net 2
11. 6. 2003
27 Ribnovo
Gotse DeltchevManoilovata peshtera cave 1000 mist-net 2 22. 6. 2000
28 Kresna
Blagoevgrad Peshterata, mine 1250 mist-net 2 1 25. 9. 2001
29 Ilindentsi
Sandanski
Sharaliiskata peshtera cave 1650 hand
1 1 7. 4. 2002
29 Ilindentsi
Sandanski
Sharaliiskata peshtera cave 1650 mist-net 1 25. 6. 2002
30 Muselievo
Nikopol
Nanin kamuk cave
140 mist-net 1 10. 6. 2002
31 Dolno Cherkovishte, HaskovoSedemte peshteri-Oreshari c. 320 observ. 1 30. 9. 2003
32 Golitsa
Dolni Chiflik oak forest (Quercus cerris) 300 mist-net 4 13. 5. 2003
33 Gabrovnitsa Svoge
Sedemte prestola Monastery 650 mist-net 3 7. 5. 2003
33 Gabrovnitsa Svoge
Sedemte prestola Monastery 650 funnel trap 5 9. 5. 2003
33 Gabrovnitsa Svoge
Sedemte prestola Monastery 650 mist-net 1 15. 4. 2006
33 Gabrovnitsa Svoge
Sedemte prestola Monastery 650 funnel trap 9 16. 4. 2006
33 Gabrovnitsa Svoge
Sedemte prestola Monastery 650 funnel trap 2 19. 4. 2006
34 Sinemorets Tsarevo
Silistar Reserve
3 mist-net 1 August 1998
No.village
Table 1. (continued)
Tab. 1. (pokračování)
Table 2. Vertical distribution of the localities of the Bechstein’s bat (Myotis bechsteinii) in Bulgaria
(1935–2006)
Tab. 2. Vertikální rozšíření lokalit nálezů netopýra velkouchého (Myotis bechsteinii) v Bulharsku
(1935–2006)
altitude (m)
sites (see Table 1)
0–300
301–600
601–900
901–1200
1201–1500
1501–1800
1, 2, 6, 7, 8, 9, 10, 21, 23, 24, 26, 30, 32, 34
4, 19, 20, 25, 31
15, 22, 33
5, 11, 12, 13, 14, 3, 16, 17, 18, 28
29
total sites
14
5
3
6
5
1
rel. share [%]
41.2
14.7
8.8
17.6
14.7
2.9
New data
In this study I report new data from eight previously unknown localities. The species was proved
to occur in the Pirin Mts. (loc. 28, 29), close to the Danube (loc. 30), in the western Stara Planina
east of the Iskar river gorge (loc. 18, 33), in the Ropotamo Reserve at the Black Sea coast (loc.
7, 24) and in the eastern Stara Planina Mts. (loc. 32). At the localities Ropotamo Reserve (Loc.
7), Vodnite dupki cave (loc. 17), Izdremets (loc. 18) and Sedemte Prestola monastery (loc. 33),
Bechstein’s bats were captured more than twice. These sites (among others) were therefore
supposed to hold higher population density and bat boxes (Scwengler 2 FN) have recently been
installed at localities No. 7, 18 and 33 (2001–2002, Petrov & Kerth unpubl.). Localities No.
18, 28 and 29 broaden the altitudinal range where Myotis bechsteinii was found in Bulgaria
(see below). Furthermore, at four localities that were already known before (5, 15, 19, and 25),
the presence of the species was confirmed.
While the majority of the other papers (cf. Benda et al. 2003) deal with the geographic distribution of the species, the present survey analyses some details of its occurrence. To provide
better description of the habitats where the species was found, 32 localities (out of 34 known)
were personally visited. Among many other bat localities in the country visited by the author,
mistnetting was also attempted at some previously known sites (e.g. loc. 8, 11, 12) but no
Bechstein’s bats were captured. Some of the known sites were visited in winter as well (e.g. loc.
9, 14, 17, 20, 22, 27, 29) but the species was not found. The maximum number of Bechstein’s
bats captured per night was 11 specimens (loc. 18, 17; Schunger et. al. 2004). Since its first
discovery in 1935, 55 females, 141 males and 2 specimens (loc. 3) of unknown sex have been
captured. Out of these numbers, 129 males and 46 females (i.e. 89% of the total number) were
caught between 1994 and 2006 (i.e. in a period when intensive regional bat surveys in Bulgaria
were carried out mostly by local researchers including the author of this paper).
Altitudinal distribution
Most of the localities (n=14; 41.2%) are situated between sea level and 300 m a. s. l. (Table 2).
This high rate of occurrence at lowland sites is partly due to the fact that numerous bat surveys in
Bulgaria were carried out in this altitudinal range. On the other hand, Bulgarian bat fauna reaches
its greatest species diversity per area between 100 m and 300 m (Pandurska 1996, Petrov 2001,
Popov & Ivanova 1995) and regional fauna can consist of 17–20 bat species, which inhabit these
185
Table 3. List of the localities of the Bechstein’s bat (Myotis bechsteinii) in Bulgaria sorted by altitude.
Composition of the dominant vegetation cover is after Bondev (1991). Temporal water presence is abbreviated as “temp.”
Tab. 3. Přehled lokalit nálezů netopýra velkouchého (Myotis bechsteinii) v Bulharsku řazený podle
nadmořské výšky. Složení dominujícího vegetačního pokryvu podle Bondeva (1991). Nestálá přítomnost
vody je naznačena zkratkou “temp.”
No. village, site
m a. s. l. dominant vegetation cover
34 Sinemorets, Silistar Reserve
3
1 Kamtchiya, river
10
2 Varna, Evksinovgrad pallace
10
6 Primorsko, Arkutino swamp
10
7 Primorsko, Ropotamo Reserve
10
24 Primorsko, Rerla residence
10–95
30 Muselievo, Nanin kamuk cave
140
26 Mladezko, Leyarnitzata N 4 cave
160
8 Karlukovo, caves, ridge 200
9 Krivnya, Bozkova dupka cave
200
10 Buhovtsi, Prilepnata dupka cave
200
21 Kresna, Kresna gorge
200
23 Ustrem, Bozkite cave
200
32 Golitsa, oak forest (Q. cerris)
300
31 Dolno Cherkovishte, Sedemte peshteri- 320
Oreshari cave
19 Ribino, Samara cave
340
20 Lakatnik, Svinskata dupka cave
500
25 Brestnitza, Seeva dupka cave
500
4 Belogradchik, Gornata propast cave
600
33 Gabrovnitsa, Sedemte prestola Mon.
650
22 Golyama Jelyazna, Toplya cave
700
15 Divchovoto, Grazdenitza cave
800
11 Leshko, Leshtanskata peshtera cave
1000
12 Gorna Bela rechka, mine
1000
27 Ribnovo, Manoilovata peshtera cave
1000
5 Yagodina, Yagodinska peshtera cave
1015
186
water m f
Quercus cerris, Q. frainetto river 1
with Mediterranean elements
Quercus cerris, Q. freinetto river 1 1
Quercus pubescnes, Q. sea
1 2
virgiliana, C. orientalis
Quercus spp., Acer campestre swamp 2
Quercus spp., Acer campestre river 1018
Querceta freinetti with sea
1 1
Mediterranean elements
arable lands, Quercus spp.,
Carpinus orientalis
river 1
Quercus cerris, Q. frainetto
with Mediterranean elements river 3
Paliureta spina-christi,
Quercus cerris, Q. frainetto river 5 4
Carpinus betulus, Q. cerris, river 1
Q. dalechampii
Carpinus betulus, Q. cerris, lake
1
Q. dalechampii
Platanus orientalis, Alnus
glutinosa
river 2
Qurcus dalechampii, Carpinus river 2
orientalis
Querceta polycarpae
river 4
Carpitneta orientalis with river 1
Mediterranean elements
river 2
Carpitneta orientalis with river 2
Mediterranean elements
Carpinteta orentalis
river 1
Carpinteta orentalis
temp. 5
Quercus cerris, Q. freinetto none 1
Fagus sylvatica moesiaca
river20
Fagus sylvatica moesiaca
river 1
Fagus sylvatica moesiaca
river 4
Quercus dalechampii (+ temp. 1
Carpinus orientalis)
Fageta sylvaticae
river 1
Pinus sylvestris, Fagus
sylvatica
river 2
Pinus sylvestris, Fagus river 10
sylvatica
13 Breze, Travninata cave
1050
14 Gintsi, Dinevata peshtera cave
1200
28 Kresna, Peshterata, mine 1250
3 Samokov, Borovetz 1350
16 Karnare, Mazata cave
1350
17 Vidima, Vodnite dupki cave
1400
18 Bov, Izdremets, mine 1450
29 Ilindentsi, Sharaliiskata peshtera cave 1650
Carpineta orientalis (+ none 1
Fagus sylvatica)
Carpineta orientalis (+
Fagus sylvatica)
river 1
Fagus sylvatica moesiaca
river 2
Picea abies, Fagus sylvatica river
(+ Pinus sylvestris)
Fageta sylvaticae
none 1
Fageta sylvaticae
river 23
Fagus sylvatica moesiaca, lake 49
Carpinus betulus
Pinus heldreichi, P. sylvestris none 2
1
1
4
1
areas seasonally or permanently. Higher population density of Bechstein’s bats in the lowlands
was reported also from Switzerland (Zingg 1982), Germany (Kerth 1997, Weishaar 1996) and
the Czech Republic (Červený & Bürger 1989). The high number of localities in the lowlands of
Bulgaria does not correlate with the species abundance. Considering all captured bats (n= 196),
the average number of individuals from the 22 localities below 1000 m a. s. l. is 4.3 specimens
per site, versus 8.2 from 12 localities between 1000 m and 1650 m (Table 1).
Two male Bechstein’s bats were caught and radio-tracked at the swarming site of Izdremets
(loc. 18) at 1450 m in August and September. On the next day after capture, both were found
to roost in beeches (Fagus sylvatica) at lower altitudes (630–680 m, loc. 33). These individuals thus migrated vertically about 770 m (2.7 km one way straight distance) for swarming
and turned back to their roosts. These observations are the first that demonstrate short-distance
vertical flights in the species.
The highest altitude at which Myotis bechsteinii was recorded in Bulgaria is 1650 m a. s. l.
(loc. 29). The species (1 female, 2 males) was encountered at this location twice in three months.
Other findings at comparable altitudes in central Spain (1500 m, Benzal & de Paz 1991), in the
Swiss Alps at Montreux (1560 m, Chapuisat & Ruedi 1993), Jumelles (grotte au Tichodrome,
1750 m) (Arlettaz et al. 1993) indicate that the species can be occasionally found near the
timberline. However in most of these high-altitude cases, bats probably went so high for swarming or feeding rather than for seasonal roosting. A skull was found at 1950 m (Pigna, cave No.
F 7-813) in Italy (Liguria, Amelio 1973). Holocene findings of the Bechstein’s bat in Austria
in caves from 1800 m (Bauer & Walter 1977) up to 2100 m (Spitzenberger 2001) show its
historic occurrence in this region. The subfossil Italian locality ‘Pigna’ together with the record
from 2100 m in Austria are the highest ever recorded localities for Myotis bechsteinii.
Seasonal occurrence and wintering
The highest number of Bechstein’s bats was captured in September, followed by August. At that
time, 100 males were caught (i.e. 71% of all males reported in this study) versus only 12 females. The highest number of females was recorded in late April (19 ind.) and in early June (16
ind.) due to the discovery of three breeding colonies (Loc. No. 7, 33) found by radio-tracking
of female bats. Bechstein’s bats were caught only twice in July (1950, 1997), though at the
187
locality No. 7 mistnetting in the forest was performed for 10 nights in 2002 and no Bechstein’s
bats were captured.
It is worth mentioning that since 1935, only two specimens have been found to hibernate in
caves (loc. 4: 6 February 1960, loc. 22: 2 February 1997). Both were male and were found in
crevices in the wall near the entrance. Only one of those records comes from the last ten years
even though bat research in the country has been relatively intensive during this period. Myotis
bechsteinii has never been observed during recent winter censuses of bats in numerous caves
(about 60), mines (about 10) and other underground roosts, which host hibernating colonies of
various vespertilionid species (cf. Benda et al. 2003).
Habitat and roost preferences
Forests and scrubs dominated mainly by Quercus spp., Carpinus spp. and occasionally by
Platanus orientalis cover all lowland localities (i.e. 0–600 m) where the species was found in
Bulgaria. In many cases (e.g. loc. 13, 17, 18, 28) the Bechstein’s bats were not caught in a roost
but at a swarming site. Thus our knowledge on the preferred habitats is biased by the seasonal
activity patterns.
Oak and oriental plane forests offer relatively thick trees and Bechstein’s bats are thought
to occupy mostly tree holes and crevices (cf. von Helversen & Weid 1990). Findings of the
Bechstein’s bat in the Carpinus forests, where suitable trees are much less common could suggest that the species uses alternative roosts such as caves, mines, bunkers, tunnels, rock fissures,
etc. Male specimens were occasionally recorded in a crevice between bricks of a rocky bridge
(loc. 19) and in a rock crevice (loc. 31).
Roosts of male Bechstein’s bats and two breeding colonies have recently been found in tree
holes (2001–2006, Petrov & Kerth, new records). In the lowland forest of the Ropotamo Reserve, most of the used roosts were discovered in the common maples (Acer campestre) and rarely
in Quercus cerris or Q. polycarpa. In the area of the Sedemte Prestola monastery (western Stara
Planina Mts.), where the third breeding colony was found, all roosts were in beeches (Fagus
sylvatica), though oak trees were also present. Single individuals were found in shallow cavities
in relatively slender trees (DBH = 13–20 cm) at the height of 0.7–5 m above the ground (Table
4). Roosting groups (5–55 ind.) were found in thicker trunks (DBH = 40–55 cm) at the height
of 5–10 m above the ground. A colony of 55 individuals has recently been observed emerging
at dusk in this region but none of the bats was captured and included in the present analysis.
This colony however is the largest known so far from the southeastern Europe.
Bechstein’s bats were found only once (4 June 2002) in a bat box (Schwegler 2FN) in the
Ropotamo Reserve. The two individuals were the first bats found in a bat box in Bulgaria, only
a year after the boxes were installed. Since then no Bechstein’s bats have been found in the boxes
(n=55) at this site but temporal roosting seems possible at times when no census was made.
The majority of the capture sites were situated next to a permanent water body: river (n=23),
sea (n=2), lake (n=2) and swamp (n=1). Only few of the localities lacked water (n=4) or had
only temporal water sources (n=2) (Table 3).
Climatic features of the localities: In Bulgaria, Myotis bechsteinii was found at sites with
high average diurnal temperature during summer (22–25 °C) and very little annual precipitation (550 mm and lower) (loc. 21, 23), as well as at sites with low summer diurnal temperature
(12–16 °C) and heavy precipitation (1000 mm and higher) (loc. 5, 17). Abundance was higher
at the latter sites, where temperatures are moderate and the climate is clearly humid throughout
the year.
188
Table 4. Tree roosts of the Bechstein’s bat (Myotis bechsteinii) in Bulgaria (2001–2006). CF – circumference of the trunk [cm]; DBH – trunk diameter at breast height [cm]; DG – distance to the ground [m]
Tab. 4. Stromové úkryty netopýra velkouchého (Myotis bechsteinii) v Bulharsku (2001–2006). CF – obvod
kmene [cm]; DBH – průměr kmene ve výši prsou [cm]; DG – vzdálenost od země [m]
site
date
tree species
Ropotamo (7)
13 September 2001
18 April 2002
20 April 2002
22 April 2002
23 April 2002
26 April 2003
27 April 2003
28 Sept. 2003
15 June 2004
Monastery (33)
September 2001
9 May 2003
16 April 2006
17 April 2006
18 April 2006
CF
DBH
DG
Acer campestre
Quercus cerris
Acer campestre
Acer campestre
Acer campestre
Quercus polycarpa
Acer campestre (dead trunk)
Acer campestre
Carpinus betulus
44 78
62
48
40
170
121
76
161
14
25 19
15
13
54
39
24
51
3.0
1.5
1.6 2.5 0.7 9.0 6.0 3.5 2.5 Fagus sylvatica
Fagus sylvatica
Fagus sylvatica
Fagus sylvatica
Fagus sylvatica
95
131
87
153
139
30
42
28
49
44
3.5 5.0 5.0 10.0 8.0 number of bats
1 male
1 male
1 male
1 male
1 male
6 females
1 female
1 male
11 females
1 male
5 females
10 females
1 female
55 individuals
Capture methods
Almost all specimens reported during the last 30 years in Bulgaria were caught using mistnets
at cave and mine entrances. Captures at cave mouths (20 cases) make up the majority of the
samples. The highest abundance of male bats was found during the swarming season (late
summer and early autumn), while mistnetting at caves and mines in old growth mesophile
mountain forests (e.g. loc. 17, 18).
In the recent years (after 2001), females from several breeding groups/colonies have been
caught at tree holes using a funnel trap. When the latter was set well, almost no escapes were
observed. Captures with bare hands happened only occasionally. Bechstein’s bat has been found
killed on the road only once in Bulgaria (Loc. No. 21), though many other species (some of
them in high numbers) have been found in a road mortality survey in the Kresna gorge (SW
Bulgaria, B. Petrov, unpubl.). The highest capture success of mistnetting in forests was proved
when the net was set above a stream or shallow riverbed. In the latter case, captures of males
were slightly prevailing (9 males, 7 females, loc. 7, 24, 33, and 34) compared to strongly male
dominated catches at the swarming sites (see below).
Discussion
Few studies on Myotis bechsteinii provide detailed data on the vegetation and habitat type in
the surroundings of its roosts. The existing data suggest that this bat is a typical inhabitant of
the old growth forests in western Europe (Kerth 1997, Schlapp 1990). In central parts of the
continent, the Bechstein’s bat also occurs in open river valleys with groves in their vicinity,
189
and in small woodland patches, which alternate with arable and parklands (Červený & Bürger
1989, Harmata 1969). In southern Europe, Myotis bechsteinii was found in more or less dense
xerophile forests of the Mediterranean type in Portugal and Spain (Benzal & de Paz 1991),
Greece (Hanák et al. 2001, von Helversen & Weid 1990) and Corsica (Libois & Franken
1981). In Bulgaria, the habitats vary with altitude, ranging from dry mixed lowland oak forests
to humid mountain beech and mixed coniferous woodlands. As Bechstein’s bats feed mostly
on non-airborne invertebrates and water independent flying insects (Dondini & Vergari 1999b,
Krochko 1990, Wolz 1993), the presence/absence of water (streams, lakes, etc.) probably does
not restrict the species distribution. On the other hand, water availability favours insect diversity,
which is presumed to promote the occurrence of bats.
The Bechstein’s bat is known as a non-migratory species. Most local movements cover less
than 10 km and the females exhibit extreme philopatry (Kerth et al. 2000, 2002). The longest
known dispersal flights reach 39 km (Haensel 1991), 43 km (Rudolf et al. 2004) and exceptionally up to 60 km (Kerth & Petit 2005). The maximum flights proved so far by radiotracking
in Bulgaria are between 2 and 3 km (loc. 7, 18, 24, and 33).
At the swarming sites with the highest capture rate in Bulgaria, more males than females were
captured. At loc. 17 only one of the 24 specimens was a female (4.2%). At loc. 18, only 4 out
of 54 specimens were females (7.5%). This could indicate higher mobility or longer dispersal
flights of the males during the pre- and postmating period compared to the females (cf. Kerth
et al. 2003).
The high number of records in late summer probably reflects both the bat research activity,
which usually is at its maximum during these months, and the fact that the intensity of bat swarming at caves and mines is highest during this period. The latter was proved in many countries
(cf. Parsons et al. 2003, Kerth et al. 2003), including Bulgaria (cf. Schunger et al. 2004).
The scarce winter records of Myotis bechsteinii in Bulgaria suggest that the species rarely
uses underground roosts in this part of its life cycle. In the Czech Republic (Šumava Mts.),
Bechstein’s bats were frequently found to hibernate in caves and galleries only during harsh
winters (Červený & Bürger 1989). In Great Britain, more records of this bat from underground
sites have been reported during severe winters (Harrington et al. 1995).
Findings of the Bechstein’s bat all over Europe are relatively rare compared to records of
the other species of the continental bat fauna (Baagøe 2001). Many of the recent peripheral
localities remain isolated from each other and presumably some marginal populations could
become allopatric. Only in a few regions of Central Europe (e.g. parts of Germany and the Czech
Republic) it occurs in higher population density and substantial signs of a dramatic decline are
not yet recognised (Červený & Bürger 1989, Kerth 1997, Schlapp 1990, Weishaar 1996). In
Bulgaria, the Bechstein’s bat ranks among the common bat species with a continuous distribution in regions with larger forest coverage (e.g. Strandja Mts. along the Black Sea coast). On
the other hand, only single individuals were caught in majority of the other sites. At present, the
species could be considered “rare but locally common” with regard to distribution and “vulnerable” with regard to its environmental sensitivity and low colonisation abilities.
Subfossil evidence from Austria (Bauer & Walter 1977, Rabeder 1972, Spitzenberger
2001), Great Britain (Yalden 1999), Bulgaria (Popov 2000), Czech Republic (Kowalski 1962),
Hungary (Topál 1959), France (Mein 1975, Sevilla 1990), Italy (Kotsakis & Petronio, 1980),
Poland (Kowalski 1956, Ochman 1999), Spain (Sevilla 1989) and Ukraine (Krochko 1990)
show that Myotis bechsteinii was one of the most common bat species in the Pleistocene and
Early Holocene deposits. Milder, humid climate and the presence of continuous deciduous wo190
odlands all over the continent and particularly in Bulgaria presumably favoured this abundance
especially during the Interpleniglacial and the Early Holocene (Peshev et al. 2004). Since then
the climate has changed many times and forests were cut in vast regions including clearance of
deciduous woodlands in Europe during the post-Neolithic (Yalden 1999). This probably led
to the present fragmentation of the species range and population decline.
One possible reason of the current rarity of the Bechstein’s bat in most parts of Europe may
be the limited distribution and size of old growth forests as well as their low connectivity.
Deforestation and unsustainable forest management practices led to a reduction of roost availability, which seems to be the most important environmental factor for the species occurrence
(Harrington et al. 1995, Hutson et. al. 2001). Another possible reason could be the concentration of bat research activities, which are not evenly spread over the species range. Although
the research coverage varies from year to year, it probably plays a significant role in accuracy
of the population assessments at local and national level.
The Bechstein’s bat is protected by law in at least 31 of the European countries, which have
joined the EUROBATS agreement, including Bulgaria. Considering the global fragmentation of forest habitats preferred by the species, its low population density and poor dispersal
abilities, Myotis bechsteinii was also classified as a “vulnerable” species in the new edition of
the Bulgarian Red Data Book (Petrov in press). Because of its sedentary life style, a “colonyorientated“ conservation approach was suggested in Germany (Meschede & Heller 2000).
Bat box projects in Europe, e.g. in Italy (Dondini & Vergari 1999a), Germany (Kerth 1997,
Taake & Hildenhagen 1989), United Kingdom (Schofield et al. 1997) and recently in Bulgaria
(Petrov & Kerth, unpublished) proved that Myotis bechsteinii is among the first bat species,
which occupy these artificial shelters, especially those that resemble woodpecker’s holes (e.g.
2 FN Schwegler’s boxes). Furthermore, restoration and planting of new forest clearings will
contribute to the connectivity of suitable habitats and thus to spreading of the Bechstein’s bat
at least in the centre of its range.
SOUHRN
První nález Myotis bechsteinii v Bulharsku pochází z roku 1935 a od té doby bylo zaznamenáno celkem
55 samic, 141 samců a dva jedinci neznámého pohlaví, avšak do současné doby byly v Bulharsku nalezeny
jen tři mateřské kolonie. Netopýr velkouchý je znám z Bulharska z celkem 34 lokalit (33 čtverců UTM)
ležících v nadmořské výšce od hladiny moře po 1650 m. Jen dva jedinci (samci) byli dosud nalezeni
zimující v jeskyních, jiné nálezy ze zimovišť nejsou známy. Většina lokalit leží ve výšce do 300 m n. m.,
nejvyšší počet jedinců byl ale zaznamenán v horských bukových a smíšených lesích ve výšce mezi 800
a 1450 m n. m. V průběhu “swarmovacího” období byly zaznamnány vertikální migrace o zhruba 770 m
mezi denním úkrytem a místem odchytu. Myotis bechsteinii byl doložen z pleistocénních a spodnoholocénních vrstev dvou jeskyní v Bulharsku, ovšem v těchto údobích představoval jednoho z nejhojnějších
a nejběžnějších netopýrů. Ochrana původních lesních porostů (tj. udržitelné lesnické hospodaření), udržování jejich propojení a obnova hospodářských lesů jsou faktory v současné době nejvíce považovány
za nezbytné pro ochranu druhu v Bulharsku.
Acknowledgments
I wish to thank T. Ivanova (NMNH, Sofia), T. Stoyanov (Sofia), A. Gueorguieva (BRPG, Sofia), B. Barov
(BSPB, Sofia), N. Simov (NMNH, Sofia), R. Pandurska-Whitcher (Sofia), C. Dietz and I. Schunger
(Tübingen), E. Tilova (Green Balkans) as well as all other colleagues who provided material or helped me
during the field research. V. Beshkov (Institute of Zoology, Sofia) offered logistic support and assisted my
191
activities with endless enthusiasm. V. Popov (Institute of Zoology, Sofia) gave some valuable suggestions
on the manuscript. I also thank A. Hutson (BCT, London), G. Dondini (Florence), H. J. Baagøe (Copenhagen), I. Rakhmatulina (Baku), K.-G. Heller (Magdeburg), M. Paunović (Belgrade), M. Zagmajster
(Ljubljana), H. Schofield (The Vincent Wildlife Trust, London), P. Boye (Bonn) and K. Safi (Zürich) for
providing me with some of the literature sources. Field travels between 2001 and 2004 were supported
by the Swiss National Science Foundation (c/o SCOPES program, grant No. 7BUPJ062292). My sincere
thanks go to G. Kerth and B. König (Zürich), who took part in two of the research trips and kindly shared
their experience and literature on the Bechstein’s bat. D. Dechmann (Zürich) and G. Kerth commented
earlier versions of the manuscript and especially D. Dechmann generously provided linguistic help.
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Lynx (Praha), n. s., 37: 197–200 (2006).
ISSN 0024–7774
First record of Myotis blythii in Poland (Chiroptera: Vespertilionidae)
Pierwsze stanowisko Myotis blythii w Polsce (Chiroptera: Vespertilionidae)
Krzysztof Piksa
Cracow Pedagogical University, Institute of Biology, Podbrzezie 3, PL–31-054 Kraków, Poland;
[email protected]
received on 30 June 2006
Abstract. The first record of M. blythii in Poland is described. It slightly extends the known distribution
range of the species in Central Europe northwards.
Three large species of the genus Myotis live in Europe. Two of them, the greater mouse-eared
bat, M. myotis (Borkhausen, 1797), and the lesser mouse-eared bat, M. blythii (Tomes, 1857),
are widely distributed throughout southern and central Europe. The distribution range of the
third one, the Maghrebian mouse-eared bat, M. punicus Felten, 1977, is restricted to several
Me­di­ter­ra­ne­an islands, besides its main range in north-western Africa (Castella et al. 2000,
Güttinger et al. 2001, Topal & Ruedi 2001). In Poland, M. myotis is widespread and its continual
distribution range covers the southern and partly the western and central parts of the country. In
addition, scattered records have been reported from most of the rest of Poland (Sachanowicz et
al. 2006). However, M. blythii has not yet been reported from Poland. In Europe, the northern
margin of its range reaches roughly 50 °N in the Czech Republic and Slovakia (Topál & Ruedi
2001), so the species has been known from near the southern border of Poland. Therefore, under
favourable circumstances, the species was likely to occur also in Poland.
During the field studies on swarming activity of bats in the Tatra and Beskidy Mts. carried
out in 1999–2005, one individual of M. blythii was recorded. An adult male of M. blythii was
cap­tu­red on 12 October 2005 into a net installed inside the Czarna Cave, several dozens of
metres below the northern cave entrance at the altitude of 1294 m, during an in-flight. This site
is situated in the Organy Massif, in the Kościeliska Valley. The total length of known corridors
of this cave is about 6,500 m, with denivelation of 303.5 m. It has three openings situated at
1326 m, 1294 m and 1404 m a. s. l. (Grodzicki et al. 1995). It is the largest bat hibernaculum
in the Tatra Mts (Piksa & Nowak 2000).
In order to identify the bat species, the basic morphometric features enabling to dif­f­e­ren­ti­a­te
between M. myotis and M. blythii were considered: forearm length (FaL), ear length (EaL),
and ear width (EaW) (see Arlettaz et al. 1991, Arlettaz 1995, Dietz & von Helversen
2004). Accro­ding to Arlettaz et al. (1991), the Z function was calculated: Z = 0.1084×FaL +
1.4166×EaL–40.5907 (if Z< 0, then M. blythii).
At first sight, the captured bat seemed smaller and more subtle than M. myotis. As far as the
coloration is concerned, it was similar to the earlier and later captured greater mouse-eared bats,
with a little bit lighter fur on the belly. The white spot on the hair on the head, which is typical
197
of some M. blythii males, was not visible. The ears were narrower and thinner. The teeth of
the captured bat were significantly rubbed off (e.g. the top left canine was broken in the distal
part), indicating that it was an adult specimen.
The right FaL was 58.1 mm, the left FaL 58.9 mm. The EaW and EaL of the right ear were 8.9
and 22.6 mm, respectively; the left EaW was 8.9 mm. The left EaL was not measured as its tip
was slightly cut. The lengths of the forearms and ears of the individual caught were com­pa­red
with the dimensions of other greater mouse-eared bats captured in the years 2004–2005 during
the research on swarming activity in the Polish parts of the Carpathians (Fig. 1). The value of
the formula compiled by Arlettaz et al. (1991) was –2.28 in this bat, i.e. the value typical for M.
blythii, and differed from the values found in the captured individuals of M. myotis (0.21–3.34).
In this respect the examined bat differed remarkably from the other mou­se-eared bats captured
at that time in the Polish Carpathians.
The Czarna Cave is the first locality of M. blythii in Poland. However, the species does not
seem to be a permanent element of the Polish fauna. Most probably, this was just an ac­ci­den­
tal­ly present individual whose occurrence, especially in the Polish Tatra Mts, is surprising. On
the southern side of the Tatra Mts. in Slovakia, the species has been recorded only at two sites
during hibernation and at lower altitudes: in winter 1964 in the Belianska Cave (890 m a. s. l.)
(Mošanský & Gaisler 1965, Gaisler & Hanák 1972) and in the winter season 1994/1995
in the Lučivianska Cave (800 m a. s. l.). However, the accuracy of identification of the latter
record has been doubted (Pjenčák et al. 2003). Therefore, the last certain record of this species in the Slovak Tatra Mts. was made more than 40 years ago. Since then, despite in­ten­si­ve
Fig. 1. Scatter plot of the forearm length against the ear length (n=63) recorded in Myotis myotis (triangles)
and M. blythii (square) captured in 2004 and 2005 in the Polish Carpathians.
Rys. 1. Relacja między długością przedramienia (Forearm length) a długością ucha (Ear length) (n=63)
u Myotis myotis (trójkąty) i M. blythii (kwadrat) schwytanych w 2004 i 2005 roku w okresie swarmingu, w Tatrach, Beskidzie Wy­spowym, Beskidzie Sądeckim, Beskidzie Niskim, Pogórzu Ciężkowickim
i Podhalu.
198
research, especially in the recent years, both in the periods of activity and of hibernation, the
species has not been found (Pjenčák et al. 2003). In other parts of Poland, in the regions close
to the northern range of M. blythii, where its occurrence should be possible, it has not yet been
recorded (author’s unpubl. data, Paszkiewicz et al. 1998, Postawa & Wołoszyn 2000, Węgiel
et al. 2001, 2004). However, it cannot be excluded that single specimens of this spe­ci­es might
occasionally occur in Poland.
Streszczenie
Dorosły samiec nocka ostrousznego Myotis blythii został odłowiony 12 października 2005 roku w Tatrach Polskich w Jaskini Czarnej (1294 m n. p. m.). To pierwsze stanowisko tego gatunku nietoperza
w Polsce.
Acknowledgements
I would like to thank the members of the Caving Club of University of Science and Technology in Cracow,
as well as Elżbieta Wiejaczka and Wojciech Gubała for their help in the field. This study was supported
by the grant from the State Committee and Scientific Research.
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Bieszczady Moun­ta­ins]. Monogr. Bieszczad., 9: 91–101 (in Polish, with a summary in English).
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F. (ed.): Handbuch der Säugetiere Europas. Band 4: Fledertiere. Teil I. Chiroptera I. Rhinolophidae,
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Węgiel A., Paszkiewicz R. & Szkudlarek R., 2001: Nietoperze Beskidu Wyspowego, Beskidu Sądeckiego, Beskidu Niskiego i Pogórza Karpackiego – letnie schronienia nietoperzy w budynkach [Bats of
Beskid Wyspowy, Beskid Sądecki, Beskid Niski and Pogórze Karpackie – summer roosts in buildings].
Nietoperze, 2: 75–84 (in Polish, with a summary in English).
Węgiel A., Szkudlarek R. & Gottfried T., 2004: Wyniki odłowów nietoperzy przy otworach niektórych
jaskiń w Beskidach [Species composition, activity and population structure of bats netted in summer
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200
Lynx (Praha), n. s., 37: 201–228 (2006).
ISSN 0024–7774
Areal and altitudinal distribution of bats in the Czech part of the Carpathians (Chiroptera)
Plošné a výškové rozšíření netopýrů v české části Karpat (Chiroptera)
Zdeněk Řehák
Institute of Botany and Zoology, Masaryk University, Kotlářská 2, CZ–611 37 Brno, Czech Republic;
[email protected]
received on 20 December 2006
Abstract. All available bat records from the Outer Carpathians, situated in the eastern part of the Czech
Republic, were summarized, divided into old (before 1956), winter and summer records and evaluated
with respect to orographical units, quadrats of grid maps and elevation. In total, 20 bat species and/or
two pairs of sibling species were recorded in 9 orographical units and 41 quadrats covering the area
under study. Hitherto, thirteen species of bats were recorded as hibernating and in 10 species maternity
colonies were found in the Czech Carpathians. The areal distribution of particular species was presented
via grid maps. Plecotus spp. (50.0% of the studied area), Myotis myotis (46.7%) and M. daubentonii
(40.0%) can be considered the most distributed of them. Altitudinal distribution of sites where bats were
recorded, expressed as medians of their elevation, shows that P. austriacus significantly preferred low
elevation both in the winter and summer periods (259 and 298 m above sea level, respectively) while the
hibernacula of the M. mystacinus group and M. nattereri prevailed at higher elevations (872 m and 1050
m, respectively). During the non-hibernation period the highest medians of elevation were recorded in
M. brandtii and again in M. nattereri (765, resp. 665 m a. s. l.). The causes of preferences for such high
elevated sites are discussed.
Introduction
The knowledge of the distribution of bats enables to estimate their species diversity over
reasonable geographical units and, consequently, to assess the ecological value of individual
segments within a given territory. At the same time, bats’ distribution reveals the differences in
conditions favourable to the life of these mammals among various areas of the same territory.
Differences in the composition and abundance of bat communities then reflect different habitat
demands of individual species the communities consist of. Information on the distribution of
bats, therefore, represents a starting point for both synecological and autecological studies of
the respective chiropteran fauna.
Faunistic review of all available records of bats in a particular area is a condition sine qua
non to analyse the distribution of the respective bat assemblage or assemblages. Within the
territory of the Czech Republic, the Carpathians were relatively neglected in this respect until
quite recently. The history of chiropterological research there, not very ample so far, was dealt
with in previous papers by the author (Řehák 1998, 2001). The first faunistic review of bats
on the territory of the Czech (Moravian-Silesian) Carpathians was published by Řehák (1998).
201
In the present paper, the data of the former one gathered up to 1998 are supplemented by new
records till 2006, both published and unpublished, to get a real image on the bat fauna of the
easternmost part of the Czech Republic. New data were obtained mainly by the results of systematic research in the Hostýnské vrchy Mts (Lučan 2000, Lučan & Svačina 2001), of acoustic
detectoring in Wallachia (Řehák 2001 and unpublished) or of an intense research of bat activity
at the Hranická propast Chasm, including a maternity colony roosting in a cave of the chasm
(Baroň & Řehák 2001, Řehák & Baroň 2002 and unpublished). Further data resulting from
long-term monitoring of hibernacula, Myotis myotis maternity colonies and bat-detectoring of
forest bats and sibling species Pipistrellus pipistrellus and P. pygmaeus (Řehák et al. 2005)
were included as well. Numerous data, even from regions omitted so far, were obtained thanks
to the improvement of species determination from bat detector samples, by tape recording the
ultrasound signals and their subsequent digitalization and computer analysis.
Although mountain and hilly elevations (up to 1323 m a. s. l.) prevail on the territory, large
underground spaces serving to bats as their natural hibernacula are relatively rare there, compared
to similar mountain regions elsewhere. Warm pseudo-karstic caves at higher elevations, situated
mainly in the eastern part of the territory, are exception. Some of them have been known for years
as bat hibernacula (Rumler 1985, Baroň & Řehák 1997, Wagner 2001). Neither the mining
was common in the Moravian-Silesian Carpathians, except for a few slate quarries and galleries (Kirchner & Řehák 1990, Řehák & Baroň 1997, Šafář & Rumler 2001). Winter records
of bats are particularly rare in the southern and western hilly grounds where only subterranean
vaults and cellars of houses are known as bat hibernacula. In contrast, summer records of bats
are relatively numerous. They concern checks of bats roosting in lofts, mainly of sacral buildings
and castles, netting of bats in the field and, during the last 15 years, acoustic bat-detectoring.
It was demonstrated that the occurrence of bats in summer was not limited to warmer lower
elevated sites with a higher food supply, but bats occurred also at higher elevations, mainly
in the second half of summer. Records of bats were made close to underground spaces where
some of them later hibernate, bat detectoring in the massif of the Lysá hora Mt. revealed bats
foraging on the mountain ridges at the elevation of 900 m and more.
There are two goals of the present paper: (1) to evaluate all records and to show the geographical distribution of bat species on maps with a standard girid, and (2) to analyse the altitudinal
distribution of individual species with respect to main periods of their life cyle.
Material and methods
Study area
As in the previous paper (Řehák 1998), the territory on the right bank of the Morava River was excluded
from the study area (Fig. 1). The area under study consists of 10 orographic units which are, approximately
in the north-south direction: Slezské Beskydy Mts, Jablunkovská brázda Furrow, Jablunkovská vrchovina
Highland, Moravskoslezské Beskydy Mts, Podbeskydská pahorkatina Upland, Rožnovská brázda Furrow,
Javorníky Mts, Hostýnsko-vsetínská hornatina Mts, Vizovická vrchovina Highland and Bílé Karpaty Mts
(Demek & Střída 1971). No research concerning bats was made in only one of them, the Jablunkovská
brázda Furrow, and there have been no records of bats there. The whole territory with its orographic and
hydrologic situation is shown by a map (Fig. 2). Mountains at the border between the Czech Republic
and Slovakia represent the highest part of the area, the elevation then decreases towards the west down to
the river valleys of Bečva and Morava. The elevation decreases also in the southward direction where it
is difficult to find the border between the Carpathians and the Dolnomoravský úval Valley.
202
Table 1. List of bat species recorded in the area of the Czech Carpathians and abbrevations for bat taxa
Tab. 1. Seznam druhů netopýrů zjištěných na území českých Karpat a zkratky jejich názvů
species
Rhinolophus hipposideros
Myotis mystacinus
Myotis brandtii
M. mystacinus seu M. brandtii
Myotis emarginatus
Myotis nattereri
Myotis bechsteinii
Myotis myotis
Myotis oxygnathus
Myotis daubentonii
Vespertilio murinus
Eptesicus nilssonii
abb.
Rhip
Mmys
Mbra
Mmys/bra
Mema
Mnat
Mbec
Mmyo
Moxy
Mdau
Vmur
Enil
species
Eptesicus serotinus
Nyctalus noctula
Nyctalus leisleri
Pipistrellus pipistrellus
Pipistrellus pygmaeus P. pipistrellus seu P. pygmaeus
Pipistrellus nathusii
Barbastella barbastellus
Plecotus auritus
Plecotus austriacus
P. auritus seu P. austriacus
abb.
Eser
Nnoc
Nleu
Ppip
Ppyg
Ppip/pyg
Pnat
Bbar
Paur
Paus
Paur/aus
Fig. 1. Map of Moravia including the W-Carpathian area under study.
Obr. 1. Mapa Moravy s vyznačením studovaného území Západních Karpat.
Explanations / vysvětlivky: A – Bohemian Highlands / Česká vysočina; B – Western Carpathians / Západní
Karpaty; a – Outer Carpathian Depressions / Vněkarpatské sníženiny; b – Outer Carpathians / Vnější Karpaty; c – Inner Carpathian Depressions / Vnitrokarpatské sníženiny; thick line / silná čára – state border /
stární hranice; middle line / středně silná čára – geomorphological system borders / hranice geomorfologických provincií; thin line / tenká čára – geomorphological subsystem border / hranice geomorfologických
podprovincií; chequered area / kostkovaná plocha – study area / studované území.
203
Material
A data base has been developed from all available sources since the twentieth of the 20th century until
the year 2006. It has 1447 items with records of 20 bat species. The species, including abbreviations of
their names, are listed in Table 1. Records with unspecified location or timing were excluded from the
evaluation of bats’ distribution. Mostly they concern old data (Remeš 1927, Gaisler 1956, Hanák 1960).
In addition to previous publication (Řehák 1998), the data base was completed by records published in the
papers by Lučan (2000), Lučan & Svačina (2001), Šafář & Rumler (2001), Wagner (2001), Jahelková
(2003) and, in certain species, missing data were assumed from Hanák & Anděra (2005). Unpublished
own data were added too.
Three categories of data are recognized. The first category is represented by old records before the year
1956 when the first thorough faunistic review of bats in the then Czechoslovakia was published (Gaisler
1956). The records made since 1956 were divided into winter and summer ones. The second category
(winter records) are those made in hibernacula and concerning torpid bats. In most cases, winter season
has formally been fixed to 15 October – 30 April (Gaisler et al. 1988, cf. Hanák & Anděra 2005). Irrespectively of this time span, findigs of torpid bats in May in hibernacula situated at high elevations were
considered winter records as well. The third category (summer records) comprises mostly the findings
made from 1 May to 14 October but, as in the case of winter records, this delimitation was not absolutely
observed. Finds and observations outside a shelter, e.g., made at the beginning of April or at the end of
October, were ranged to this category. Records of colonies were specified and evaluated separately from
the rest of summer records. Except in Vespertilio murinus, they concerned maternity colonies. All records
Fig. 2. Geographical map of the area under study.
Obr. 2. Geografická mapa studovaného území.
204
were assigned to the orographic units, quadrats of the mapping grid and their elevation was determined
(see Řehák 1998).
Methods
Various methods were used to get the data. Winter records were made either by occasional checks in
house cellars and vaste subterranean vaults of castles (Hanák 1960, Vlašín et al. 1993, 1995), or by
regular winter monitoring in former mine galleries (Baroň & Řehák 1997) and natural pseudo-karstic
clefts (ibid., Wagner 2001). All winter records of Vespertilio murinus came from above ground rooms of
buildings (Řehák 1998, Kašpar in litt.). Summer records were made by searching in lofts and under roofs
of buildings, namely churches and castles, by netting at the entrances to underground or on potential bat
hunting grounds, namely above brooks and on banks of water reservoirs. Important data have also been
obtained by the collection and analysis of owl pellets. Except in some cases, the detection of echolocation
signals yielded important data. Acoustic determination was often limited in cases where species identification based on external morphology was difficult as well. In sibling species such as M. mystacinus and
M. brandtii, or P. auritus and P. austriacus, visual identification from a distance is problematic (cf. Bar­
tonička 2004), in Pipistrellus bats it is nearly impossible vithout manipulation of them. In P. pipistrellus
and P. pygmaeus their identification based on morphology is problematic in any case (Řehák et al. 2005)
but they can be recognized by detecting their ultrasound signals. Acoustically based determination of the
two Pipistrellus species mentioned above, however, has been done only since 2000, some earlier records
therefore have to be assigned to Pipistrellus pipistrellus sensu lato. The relatively recently discovered
species P. pygmaeus (Jones & van Parijs 1993, Barratt et al. 1997) was acousticly detected only in three
places at lower elevations on the periphery of the territory studied (own unpublished data). Therefore,
records prior to 2000 made at higher elevations are assigned to P. pipistrellus sensu stricto.
Areal distribution of records belonging to the respective category was visualized with the aid of the
mapping grid (cf. Řehák et al. 2003). Relative representation of each species is then calculated as a percent
of quadrats where it was recorded out of the total of all quadrats of the area. Altitudinal distribution was
evaluated after the variance of elevations of the localities, divided again according to summer records,
winter records and records of maternity colonies. Non-parametric tests were applied, the Kruskal-Wallis H
test to the comparison among species and the Mann-Whitney U test to compare species pairs. All statistical
operations were made using the software Statistica for Windows.
Results
Bat fauna in particular orographical units
The number of localities with records of bats in particular orographic units, supplemented by the
number of recorded species, are given in Table 2. It is evident from the table that the Podbeskydská pahorkatina Upland and the Hostýnsko-vsetínská hornatina Mts have the most rich bat
fauna. Both orographic units are situated in the north-western part of the territory which gradually fades away into the lowlands of Outer Carpathians. There the number of both hibernacula
and localities with summer occurrence of bats is the highest which is reflected in the highest
number of bat species in winter as well as in summer. In contrast, Slezské Beskydy Mts and
Jablunkovská vrchovina Highland in the northern part of the territory have the lowest number of
both localities and bat species. In most orographic units, the number of localities with summer
records is higher than that with winter records (Table 2). All localities arranged according to
their situation in individual orographical units and quadrats are listed in Appendix 1.
205
Table 2. The total number of bat species and localities with records of bats in particular orographical
units
Tab. 2. Celkový počet druhů netopýrů a lokalit s nálezy netopýrů v jednotlivých orografických celcích
Explanations / vysvětlivky: NoL – number of localities / počet lokalit; NoS – number of bat species / počet
druhů netopýrů; W – winter season – records of torpid bats at hibernacula and all winter records of V.
murinus / zimní období – nálezy letargujících netopýrů na zimovištích a všechny zimní nálezy V. murinus;
G – growing season – other records of bats, especially from summer / mimohibernační období – ostatní
nálezy netopýrů, především letní
orographical unit / orografická jednotka
W
NoL
G
W
NoS
G
W+G
Slezské Beskydy Mts
Jablunkovská vrchovina Highland
Moravskoslezské Beskydy Mts
Podbeskydská pahorkatina Upland
Rožnovská brázda Furrow
Javorníky Mts
Hostýnsko-vsetínská hornatina Mts
Vizovická vrchovina Highland
Bílé Karpaty Mts
0
1
6
10
1
1
9
9
2
2
0
20
53
25
12
41
30
34
0
3
9
10
1
4
11
8
4
3
0
14
17
15
14
18
13
13
3
3
14
17
15
14
18
13
14
Areal distribution
The area studied covers altogether 60 quadrats of the mapping grid but 19 of them by less than
50% only. Some of the peripheral quadrats belong to the area by a very small territory. There
are no records of bats in 19 quadrats, 15 of them are the same as above. Bats were recently
found in all quadrats with old records prior to 1956. Winter records cover 19 quadrats (31.6%),
in one of them, 6478, only hibernating bats were recorded (Na Gírové cave in the Jablunkovská
vrchovina Highland). There are 6 quadrats (10%) with five and more hibernacula in each or
with an important hibernaculum where 10 or more individuals of one bat species were recorded
during a single check. The most important natural hibernacula include the pseudo-karstic cleft

Fig. 3. Distribution maps of individual bat species or pairs of sibling species.
Fig. 3. Mapy rozšíření jednotlivých druhů nebo dvojic podvojných druhů netopýrů.
Explanations / vysvětlivky: empty square / prázdný čtverec – records only before 1956 / jen nálezy před
rokem 1956; black square or semisquare / černý čtverec nebo půlčtverec – records of nursery colonies /
nálezy letních reprodukčních kolonií; grey square / šedý čtverec – records of male colonies (only V. murinus)
/ nálezy samčích kolonií (jen V. murinus); little empty circle or semicircle / malý prázdný kruh nebo půlkruh
– records of less than 10 individuals in 1–4 hibernacula / nálezy méně než 10 jedinců na 1–4 zimovištích;
big empty circle or semicircle / velký prázdný kruh nebo půlkruh – 5 or more hibernacula or records of
10 or more individuals in one hibernaculum during one check at least / 5 a více zimovišť nebo nálezy 10 a
více jedinců nejméně na 1 zimovišti během jedné kontroly; little full circle or semicircle / malý plný kruh
nebo půlkruh – less than 5 sites of summer records / méně než 5 lokalit s letními nálezy; big full circle or
semicircle / velký plný kruh – 5 or more sites of records of bats or records of 10 or more individuals (excl.
colonies) at a site per one survey during growing season / 5 a více lokalit s nálezy netopýrů nebo nálezy
10 a více jedinců (vyjma kolonií) na lokalitě při jedné akci během mimohibernačního období.
1 – Rhinolophus hipposideros, 2 – Myotis mystacinus, 3 – Myotis brandtii, 4 – Myotis mystacinus seu M.
brandtii, 5 – Myotis emarginatus, 6 – Myotis nattereri.
206
1
2
3
4
5
6
207
Table 3. Areal distribution of particular bat species or species groups
Tab. 3. Plošné rozšíření jednotlivých druhů netopýrů či jejich skupin
Explanations / vysvětlivky: NoQ – number of quadrates where bats were recorded (Myotis mystacinus
group includes M. mystacinus and M. brandtii, Pipistrellus pipistrellus group includes P. pispistrellus and
P. pygmaeus and Plecotus group includes P. auritus and P. austriacus) / počet kvadrátů s nálezy netopýrů
(Myotis mystacinus group zahrnuje M. mystacinus a M. brandtii, Pipistrellus pipistrellus group zahrnuje
P. pipistrellus a P. pygmaeus a Plecotus group zahrnuje P. auritus a P. austriacus)
species
Rhinolophus hipposideros
M. mystacinus group
Myotis emarginatus
Myotis nattereri
Myotis bechsteinii
Myotis myotis
Myotis oxygnathus
Myotis daubentonii
Vespertilio murinus
Eptesicus nilssonii
Eptesicus serotinus
Nyctalus noctula
Nyctalus leisleri
P. pipistrellus group
Pipistrellus nathusii
Barbastella barbastellus
Plecotus group
winter season
NoQ
%
15
8
6
3
4
10
0
7
2
2
3
0
0
0
0
3
14
25.0
13.3
10.0
5.0
6.7
16.7
0
11.7
3.3
3.3
5.0
0
0
0
0
5.0
23.3
growing season
NoQ
%
19
16
10
9
9
26
1
22
6
13
18
15
9
21
5
10
24
31.7
26.7
16.7
15.0
15.0
43.3
1.7
36.7
10.0
21.7
30.0
25.0
15.0
35.0
8.3
16.7
40.0
total
NoQ
%
20
18
13
9
9
28
1
24
6
13
19
15
9
21
5
10
30
33.3
30.0
21.7
15.0
15.0
46.7
1.7
40.0
10.0
21.7
31.7
25.0
15.0
35.0
8.3
16.7
50.0
caves in the Moravskoslezské Beskydy Mts (Cyrilka, Kněhyňská jeskyně and Ondrášovy díry
caves), Javorníky Mts (Pod kazatelnou I Cave) and Vizovická vrchovina Highland (three caves
in the hill Kopce near Lidečko). Important artificial hibernacula include the Sintrová Gallery
near Liptál (Vizovická vrchovina Highland), the subterranean vault of the Holešov Castle and
the cellars of buildings in Lukov. Except the Kněhyňská and Ondrášovy díry caves dominated
by Myotis myotis, all important hibernacula harbour Rhinolophus hipposideros as the most
abundant species in winter.
Summer records cover altogether 40 quadrats (66.7%); maternity colonies were found in 24
of them (40.0%). Summer male colonies of Vespertilio murinus were recorded behind window
shutters at two localities in the Hostýnsko-vsetínská hornatina Mts (quadrats 6572, 6672). In
addition to the quadrats with records of bat colonies, there are 6 quadrats with at least five localities in each or with an important summer locality where 10 or more individuals of one bat
species were recorded during a single check. Numerous bats were netted in late summer at the
entrances to caves serving also as hibernacula in three of the quadrats (6575, 6672, 6774). Bats

Fig. 3. Distribution maps of individual bat species or pairs of sibling species (continued).
Fig. 3. Mapy rozšíření jednotlivých druhů nebo dvojic podvojných druhů netopýrů (pokračování).
7 – Myotis bechsteinii, 8 – Myotis myotis, 9 – Myotis oxygnathus, 10 – Myotis daubentonii, 11 – Vespertilio
murinus, 12 – Eptesicus nilssonii.
208
7
8
9
10
11
12
209
were recorded at more localities in each of the remaining three quadrats. There the records were
made by either systematical bat detectoring (quadrats 6377, 6574) or by regular mist netting
(6472 – Hranická propast Chasm).
The distribution of records of individual bat species according to three categories of data
reflecting the time period, number of localities and number of bats is shown in Fig. 3 (1–23).
The data on areal distribution of all species are summarized in Table 3. In cases when sibling
species were not distinguished from each other or their determination was questionable, they
were lumped together in the table. With respect to the absolute and relative number of quadrats
with bat records, the species or groups of species with the largest distribution are Plecotus spp.,
Myotis myotis and M. daubentonii evidenced on at least 40% of the territory. In addition to them,
Pipistrellus pipistrellus s. l. and Rhinolophus hipposideros were recorded on one third or a little
more of the territory. In contrast, Pipistrellus pygmaeus (3 quadrats and localities) and P. nathusii
(5 quadrats and 6 localities) are very rare. Myotis oxygnathus, recorded only once and in one
locality, is actually the rarest known bat. In the hibernacula, R. hipposideros and Plecotus spp.
have the largest distribution covering ca 25% of the territory. The only other species covering
more than 15% of the territory is M. myotis. There are no records of hibernating individuals in
Nyctalus and Pipistrellus species. Summer records of M. myotis, Plecotus spp., M. daubentonii
and P. pipistrellus s. l. cover more than 1/3 of the territory. Other species common in summer
are R. hipposideros and Eptesicus serotinus. In E. nilssonii and Barbastella barbastellus the
summer records cover a significantly larger area than the winter ones.
Altitudinal distribution
Localities situated at the lowest elevations comprise bat hibernacula in house cellars in the
Podbeskydská pahorkatina Upland and Vizovická vrchovina Highland and a wine-vault in the
slopes of Bílé Karpaty Mts, all under 250 m a. s. l. Single P. austriacus, in one case P. auritus,
were recorded in a total of six cellars. In contrast, five pseudo-karstic caves in the massif of
highest peaks of the Moravskoslezské Beskydy Mts, i. e. Lysá hora, Kněhyně and Radhošť,
represent hibernacula situated at the highest elevations (945 – 1100 m), absolutely highest
situation having the Čertova díra Cave, 1100 m a. s. l. However, bats go up to such elevations
even in summer. In the Moravskoslezské Beskydy Mts, they were visually observed on the
Malá Stolová Mt. and acousticly detected on Lysá hora Mt., both at 900 m a. s. l. or higher and
in late summer bats were netted at the entrances of three pseudo-karstic caves (945, 1050 and
1050 m a. s. l).
The range of elevations and their median are shown in Fig. 4 (a–c). The variance of elevations is given for individual species as well as for the whole sample. The sample consists of 38
hibernacula and 14 bat species or species groups but M. bechsteinii, V. murinus, E. serotinus
and P. auritus seu austriacus were recorded in less than five hibernacula and were excluded
from the evaluation. The species M. mystacinus and M. brandtii were lumped together due to
the problems of their differentiation. Significant differences among the species were revealed
in the distribution of elevations of their hibernacula [Kruskal-Wallis test, H(9, N=112)=33.3,

Fig. 3. Distribution maps of individual bat species or pairs of sibling species (continued).
Fig. 3. Mapy rozšíření jednotlivých druhů nebo dvojic podvojných druhů netopýrů (pokračování).
13 – Eptesicus serotinus, 14 – Nyctalus noctula, 15 – Nyctalus leisleri, 16 – Pipistrellus pipistrellus, 17
– Pipistrellus pygmaeus, 18 – Pipistrellus pipistrellus seu P. pygmaeus.
210
13
14
15
16
17
18
211
Fig. 4a. Box-plots of altitudinal distribution of hibernacula. Abbreviations – see Table 1.
Obr. 4a. Krabicové diagramy výškové distribuce zimovišť. Zkratky – viz tab. 1.
p=0.0001]. From among all species, P. austriacus hibernates at the lowest situated localities
(median 259 m a. s. l.), being the only species evaluated in which the median of altitude lies
below 400 m. The distribution of elevations of hibernacula with P. austriacus differs significantly
from that in other 9 species (Mann-Whitney tests: U=3.5–27.0, Z=2.17–4.11, p<0.0301). On
the contrary, M. mystacinus seu M. brandtii and M. nattereri hibernate at the highest situated
localities (median 872 m a. s. l. and 1050 m a. s. l., respectively). The differences between the
elevations of hibernacula of M. nattereri and other species except P. austriacus are insignificant
(Mann-Whitney tests) due to the small sample size. The differences between the elevations of
hibernacula in the M. mystacinus group and P. austriacus as well as R. hipposideros and B.
barbastellus are significant (Mann-Whitney U tests: U=3.5, Z=3.814, p=0.000134, U=53.0,
Z=2.532, p=0.0113 and U=9.0, Z=1.967, p=0.049). The medians of elevations of hibernacula
range from 400 to 800 m a. s. l. in 7 species, being 400–600 m in R. hipposideros and B. bar­
bastellus (415 and 440 m, respectively) and 600–800 m in the remaining 5 species (Fig. 4a).

Fig. 3. Distribution maps of individual bat species or pairs of sibling species (continued).
Fig. 3. Mapy rozšíření jednotlivých druhů nebo dvojic podvojných druhů netopýrů (pokračování).
19 – Pipistrellus nathusii, 20 – Barbastella barbastellus, 21 – Plecotus auritus, 22 – Plecotus austriacus,
23 – Plecotus auritus seu P. austriacus.
212
19
20
22
21
23
213
Fig. 4b. Box-plots of altitudinal distribution sites with occurrence of bats during growing season. Abbreviations – see Table 1.
Obr. 4b. Krabicové diagramy výškové distribuce lokalit s výskytem netopýrů v mimohibernačním období.
Zkratky – viz tab. 1.
In 240 localities 22 species or species groups were recorded in summer. Three couples of sibling
species and P. pygmaeus (3 localities only) are excluded from the evaluation. In 11 out of the
18 evaluated species, the maximum median of elevations is 400 m, it exceeds this value in the
remaining species (Fig. 4b). Significant differences were found in the distribution of elevations
of summer localities among the species [Kruskal-Wallis test: H(17, N=435)=65.6, p<0.0001].
As in the hibernacula, the lowest median of summer elevations concerns P. austriacus (298
m). Again, the distribution of elevations of summer localities with P. austriacus differs significantly from that in most other species (Mann-Whitney tests: U=4.0–338.0, Z=2.264–5.159,
p<0.0236), the only exception being P. nathusii (p>0.05). The distribution of elevations of P.
nathusii summer localities (median 332 m) differs significantly only when compared with that
of M. brandtii (U=5.0, Z=2.286, p=0.0223), M.bechsteinii (U=14.5, Z=2.149, p=0.0317) and E.
nilssonii (U=37.0, Z=2.307, p=0.0210). The highest values of medians concern the elevations
of localities in M. brandtii (765 m) and M. nattereri ( 665 m). The difference between the data
of M. brandtii and that of most other species are significant (Mann-Whitney tests: U=3.5–50.0,
Z=2.185–3.780, p<0.0289), exceptions are that of M. nattereri and M. bechsteinii, both insignificant. The distribution of elevations of M. nattereri summer localities differs significantly
from that in 9 species (Mann-Whitney tests: U=22.5–195.5, Z=2.239–3.691, p<0.0252), while
the differences against the remaining 8 species are insignificant. Following are the results when
214
comparing two related species of the same genus: P. auritus prefers higher situated summer
localities than P. austriacus (U=125.0, Z=3.985, p=0.000068), M. brandtii than M. mystacinus
(U=30.0, Z=2.308, p=0.0210) and E. nilssonii than E. serotinus (U=363.5, Z=2.849, p=0.00438).
Only the differences between elevations of summer localities in N. leisleri and N. noctula are
insignificant.
No significant differences, probably due to a small sample size, were revealed when comparing
the elevations of 43 shelters of summer colonies among 9 bat species (Kruskal-Wallis test). B.
barbastellus was not evaluated due to the evidence of only one colony. Maternity colonies at
the highest elevations were found in E. nilssonii (n=3, median 500 m a. s. l.), maternity colonies
at the lowest elevations in P. austriacus (n=4, median 295.5 m a. s. l.).
Discussion
So far, 20 bat species were recorded on the studied territory belonging to the province of W Carpathians. At least 13 of them were found hibernating, this number being similar to that of species
recorded in hibernacula of near Oderské vrchy Mts where 11 bat species were recorded (Šafář
& Rumler 2001, Wagner 2001, 2004). No winter occurrence of Myotis brandtii was recorded
Fig. 4c. Box-plots of altitudinal distribution of sites with occurrence of nursery colonies. Abbreviations
– see Table 1.
Obr. 4c. Krabicové diagramy výškové distribuce lokalit s výskytem reprodukčních kolonií. Zkratky – viz
tab. 1.
215
so far but it is probable. When checking torpid bats in hibernacula visually without touching
them it is impossible to distinguish safely between M. brandtii and M. mystacinus and, therefore, M. brandtii could be overlooked between bats labelled as M. mystacinus (cf. Bartonička
2004). Most records of M. brandtii concern bats netted in late summer at entrances to caves
where the bats later hibernate, including M. mystacinus, which corroborates the hypothesis of M.
brandtii hibernation in the area studied. Due to this potential confusion, all winter records of M.
mystacinus were considered records of the M. mystacinus group. Another confusion concerning
M. brandtii possibly results from the fact that the localities of its netting records are situated
at high elevations, M. brandtii thus appearing to be a mountain species. Similar problem of
distinguishing in winter concerns the siblings P. auritus a P. austriacus and, accordingly, their
winter records were consider records of the P. auritus group. V. murinus, although recorded in
winter, was never found in an underground hibernaculum or even in a deep torpidity. Its recods
invariably concern single individuals in rooms of buildings above ground or observations of
flying bats. Compared to previous data (Řehák 1998) new hibernating species was found, viz.,
E. serotinus which was repeatedly recorded in castle vaults of Bystřice pod Hostýnem and
Holešov (Lučan & Svačina 2001). So far, no winter records of Nyctalus and Pipistrellus bats
in the area of study exist. Since bats of the genus Nyctalus do not hibernate underground, they
possibly either migrate to warmer regions of southern Europe or they hibernate in large tree
cavities. N. noctula, however, is known to hibernate in rock crevices and in various fissures
under the roofs and in the outer walls of buildings including the relatively newly built blocks
of prefabricated panel houses in towns and cities (Gaisler 1997, Lehotská & Lehotský 2000).
Such records are missing in the area so far. Low number of winter records of psychrophilous
hibernants, namely B. barbastellus and E. nilssonii, is probably due to the character of pseudo-karstic caves and galleries, all of them being relatively warm in winter with temperatures up
to 8 °C (Baroň & Řehák 1997), which are too high to ensure their winter torpor. On the other
hand, such hibernacula suit well to species which are dominant in the study area – M. myotis
and mainly R. hipposideros. Both species were found to be relatively abundant in all larger
hibernacula (Řehák 1998, Wagner 2001). The situation is very different from that in the near
Oderské vrchy Mts and Jeseníky Mts with vaste systems of abandoned galleries where winter
temperature is close to 0 °C at places and both B. barbastellus and E. nilssonii are very abundant
(Rumler 1985, Řehák & Gaisler 1999, Wagner 2001, 2004). At the same time, the hibernating
bat assemblages are much larger in that two mountain systems and include high numbers of
both eurytherm M. myotis and thermophile R. hipposideros (Buřič & Šefrová 2001, Řehák &
Gaisler 2001, Šafář & Rumler 2001). The pseudo-karstic cleft caves of the Moravskoslezské
Beskydy Mts are among the highest situated bat hibernacula in the Czech Republic. In four of
them, their elevation is above 1000 m. There is only one further hibernaculum in the Czech
Republic situated at such elevation, in the Jeseníky Mts (Souček 1969).
All 20 bat species evidenced in the area of study were recorded in the summer season.
Compared to previous data on bats in the Czech Carpathians (Řehák 1998), P. pygmaeus is
a new species which can be reliably determined in the field by the detection of its ultrasound
signals (Jones & van Parijs 1993). So far, this species was recorded in only three localities at
low elevations, invariably close to water streams or reservoirs (cf. Řehák et al. 2004, 2005).
Its sibling P. pipistrellus, in contrast, is abundant from lowlands to mountains over 900 m
a. s. l. (Řehák ibid.). As the probability of former recording P. pygmaeus at medium and higher
elevated localities is negligible, most older records are considered to represent the species P.
pipistrellus. Reproduction was evidenced in 10 bat species in which maternity colonies were
216
recorded. Although some of the colonies disappeared during the research the number of reproducing bat species is still high with respect to the relief of the area studied. Most maternity
colonies were recorded at low elevations, only three colonies of two species (E. nilssonii and
M. myotis) were discovered in the lofts of buildings at elevations higher than 500 m. In the
mounatins, maternity colonies were typically coupled to human settlements in river valleys.
The largest number of maternities (20) was found in Myotis myotis, at the same time being the
largest bat assemblages in the region (Řehák 1998). As elsewhere, large lofts of churches and
castles shelter maternity summer colonies of M. myotis. Large maternity colony in the nearly
inaccessible (to humans) Rotunda Cave belonging to the system of Hranická propast Chasm
is the only exception. It is the only maternity colony of M. myotis known from a cave in the
Czech Republic (Baroň & Řehák 2001, Řehák & Baroň 2002). This M. myotis summer colony
is unique in many respects, e.g. concerning its size: the maximum number of pregnant females
(young excluded) was estimated to 1,044 individuals, that of lactating females plus young to
more than 2,000 individuals (Řehák 2006, unpublished).
With respect to the landscape of the area studied, the total of 20 bat species is high and ranges
well to the lowland and middle elevated regions where bats have been studied for a long time.
There were only a few mountain systems in the Czech Republic the chiropteran fauna of which
was studied systematically, namely the Šumava Mts (17 species, Anděra & Červený 1994),
Žďárské vrchy Mts (15 species, Řehák et al. 2000) and Železné hory Mts (13 species, Benda et
al. 1997). Nineteen bat species were reported from the Krkonoše Mts (Anděra et al. 1974) but
the records of two of them (N. leisleri, R. ferrumequinum) are very old and doubtful.
Recently, faunistic data on bats were published from eastern Bohemia (Lemberk 2004) and
southwestern Moravia (Reiter et al. 2003). In the former region, 21 bat species and 14 species
with maternity colonies were recorded. In the latter region, 19 bat species were evidenced of
which in 17 the reproduction was proved. The only large region with a more rich chiropterofauna is represented by southern Moravian lowlands in which 22 or 21 species (if one excludes
the problematic N. lasiopterus) were recorded and in 11 species reproduction was evidenced
(Gaisler et al. 1990, 2002, Řehák et al. 2003). Concerning the lowlands of northern Moravia
which, similarly to southern Moravian lowlands, are situated close to the Carpathians, there are
faunistic data on bats from the Poodří and Litovelské Pomoraví. In the lowlands of Poodří, 17
species including P. pygmaeus (Řehák & Bryja 1998, unpubl. data) and in that of Litovelské
Pomoraví as many as 18 species were recorded (Bartonička et al. 2002). It can be concluded
that the regions are mutually similar concerning the numbers of their bat species.
In the orographic units, the representation of localities and bat species is uneven. There may
be double reasons of it, the differences in their geomorphology, mainly in the elevation, and
the differences in research activity concerning the bat faunistics. Important bat hibernacula in
the Moravskoslezské Beskydy Mts, Hostýnsko-vsetínská hornatina Mts and Vizovická vrchovina
Highland were systematically studied, while hibernacula suitable to such a study are missing in
the southern part including the Bílé Karpaty Mts. Summer records reflect mainly the search of
bats in large lofts of churches and castles which also are unevenly distributed, this fact again may
be responsible of the differences between the orographic units concerning their bat fauna.
The rank of species most common from the point of view of areal distribution is similar as
in other regions where the mapping grid was applied. The Czech Carpathians (60 quadrats in
total) can be compared with the Šumava Mts (50 quadrats, Anděra & Červený 1994), eastern
Bohemia (68 quadrats, Lemberk 2004) and southwestern Moravia (38 quadrats, Reiter et al.
2003). Generally, the quadrats of Carpathians are covered less by the records of bats than that
217
in other regions mentioned above what can be due to both worse living conditions for bats and
lesser activity of researchers in the Carpathians (Řehák 1998, 2001).
Plecotus species were recorded on 50% of the Carpathian territory but their abundance has
to be estimated with caution since the taxon comprises two species. Further bats common in
the Carpathians are M. myotis and M. daubentonii. The situation is similar in eastern Bohemia
and southwestern Moravia. In eastern Bohemia, P. auritus and M. myotis are the most distributed species while P. austriacus ranges fifth; in southwestern Moravia, P. austriacus ranges
first followed by M. myotis while P. auritus together with M. daubentonii occupy the third and
fourth places. The differences in abundance of P. austriacus and P. auritus in eastern Bohemia
and southwestern Moravia probably reflect different geomorphological situation and climatic
conditions between these two regions. While P. austriacus prefers warmer open lowland, P.
auritus is more common in colder woodland of highlands and mountains. As shown by this paper,
in the Carpathians the former species is more common in the warmer south at low elevation
and few woods, the latter is more common at higher elevated woodland of the northeast. M.
daubentonii followed by P. auritus represent the two most common species in the Šumava Mts
where P. austriacus, by the number of quadrats where it was recorded, ranges as far as sixth.
This can easily be explained by the lack of warm Pannonian lowlands in the region of Šumava
and its foothills. In southern Moravia, in contrast, the impact of warm lowland habitats on the
chiropteran fauna is evident (Reiter et al. 2003).
When evaluating the altitudinal distribution, uneven representation of most bat species is
striking. Often bats were recorded in lowland habitats and then relatively high in the mountains.
This in part reflects the situation of most important and regularly checked hibernacula at higher
elevations. Therefore the average elevation of winter bat records is higher than that of summer
records. Nevertheless, certain species were recorded at high elevations even in summer as in the
case of acoustic detection of bats foraging on the massif of the Lysá hora Mt., more than 900 m
a. s. l. (Řehák & Bartonička, unpubl.). Summer netting at the entrances to high situated caves
of the Beskydy Mts then raised the upper limit of summer occurrence in 10 bat species up to
1050 m a. s. l. which influenced the hypsometric distribution mainly of rare species difficult to
monitor by methods other than netting (M. brandtii, M. nattereri). In contrast, summer maternity
colonies were significantly more often found at lower elevations.
In any study of the distribution of bats over a large territory, the impact of the respective
scientist or scientists can not be excluded: the number of records is not only a function of bats’
occurrence but also of human activity. The number of records reflects the extent of monitoring
in both space and time. Acoustic bat-detectoring, in spite of certain limits (difficult or even
impossible determination of some species), substantially increases the capacity of field work
with bats enabling to monitor even less attractive habitats and localities neglected so far.
Souhrn
V práci jsou shrnuty dostupné údaje o výskytu netopýrů pocházejících z území Vnějších Karpat ve východní
části České republiky. Nálezy jsou rozděleny na zimní, letní a na nálezy letních kolonií a poté hodnoceny
podle jejich příslušnosti k orografickým jednotkám, kvadrátům síťové mapy a podle nadmořské výšky.
Celkem bylo v 9 orografických jednotkách a 41 kvadrátu, pokrývajících studované území, evidováno
20 druhů netopýrů a dvě dvojice podvojných druhů. Dosud bylo na zimovištích v českých Karpatech
zjištěno 13 druhů netopýrů a u 10 druhů byly v létě nalezeny reprodukční kolonie samic.
218
Plošnou distribuci jednotlivých druhů znázorňují síťové mapy. Za nejrozšířenější druhy lze považovat
netopýry rodu Plecotus (50,0 % kvadrátů pokrývajících studované území), Myotis myotis (46,7 %) a M.
daubentonii (40,0 %).
Výšková distribuce lokalit, kde byli nalezeni netopýři, je vyjádřena mediány a kvantily jejich nadmořských výšek a znázorněna pro jednotlivé druhy krabicovými diagramy podle období nálezů (zimní,
letní). Lokality s letními koloniemi jsou hodnoceny samostatně. P. austriacus preferoval průkazně nižší
polohy než ostatní hodnocené druhy, a to jak v zimě, tak v létě (medián 259, resp. 298 m n. m.). Naopak,
výše položená zimoviště využívali nejvíce netopýři skupiny M. mystacinus a M. nattereri (medián 872 m,
resp. 1050 m n. m.). Vysoké hodnoty jsou ale ovlivněny relativně nízkým počtem zimovišť vhodných pro
hibernaci těchto těžko nalezitelných druhů. Většinou se jedná o vysoko položené pseudokrasové jeskyně.
V mimohibernačním období byli ve vyšších polohách nalézáni M. brandtii a opět M. nattereri (medián
765, resp. 665 m n. m.). Výšková distribuce letních nálezů je u méně častých druhů do jisté míry ovlivněna odchytem před vchody do jeskyní, nalézajícími se ve velkých nadmořských výškách. Zvláště je to
patrné u druhů, které se jinými metodami v létě zjišťují obtížně. Rozdíly v nadmořských výškách lokalit
s letními koloniemi jsou vzhledem k nízkému počtu lokalit statisticky neprůkazné.
Acknowledgement
I am oblidged to Prof. Jiří Gaisler for all his comments on the text and its translation to English. I am
also grateful to Dr. Tomáš Bartonička for his help with statistical evaluation. The study was supported
by the Long-term Research Project of Ministry of Education, Youth and Sports of the Czech Republic
No. MSM0021622416.
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Appendix
The list of localities with records of bats / seznam lokalit s nálezy netopýrů
Explanations / vysvětlivky: SN – quadrate number of the Czech mapping grid / číslo kvadrátu české
mapovací sítě; SPN – number of bat species / počet druhů netopýrů; S – season / období; G – growing
season – “summer” records of bats / mimohibernační období – “letní” nálezy netopýrů; W – winter season – records of torpid bats at hibernacula and all winter records of V. murinus / zimní období – nálezy
letargujících netopýrů na zimovištích a všechny zimní nálezy V. murinus.
SN
site / lokalita
habitat / stanoviště
SPN
Slezské Beskydy Mts
6378 Nýdek
cottage
1
in the open air
3
Jablunkovská vrchovina Highland
6478 Girová, Na Girové cave
cave
3
Moravskoslezské Beskydy Mts
6377 Řeka
in the open air
2
Smilovice
above a stream
1
Komorní Lhotka
above a stream in a village centre 1
near a stream at the end of a village
3
6475 Frenštát p. R.
church loft
1
Malá Stolová, Leopoldka
in the open air
1
6476 Čeladná
cottage
1
Lukšinec, Ondrášovy díry cave cave
6
cave entrance
8
Lukšinec
mountain forest
1
Lysá hora, Malenovice canyon
mountain forest
7
Lysá hora, Pod Ivančenou
mountain forest and meadow
6
Ostravice
above a stream in a village
1
Ostravice, Mazák
above a stream
1
6477 Morávka
cottage
1
6575 Čertův mlýn, Čertova díra cave cave
7
Dolní Bečva, Kamenné
in the open air
1
cottage celllar
1
Kněhyně, Kněhyňská cave
cave
7
cave entrance
10
Kněhyně, Kyklop cave
cave
2
Pustevny, Cyrilka cave
cave
8
cave and cave entrance
9
Vasko
cave
2
6576 Staré Hamry
church loft
1
house roof
1
Podbeskydská pahorkatina Upland
6277 Ropice
above a stream
4
Třanovice
above a stream
1
6375 Hukvaldy
cottige roof
1
a game preserve
1
6376 Baška
dam
1
Dobrá
unknown
1
Frýdek-Místek
free on a pavement
1
S
G
G
W
G
G
G
G
G
G
G
W
G
G
G
G
G
G
G
W
G
G
W
G
W
W
G
W
G
G
G
G
G
G
G
G
G
223
6377 Hnojník
Střítež
6471 Helfštýn
Lipník nad Bečvou
6472 Černotín
Hranice
Hranice, Hůrka
Lipník nad Bečvou
Teplice nad Bečvou
6473 Hustopeče nad Bečvou
Hustopeče nad Bečvou, Na Valše
Lešná u Valašského Meziříčí
Perná-Bučí
Poruba
Starý Jičín
6474 Hodslavice
Nový Jičín
Štramberk
Štramberk, Bílá hora
Štramberk, Kotouč
Štramberk, Na Horečkách
Štramberk, Šipka
Štramberk, Trúba
Veřovice-Padolí
6475 Kunčice p. O.
Kunčice p. O., Skalka
6572 Kelč
6573 Branky na Moravě
Poličná
Poličná-Junákov
Poličná-Juřinka
Valašské Meziříčí
224
above a stream
above a stream, under a bridge
in the open air
above a river
gallery in a quarry
above a river
above a river
above a street-lamp near a autocamp
chasm
cellar
above a river
boiler room in a spa building
cave
cave
castle loft, pellets of T. alba
above a pond
castle cellar and corridor
castle loft
church loft
house roof anf loft
house
castle
church loft, pellets of T. alba
cellar
free in a castle park and a city
above a pond
above a swimming pool
in the open air in a park
garden in an abandoned quarry
in the open air in the Kamenárka quarry
gallery in a quarry
galleries
cave and cave entrance
near a cottage
above a stream
near a cottage
above a pond
house
cottage, garden
church and castle lofts, pellets of T. alba
castle loft
house loft
farm loft
stream bank - pellets of S. aluco
pellets of S. aluco
buildings, in the open air, pellets of T. alba
buildings and in the open air
castle cellars
castle loft
castle park
house loft
1
3
1
3
7
5
4
2
14
1
2
1
2
1
4
2
2
5
3
5
1
2
3
1
1
2
1
3
3
1
2
2
4
1
1
1
1
1
2
3
1
1
1
1
1
4
1
1
2
2
2
G
G
G
G
W
G
G
G
G
W
G
G
W
G
G
G
W
G
G
G
G
G
G
W
G
G
G
G
G
G
W
W
G
G
G
G
G
G
G
G
G
G
G
G
G
G
W
W
G
G
G
6671 Holešov
house loft
church loft
castle cellar
house cellars
Němčice
house cellar
Rožnovská brázda Furrow
6574 Hrachovec
above a river
Rožnov pod Radhoštěm
bulding
buildings
owl box, pellets of S. aluco
park and above a river
Rožnov p. R., Dolní Paseky
beech wood
Rožnov p. R., Hradisko
pellets of S. aluco
Střítež
in the open air
Velká Lhota
in the open air
Veselá
in the open air
Vidče
church loft
in the open air
Zašová
in the open air
church loft, owl pellets
6574 Zubří
church loft, owl pellets
above a stream
village and in the open air
Zubří-Hamry
tree hole, owl pellets
pond banks and in the open air
6575 Dolní Bečva
river bank
in the open air
Horní Bečva
church loft
in the open air
above a dam
Rožnov p. R., Rysová
owl box, pellets of S. aluco
Vigantice
in the open air
Javorníky Mts
6674 Hovězí
in the open air
Huslenky
church tower
above a river
6675 Velké Karlovice
village
6676 Malé Karlovice-Tísňavy
school loft, garden
Velké Karlovice-Podťaté
house
6774 Francova Lhota
cottage loft
Lužná
in the open air
Pulčín-Hradisko
rocks
Pulčín-Hradisko, Pod Kazatelnou cave
cave entrance
6874 Francova Lhota
in the open air
Zděchov
church loft
Hostýnsko-vsetínská hornatina Mts
6572 Bystřice p. Hostýnem
castle cellars
castle park
church loft
1
1
3
2
1
G
G
G
G
W
1
1
2
1
6
1
1
2
1
3
1
3
6
1
2
1
2
4
6
1
2
1
3
2
1
1
G
W
G
G
G
G
G
G
G
G
G
G
G
G
G
G
G
G
G
G
G
G
G
G
G
G
2
1
1
3
3
1
1
2
1
4
10
4
1
G
G
G
G
G
G
G
G
G
W
G
G
5
2
1
W
G
G
225
6573 Bystřička
school loft
1
Podhradní lhota
cadaver on the road
1
Polomsko
cottage
1
Rajnochovice
buildings
1
church loft
1
Rajnochovice, Juhyně valley
above a stream
2
6574 Bystřička-Zadrhlov
gallery
4
Malá Lhota, Páleniska
house loft
1
6671 Dobrotice-Lysina
cottage
1
6672 Chvalčov
road
1
village
3
Chvalčov, Bystřička valley
above a stream
8
clearing
1
Chvalčov, Smrdutá
cave
3
cave entrance
10
Rajnochovice, Košový
building
1
Rajnochovice, Uhliska
above a stream
1
Rusava
Protestant church, loft
1
Catholic church - loft
1
cadaver on the road
1
Rusava, Ráztoka
banks of a water reservoir
1
Rusava, Čecheřinka cave
cave
3
cave entrane
8
6673 Jablůnka
river bank
1
Kateřinice
school loft
1
Křížový, Zbojnická cave
cave
3
Pržno
church loft
1
Růžďka
loft
1
Vsetín
town, above a river
4
cellar
1
loft
1
6674 Halenkov
railway station, in the open air
1
Janová
in the open air
5
Vsetín-Hluboké
in the open air
1
6675 Karolinka
in the open air
1
Nový Hrozenkov
church loft and tower
2
Velké Karlovice, Jezerné
above a pond
7
6676 Velké Karlovice, Miloňov
loft of a hotel
1
6772 Kašava
church loft
1
Lukov
house cellars
1
Lukov, castle
castle cellars
3
6
Lukov - Hradisko c.
cave
3
cave entrance
2
6773 Liptál
Protestant church, loft
2
Catholic church, loft
1
school loft
1
Vizovická vrchovina Highland
6771 Zlín
in the open air
1
6772 Lešná u Zlína
cellar
1
Slušovice
church loft
1
226
G
G
G
G
G
G
W
G
G
G
G
G
W
G
G
G
G
G
G
G
W
G
G
G
W
G
G
G
W
G
G
G
G
G
G
G
G
G
W
W
G
W
G
G
G
G
W
G
6773 Bratřejov
church loft
3
Jasenná
church loft
1
Liptál, Malá gallery
mine
2
Liptál, Sintrová gallery
mine
5
Pozděchov
Protestant church, loft
3
Catholic church, loft and tower
4
Vizovice
castle cellar
2
castle loft
2
church loft
1
hospital loft
1
6774 Kopce u Lidečka, Naděje cave
cave, cave entrance
8
Kopce u Lidečka, Ďáblova díra c. cave
2
Kopce u Lidečka, Kolonie cave cave
5
Kopce u Lidečka
3 cave entrances
9
Leskovec
in the open air
1
Lidečko
church loft
2
in the open air
3
6872 Březnice
church loft
1
Doubravy
in the open air
1
Horní Lhota
church loft
1
Kaňovice
chapel turret
1
Luhačovice
castle cellar
2
castle loft
3
Pozlovice
church loft
1
Velký Ořechov
church loft
1
6873 Újezd
church loft
1
6874 Horní Lideč
in the open air
3
Lačnov
church loft
1
above a pond
1
6971 Uherský Brod
house cellar
1
church loft
1
7070 Blatnice
church loft
1
7071 Blatnička
church loft
1
above a pond and a stream
1
7072 Bánov
school loft
1
Nivnice
church tower
1
Bílé Karpaty Mts
6874 Nedašov
church loft
1
Poteč
barn
1
Valašské Klobouky
house loft
1
church loft
1
mill loft
1
town hall loft
1
in the open air
2
6972 Bojkovice
castle cellar
2
castle loft
1
Nezdenice
church loft
1
Rudice
house loft
1
Záhorovice
house roof
1
6973 Slavičín
castle loft
2
church loft
1
G
W
W
G
G
W
G
G
G
W
W
W
G
G
G
G
G
G
G
G
W
G
G
G
G
G
G
G
W
G
G
G
G
G
G
G
G
G
G
G
G
G
W
G
G
G
G
G
G
227
Štítná nad Vláří
Štítná nad Vláří, Popov
6974 Brumov
7070 Lipov
7071 Boršice u Blatnice
Horní Němčí
7072 Březová
Komňa
Strání
7073 Starý Hrozenkov, Rasová quarry
7170 Hrubá Vrbka
Kněždub
Kútky
Kuželov
Měsíční údolí
Radějov
Tvarožná Lhota, Horní mlýn
228
church loft
church loft
old mill
house loft
cottage
wine cellar
church loft and tower
below house roof
church tower
church tower
above a pond and around street lamps
above a pond
church loft and tower
church loft
cadaver on the road
chaple loft
above a pond
13 wooden hides
church, external wall, loft and tower
wood hide
church tower
above a swimming pool
1
1
1
2
1
1
1
1
1
1
4
1
2
1
1
2
2
2
2
1
1
4
G
G
G
G
G
W
G
G
G
G
G
G
G
G
G
G
G
G
G
G
G
G
Lynx (Praha), n. s., 37: 229–240 (2006).
ISSN 0024–7774
Dynamics of the Pleistocene bat fauna from the Matuzka Paleolithic site
(Northern Caucasus, Russia) (Chiroptera)
Dynamika netopýří fauny paleolitického stanoviště Matuzka (severní Kavkaz, Rusko)
(Chiroptera)
Valentina V. Rossina1, Gennadiy F. Baryshnikov2 & Bronislaw W. Woloszyn3
Paleontological Institute, Russian Academy of Sciences, Moskva, Russia; [email protected]
Zoological Institute, Russian Academy of Sciences, 199 034 Sankt Petersburg, Russia
3
Institute of Systematics and Evolution of Animals PAS, Sławkowska 17, PL–31-016 Kraków, Poland
1
2
received on 31 May 2006
Abstract. The upper Pleistocene sedimentary series in Matuzka cave (northern Caucasus) covering a period from MIS6 to MIS2 provided remains of 18 species of bats. The bat record is particularly rich in the
layers corresponding to Eemian and early Vistulian. It is characterized with appearance of thermophilous
elements Rhinolophus ferrumeqiunum and Miniopterus schreibersii and a broad spectrum of taxa including dendrophilous and demanding elements such as Plecotus auritus, Myotis brandtii, M. emarginatus,
M. nattereri and M. blythii. The lithophilous forms Eptesicus serotinus, Vespertilio murinus and Nyctalus
noctula appear continuously in all layers and represent a dominant component of the assemblage. In Eemian layers they are supplemented also with Hypsugo savii and Pipistrellus cf. kuhlii which absent from
the upper layers while Pipistrellus pipistrellus appear as late as in the Early Holocene.
Introduction
During the last decades, many Paleolithic sites have been discovered on the northwestern Caucasus. The rich bone material from these sites includes numerous records of bats.
The Caucasus is a unique region particularly in respect to its climatic specificities and biogeographical role. The Caucasian mammal fauna is characterized by considerable species richness
and high degree of endemism. This holds true also for bats: the northern Caucasus is presently
inhabited by one of the richest bat fauna of Russia. Unfortunately, until now almost no information was available on the history if the Caucasian bat fauna except for rather episodical and
isolated records (comp. Vereščagin 1959). To gain a better understanding of the development
of bat faunas, the study of new paleontological data on this group is required.
The present paper describes the material of Late Pleistocene bats collected by G. F. Bary­
shnikov during archeological excavations of the Matuzka Paleolithic site. This study provides
insight into the major stages of the formation of Caucasian bat communities during the essential
part of the last glacial cycle, Eeminan and Vistulian.
229
Material and Methods
The cave site Matuzka (42° 26’ N, 39° 45’ E) is located at 720 m above sea level, at the northern margin
of the Lagonakskoe Plateau on the right bank of the Matuzka River, a right tributary of the Pshekha River, 27 km south-southeast of Apsheronsk (Fig. 1). The cave cavity is in Upper Jurassic limestones, has
a dome-shaped roof and karstic niches in the walls and roof. The cave is up to 35 m wide and about 40 m
deep, the entrance faces southwest and is 20 m high.
According to geomorphologic analysis performed by Nesmeyanov (1999), the primary cave cavity, which
is most likely of karstic-erosive origin, was formed about 150–130 thousand years ago. The analysis of
limestone pack deformation has shown that the cave initially had two, or possibly, three layers.
The three-floor primary cavity of Matuzka cave, with the total height up to 30–35 m, suggests a long
time of the cavity formation, with the participation of lateral river erosion. This process apparently resulted
in the large final size of Matuzka cave (Nesmeyanov 1999). The cave is presently a large grotto.
The site was excavated from 1985 to 1988. The 6-m-deep sequence of Pleistocene-Holocene deposits
was exposed in the site. The section of excavation comprises 8 major lithologic beds (Table 1; Baryshni­
kov & Golovanova 1989).
The sequence is subdivided by lithologic features into four units. Three lower units (beds 7–3a) contain
a Pleistocene mammal fauna and Mousterian artifacts (Golovanova et al. 1995).
The first lower unit is 0.6 m thick (beds 8a–7b) includes the deluvial deposits composed of slightly
inclined laminated loam with insertion of gruss. Apparently, it was formed under conditions of low-activity washout.
The second and third units (beds 7a–3a) are composed of rubbly-clumpy material with varying content
of loamy-gruss-rubbly filler. The second unit (beds 7a–5) is 3 m thick, of land-rockslide genesis, composed of yellow loam. The third unit (beds 4d–3a) is formed by gray loam. Bed 4b is 0.1 m thick, contains
charcoals of fire spots (Nesmeyanov 1999), with the absolute date 34200±1410 BP (Golovanova 1996).
There are no absolute dates for the others layers. However, based on rodent fauna (Nadachowski &
Baryshnikov 1991) beds 7–6 are dated to terminal Middle Pleistocene, beds 5–3 are assigned to the time
of the last glaciation, and beds 2–1 are dated Upper Pleistocene-Holocene boundary (Zaitsev & Osipova
2004). The fourth unit (beds 2–1) is 0.6 m thick, composed of lumpy-rubbly loam and contains of ceramics
fragments and burnt bones of domestic animals (Nesmeyanov 1999).
Fig. 1. The region of investigation with a location of the Matuzka cave (asterisk).
Obr. 1. Oblast výzkumu s lokalisací jeskyně Matuzka (hvězdička).
230
In addition to the main sites, excavations were performed in further trench assigned as trench 3. Based
on the mammal fauna and bone fossilization pattern, the layers of trench 3 were dated to Pleistocene-Holocene boundary, i.e. they correspond roughly to beds 3 and 1–2.
A total of 217 skull fragments and isolated teeth of bats were examined (Table 1). The material was
examined by binocular microscope, identified and compared with the samples of Recent species. The
parataxonomic categories were applied in few cases in which considerable fragmentation and poor preservation of specimens did not allow exact identification: “small-sized Myotis” means of the same size as
M. brandtii, “medium-sized Myotis” means of the M. emarginatus size.
Results
Structure of the oryctecoenoses
The complete list of species found in individual samples from the Late Quaternary beds of Matuzka cave is in Table 1. In total, 17 species were recorded: Eptesicus serotinus (Schreber, 1774),
incl. cf. E. serotinus; Vespertilio murinus Linnaeus, 1758 (cf. V. murinus); Nyctalus noctula
(Schreber, 1774) (incl. cf. N. noctula); Nyctalus leisleri (Kuhl, 1817); Myotis blythii (Tomes,
1857) (M. cf. blythii); Barbastella barbastellus (Schreber, 1774); Plecotus auritus (Linnaeus,
1758); Rhinolophus ferrumequinum (Schreber, 1774); Miniopterus schreibersii (Kuhl, 1817);
Pipistrellus pipistrellus (Schreber, 1774); P. nathusii (Keyserling et Blasius, 1839); P. cf. kuhlii
(Kuhl, 1817); Hypsugo savii (Bonaparte, 1837); Myotis brandtii (Eversmann, 1845) (Myotis
cf. brandtii); M. nattereri (Kuhl, 1817) (M. cf. nattereri); M. bechsteinii (Kuhl, 1817) (M. cf.
bechsteinii); M. emarginatus (Geoffroy, 1806) (M. cf. emarginatus). The particular remains
were directly compared to the samples of Recent specimens and apparently fall in variation of
the respective Recent species except for several Pleistocene items which appear to be somewhat
larger than reported for the Recent Caucasian material.
The most abundant species of the oryctocenosis were E. serotinus and V. murinus (30% and
26% of all remains, respectively). N. noctula, Plecotus auritus (both about 7%), B. barbastellus
and R. ferrumequinum (about 5%) were recedent elements while Myotis blythii (with somewhat
more than 2%) and all remaining species (with less than 1.9%) represent the subrecedent elements of the sample.
As concerns the structural characteristics of the oryctcenoses the whole set clearly splits into
two markedly different units: (I) The faunal association of the bed 7 (accumulated during the
Mikulino Interglacial = Eemian in the Western Europe) that is characterized by considerable
species richness and includes thermophilous elements such as R. ferrumeqiunum, M. schrei­
bersii, P. kuhlii, H. savii. Its essential characteristics are partly retained in the overlaying bed
6, where Miniopterus or H. savii are absent, of course (Fig. 2). The distribution of these species
in the section marks the boundary between the Mikulino Interglacial and the onset of the Valdai
Glacial Period (= Würm). (II) Beds 5–3a (accumulated during the Valdai glaciation) yielded the
bat fauna that is slightly poorer both in species richness and abundance. It also differs from (I)
by presence of M. nattereri, M. emarginatus and M. bechsteinii and well-pronounced instability
in the proportions of particular taxa (Fig. 2). It is dominated with the mesophilous elements,
among other Plecotus auritus and B. barbastellus, and Nyctalus noctula which reaches peak
of its abundance in the middle Valdai horizons.
As demonstrated in Fig. 3, there is a clear correlation between the contribution of bats to the
total mammalian community of a layer and contributions of the rodents inhabiting either forest
or open-ground habitats: a positive correlation exists with the proportion of rodents inhabiting
mountain forest formations and negative correlation with the proportion of rodents living in
231
Table 1. Chronological position of Matuzka cave strata and numbers of Chiroptera records
Tab. 1. Chronologická posice vrstev jeskyně Matuzka a počty v nich nalezených netopýrů
time
MA
O-isotope mammal age
stage
Eastern Europe Caucasus
Matuzka cave
layer
Chiroptera (217)
0.025
2
Sungilian
Akhtyr
1–2 + shaft 3
P. pipistrellus (3),
P. nathusii (1)
E. serotinus (13)
V. murinus (4), P. auritus (1)
M. nattereri (1)
M. bechsteinii (2)
M. emarginatus (1)
3
Chasovali
3
E. serotinus (3), N. leisleri (1)
B. barbastellus (2)
M. brandtii (1)
medium-sized Myotis (2)
0.073
4
Shkurlatian
4
E. serotinus (19)
V. murinus (6), P. auritus (7)
B. barbastellus (8)
medium-sized Myotis (1)
small-sized Myotis (1)
M. nattereri (1)
M. bechsteinii (2)
M. emarginatus (2)
N. noctula (12)
0.116
5a–5d
5–5b
E. serotinus (2)
V. murinus (3), P. auritus (1)
M. nattereri (1)
M. emarginatus (1)
M. blythii (1), N. noctula (1)
0.128
5e
Binagady
6
P. cf. kuhlii (1), N. leisleri (1)
E. serotinus (13)
V. murinus (21), P. auritus (1)
M. nattereri (3), M. blythii (1)
M. emarginatus (1)
R. ferrumequinum (1)
0.195
6
Khazarian
Kvaisi
7
P. cf. kuhlii (1), H. savii (1)
E. serotinus (20)
V. murinus (23), P. auritus (4)
B. barbastellus (2)
M. brandtii (2), M. blythii (3)
R. ferrumequinum (10)
N. noctula (2)
M. schreibersii (2)
open landscapes, such as mountain-steppe and shrub-and-grassland habitats. This suggests
considerable retreat of bat populations in periods of the most pronounced glacial conditions
(indicated by percentage of open-ground elements).
232
Fig. 2. Percentual distribution of a total number of individuals of particular bats species in sequence of
layers of the Matuzka cave.
Obr. 2. Procentuální rozložení celkového počtu jedinců jednotlivých druhů netopýrů ve sledu vrstev
jeskyně Matuzka.
Fig. 3. Percentages of the bats in the total mammalian samples of particular layers (bold line) compared
to those of forest and open-ground rodents.
Obr. 3. Procentuální zastoupení netopýrů v celkovém vzorku savců (silná čára) ve srovnání se zastoupením
lesních druhů hlodavců a hlodavců otevřené krajiny.
233
Taphonomy
Holocene bones from bed 1 have natural white colour. Most of the Pleistocene bone specimens
are light brown, some are colored with manganese. Some bones (mostly teeth) were damaged
by the digestive juice (enzymes): bed 4b (N. noctula), trench 3 (bed 3, V. murinus), bed 6
(3 specimens of E. serotinus (2) and P. auritus (1)), bed 7 (10 specimens; 4 V. murinus, 5 E.
serotinus and 1 M. cf. brandtii). This suggests that a significant part of bat sample may represent
a taphocenosis. The predominating representation of the lithophilous elements in the samples
(Eptesicus serotinus, Vespertilio murinus, Nyctalus spp. Pipistrellus spp., Hypsugo savii) suggests at the same time that a considerable part of the material may originated from autochtonous
sources, supposedly winter colonies of the respective species roosting in the ceiling fissures in
the cave entrance and/or in the rocky walls surrounding the cave.
Discussion
Out of 23 bat species presently inhabiting the northern Caucasus, 17 have been recorded in
the fosil assemblages under study. The species: Rhinolophus hipposideros, R. euryale, Myotis
daubentonii, M. aurascens, M. mystacinus and Nyctalus lasiopterus are absent from the Matuzka
orictocenosis. It is very important to recognize the reasons for their absence for analysis of the
species diversity in the Pleistocene bat community.
At present, Rhinolophus hipposideros is a common bat species in Caucasia which range is
restricted to the forest zone (Gazaryan 2002). Though it frequently occurs in cave roosts it
does not form large colonies but prefers smaller caverns for roosting which would indicate its
possible incidence in Matuzka cave. In respect to the positive prediction on its appearance in
fossil record, its absence may suggest that the pattern of abundance and geographical range of
this species differed from those in the present time.
In the case of R. euryale, a strict cave-dweller, extremely rare in the area under study (only
one record by Bobrinskoj et al. 1965), its incidence in the site under study seems to be quite
improbable, similarly as in the case of the migratory bat Nyctalus lasiopterus, a tree-dweller
closely associated with forest vegetation. Also the remaining species of the Recent list which
absents in the site are not too probable to appear there.
The group of small-sized Myotis (M. brandtii, M. daubentonii, M. aurascens and M. mysta­
cinus) is scarce in the oryctocenosis, it is represented by ten specimens only. It is noteworthy
that all the listed species are considered abundant in the Recent of Caucasia (Gazaryan 2002,
2003). The total abundance of Myotis daubentonii, extent of its range and degree of synanthropy
substantially increased during past decades (Gazaryan 2003). The natural habitats of this species
are confined to the foothills and river valleys. This species seems to avoid the altitudes above
1000 m above sea level (Gazaryan 2003). The sibling species M. aurascens and M. mystaci­
nus were only recently distinguished as the separate species based on detailed morphological
analysis (Benda & Tsytsulina 2000). At present, M. aurascens inhabits lowlands and plains
of Western Caucasus; it has not been recorded higher than 400 m above sea level (Gazaryan
2002). M. mystacinus and M. brandtii are very similar ecologically, both inhabiting mountain
forests near water bodies. They coexist in the region under study (Gazaryan 2002). In case
of all these species, their habitat preferences and abundances do not suggest particularly high
probability of their incidence in Matuzka cave – therefore, their absence in the fossil sample
need not be of a serious indication value.
234
Fig. 4. Percentual contributions of particular bat species to a community structure of individual layers of
the Matuzka cave.
Obr. 4. Procentuální zastoupení jednotlivých druhů netopýrů ve sledu vrstev jeskyně Matuzka.
Single records of P. pipistrellus and P. nathusii in the early Holocene sediments of Matuzka
cave are of the particular significance. They suggest the spread of these species in that period
(especially in comparison to their absence in the earlier strata) which corresponds well to the
scenario presented for their spread in central and Western Europe by Horáček & Jahelková
(2005).
There are more factors which supposedly might considerably influenced structure of the record and actual dynamics of bat communities recorded in Matuzka cave. Here we will discuss
briefly three of them: (1) activity of the Paleolithic man; (2) aspects of bone accumulation; and
(3) the climatic and environmental dynamics of the period under study.
(1) The reconstruction of bat fauna history in the northwestern Altai region from the Pleistocene to the present time demonstrated that the historical development of cave-dwelling
bat community was considerebly affected by cave-dwelling effort of the Paleolithic humans
(Agadjanian & Rossina 2001, Rossina 2004). Occupation of caves by the humans apparently
reduced the population of the majority of bat species because of the smoke produced by human
fires. Thus, humans were a limitative factor for the cave-dwelling bat community since the
Pleistocene (Rossina 2005).
The Mousterian man did not produce tools in Matuzka cave, but brought there artifacts ready
for use (Nesmeyanov 1999). However, Paleolithic humans regularly visited Matuzka cave for
many ten thousand years, which is evident from the regular distribution of tools in beds 7–3
(Golovanova et al. 1995). The analysis of large mammal fauna leads to the same conclusion;
the Mousterian man used Matuzka cave as a short-term shelter for humans during hunting
(Baryshnikov & Golovanova 1989). Thus, in the case under consideration, human activity
exerted a little effect on the bat community.
(2) It is commonly accepted that the main source of fossil bats in cave deposits is dying
bats from seasonal cave-dwelling colonies (Ovodov 1974, Tiunov 1997, Filippov & Tiunov
235
1999). But it is doubtful in the case under consideration. During the past 120 thousand years,
Matuzka cave was a large grotto (Potapova 1992, Nesmeyanov 1999). Thus it seems largerly
improbable that it may serve a roost for large winter colonies of cave-dwelling bat species,
such as Myotis emarginatus or Miniopterus schreibersii, though occasional appearance of these
bats during vegetation period (which used the cave e.g. as a night roost) or in transient period
cannot be excluded, of course. On contrary, the regular appearance of the species inhabiting
rocky fissures is here quite a probable, and their considerable representation in the sample fits
to that possibility quite a well.
The considerable fragmentation of all specimens and signs of treatment by the digestive juice
indicate that pellets of birds of prey may were possibly a significant source of fossil material
from Matuzka cave. Such a possibility would be indirectly supported also by the conclusion
resulting of the study of fossil birds from Matuzka cave which demonstrated that a degree and
character of damage of birds bones as well as the species composition of the sample indicate
that fossils come from owl pellets (Potapova 1992). In these connection, the specificities of the
two most abundant bat species, E. serotinus and V. murinus are worth mentioning. At present,
they are synanthropic species which roost preferably in human buildings while the records from
their natural roosts are generally rare. Correspondingly, only few winter records of E. serotinus
and V. murinus are known from the western Caucasus (Kuzjakin 1950, Gazaryan 2002). V.
murinus similarly like Nyctalus noctula and N. leisleri are seasonal migrants. Particularly worth
of attention is that the large seasonal colonies of these bats may become a hunting objects of
owls and the frequency of these species in owl diet may thus locally grow quite a high (Schmidt
& Topal 1971, Ruprecht 1990, 2005, Obuch 1989, original data). It should be mentioned that
also in several teeth of P. auritus and M. cf. brandtii the enamel shows clear signs of corrosion
by the digestive juice. Note that Plecotus along with E. serotinus and N. noctula is the most
favorite object of owls hunting (Kowalski 1995).
Unfortunately, the material is too small to allow a detailed statistical analysis of possible taphonomic effects. In any case, it is clear that the fauna is to be considered a true oryctocenosis
contributed both by taphocenoses and thanatocenoses. Despite the, of course, the stratigraphic
setting of the site and the particular records prove that the records under study come from
a well defined stratigraphical context and bear the respective stratigraphical and paleoecological
information. The comparison of the absolute amount of bat records and percentages of habitat
specialists in rodents (Fig. 3) is particularly significant in these connections. An increase of bat
percentage in layers with increased frequency of woodland elements conforms well to a genral
expectations on the paleoclimatic significance of chiropteran record. The general dynamics of
the bat fauna in study thus apparently reflects changes in landscape and climatic conditions in
the source area. In this sense the analysis of the structure of fossil bat assemblages not only
provides additional information on environmental changes, but, in some cases, it promises to
regard some aspects not accessible from other evidence (thermal balance of rocky massif as
a key factor for hubernation of fissure-dwelling bats etc.). In addition, the analysis of faunal
structure of Pleistocene bats provides important stratigraphic results.
(3) The greatest proportion and taxonomic diversity of Chiroptera (about 30% of all specimens) is recorded in beds 7–5b. The absolute maximum of bats is in bed 7.1 (Figs. 2, 4). The
oryctocenosis contains forest species of Pipistrellus and Nyctalus and the forest-steppe Myotis
blythii (Gazaryan 2002). Apparently, during the accumulation of beds 7–6, a vast area was
occupied by forests. The small mammal fauna includes forest species of the genera Pitymys and
Dryomys. However, the proportions of mountain-steppe and shrub-and-grass rodent species are
236
also high (Baryshnikov & Golovanova 1989, Baryshnikov et al. 1995). The climate was warm;
this is evident from the highest diversity of the bat assemblage from these beds (14 bat species
out of 17 are recorded) and the presence of migratory species of Pipistrellus and thermophilous
species, such as Rhinolophus ferrumequinum and Miniopterus schreibersii (more then 40% of
all bat specimens; Fig. 4).
At the boundary between beds 6 and 5b, the bat populations sharply decreased in number, with
the absolute minimum in bed 5a (Figs. 2, 3). Apparently, at that time, the area of open landscapes
increased and a more mosaic landscape structure was formed. This is supported by an increase
in the number and proportion of mountain-steppe rodents (Fig. 3), such as Spermophillus cf.
musicus, Spalax microphtalmus and Cricetulus migratorius guamensis. The climate became
somewhat cooler (Baryshnikov & Golovanova 1989).
During the accumulation of beds 5–4d–4c, the abundance and proportion of bats gradually
increased (Figs. 2, 3). The proportion of forest rodent species increased, while rodents of open
landscapes decreased in number (Fig. 3). This suggests an increase in the area of forest landscapes and, as follows from palynological data, the mountain pine forests spread, although they
were more xerophilous than at the present time (Baryshnikov et al. 1995).
Bed 4b shows a decrease in the proportion and abundance of bat remains, while the proportions of forest and steppe rodent species are approximately equal (Fig. 3). According to
palynological data, the forest-steppe or steppe with mixed broad-leaved and pine forests were
widespread at that time. However, grasslands dominated. The climate was relatively arid and
temperate (Baryshnikov & Golovanova 1989, Baryshnikov et al. 1995). Against the background
of a general increase in the proportion of rodents in bed 4a, the diversity and abundance of
bats also increased (more then 16% of all specimens) (Figs. 2, 3). Apparently, the landscapes
showed a mosaic pattern of distribution, so that mountain forests alternated with steppes and
grasslands. The pollen spectra suggest the presence of mixed pine-birch forests with alder-tree
and grass meadows (Baryshnikov et al. 1995).
Bed 3c yielded only Barbastella barbastellus and medium-sized Myotis (Fig. 2). In bed 3b, the
proportion of bat remains is equal to that of forest rodents (Fig. 3). At that time, the proportion
of Chionomys increased and steppe rodent taxa predominated (Baryshnikov & Golovanova
1989). Palynological data suggest that the subalpine grasslands and mesophilic motley grasses
dominated (Baryshnikov et al. 1995). The finds of bats and forest rodents are sporadic in bed
3a but steppe rodents predominate here as well (more than 90%; Fig. 3). Apparently, open
landscapes dominated at that time. Floral samples contain underdeveloped pollen of trees and
traces of turf-uncovered slopes, and Woodsia alpina. This suggests that at that time the cave
was in the Alpine belt, close to the upper boundary of forests (Baryshnikov et al. 1995).
As demonstrated above, the bat assemblage from Matuzka cave is divisible into two faunal
associations, which describe different climatic periods of the terminal Pleistocene.
Particular elements of the bat fauna can be used as biostratigraphic markers of time boundaries in Late Quaternary sediments of the Caucasus.
Conclusions
(1) Out of 23 bat species presently inhabiting the western Caucasus, 17 have been found in
the fossil record. Absence of some species in Matuzka oryctocenosis (Rhinolophus hipposide­
ros, R. euryale, Myotis daubentonii, M. aurascens, M. mystacinus and Nyctalus lasiopterus)
is most likely caused by taphonomic factors. Thus, by the end of the Middle Pleistocene, the
237
general appearance of the bat fauna had already been formed and remained almost constant to
the present time.
(2) Considerable part of the fauna is formed by lithophilous forms Eptesicus serotinus, Vespertilio
murinus, Nyctalus noctula, supposedly inhabiting rocky fissures in the site. At least part of the
sample apparently represents a taphocenosis coming from bird pellets.
(3) The general dynamics of the number and structure of Pleistocene bat communities from
Matuzka cave are in accordance with those of rodents inhabiting different landscapes and, hence,
indirectly reflect environmental changes in the area of Matuzka cave. The bat fauna apparently
decreased in time of spread of open-ground habitats.
(4) In the Eemian Interglacial, the fauna of bats was the richest and included thermophilic
R. ferrumeqiunum and Miniopterus schreibersii, besides of the records of Hypsugo savii and
Pipistrellus kuhlii, which demonstrate in the Eemian the ranges corresponding to their Recent
distribution.
(5) The Valdai glaciation (the time of beds 6–3a accumulation) is characterized by a slightly
poorer and less numerous bat fauna, which includes Myotis nattereri, M. emarginatus and
M. bechsteinii, and is distinguished by well-pronounced fluctuations of the proportions of taxa.
The proportion of psychrophilic faunal elements, such as Plecotus airutus and B. barbastellus,
noticeably increased.
SOUHRN
Svrchnopleistocenní sled vrstev sedimentů jeskyně Matuzka, která se nachází na severním Kavkaze (Rusko), postihuje údobí od MIS6 po MIS2. V těchto vrstvách byly nalezeny zbytky nejméně 217 jedinců 18
druhů netopýrů. Faunový nález netopýrů je nebývale bohatý ve vrstvách odpovídajících Eemu a spodnímu
Vistulianu. Ten je typický přítomností thermofilních prvků (Rhinolophus ferrumeqiunum a Miniopterus
schreibersii) a širokým spektrem taxonů včetně dendrofilních a vzácných prvků (Plecotus auritus, Myotis
brandtii, M. emarginatus, M. nattereri a M. blythii). Lithofilní formy (Eptesicus serotinus, Vespertilio
murinus a Nyctalus noctula) se objevují nepřetržitě ve všech vrstvách a představují dominující složku
společenstva. V eemských vrstvách jsou ale navíc druhy Hypsugo savii a Pipistrellus cf. kuhlii, které chybějí
ve svrchních vrstvách, zatímco Pipistrellus pipistrellus se objevuje nejdříve až ve spodním holocenu.
Acknowledgments
We are heartily grateful to Professor A. K. Agadjanian (PIN RAS) for an all-round support in our research.
We would also like to thank G. S. Rautian (PIN RAS) for valuable comments on the manuscript. This
study was supported by the Presidium of the Russian Academy of Sciences (Program no. 25, “Biosphere
Origin and Evolution”) and “Historical Dynamics of Bioresources As a Prerequisite for Their Conservation
and Harmonious Exploitation,” the Russian Foundation for Basic Research (project nos. 04-05-64805
and 05-04-48493), and the Russian State Program for Support of Leading Scientific Schools (project no.
NSh-6228.2006.4).
238
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Lynx (Praha), n. s., 37: 241–246 (2006).
ISSN 0024–7774
Plecotus macrobullaris – new bat species for Albanian fauna
(Chiroptera: Vespertilionidae)
Plecotus macrobullaris – nowy gatunek w faunie nietoperzy Albanii
(Chiroptera: Ve­sper­ti­li­o­ni­dae)
Konrad SACHANOWICZ1 & Mateusz CIECHANOWSKI2
Department of Animal Ecology, Nicolaus Copernicus University, ul. Gagarina 9, PL–87-100 Toruń,
Poland; [email protected]
2
Department of Vertebrate Ecology and Zoology, University of Gdańsk, al. Legionów 9,
PL–80-441 Gdańsk, Poland; [email protected]
1
received on 27 July 2006
Abstract. Plecotus macrobullaris was recorded in the mountains of northern Albania (Pukë district) on
9–10 August 2003. This represents the new species of bat in the poorly known mammal fauna of Albania.
At the reported locality, P. macrobullaris occurred in syntopy with P. auritus.
Introduction
The bat fauna of Albania includes ca. 24 species and belongs to the least studied among Eu­
ro­pean countries, with ca. 60% of species reported from single localities (Uhrin et al. 1996).
Moreover, the presence of several species in the country is doubtful after recent changes in
taxonomy of Myotis mystacinus and Pipistrellus pipistrellus complexes, as well as the genus
Plecotus (Benda & Tsytsulina 2000, Jones & Barratt 1999, Kiefer & Veith 2001, Spit­zen­
ber­ger et al. 2001, 2003). Actually, four species of Plecotus are recognized in the con­ti­nen­tal
Europe and all of them can be identified on the basis of external characters (Kiefer & von
Helversen 2004, Tvrtković et al. 2005, Spitzenberger et al. 2006). Their geographical ranges
overlap largely over the Balkan Peninsula.
Previous records of Plecotus auritus (Linnaeus, 1758) and P. austriacus (Fischer, 1829) pu­b­
lis­hed for Albania (Hanák et al. 1961, Hanák 1964, Lamani 1970) have not been do­cu­men­ted
sufficiently to reject the bat misidentification for their newly defined sibling species P. macro­bul­
la­ris Kuzyakin, 1965 and P. kolombatovici Ðulić, 1980. The presence of P. au­ri­tus – a female
collected in 1914 in Vermosha (Malësi e Madhe district) – has been confirmed recently in the
result of molecular investigation (Spitzenberger et al. 2001). The occurrence of P. austriacus
in Albania, although very probable, demands confirmation.
Herein, we report the first record of P. macrobullaris from Albania and comment its iden­ti­fi­
ca­ti­on in the field. 241
Records
On 9–10 August 2003, 7 individuals of long-eared bats (4 females, 3 males) identified as P. macrobullaris
were mist-netted in the entrance of a small adit (42° 06’ N, 20° 07’ E), 787 m a.s.l., east of Qafëmal village, by the road Shkodra – Kukës (N Albania, Pukë district). The adit, cut in serpentinite rocks, has been
used as a local water intake, containing a spring and a small pond in the entrance, which is located on the
rocky mountain slope, sparsely overgrown with low black pines Pinus nigra and bushes (Fig. 1). There
have been a small settlement and cultivations in the area nearby, although the dominating habitat in the
sur­roun­ding was heavily overlogged mountain forest of the black pine and the beech Fagus sylvatica. Two
adult females of P. auritus were caught at the same site, what allowed comparison of characters of both
species held in a hand. After being identified, sexed, measured and photographed all bats were released.
For the species identification in the field we have used characters given by Spitzenberger et al. (2002),
Mucedda et al. (2002) and later confirmed by Tvrtković et al. (2005).
Fig. 1. Entrance of the small adit near the Qafëmal village (K. Sachanowicz).
Rys. 1. Wejście do sztolni w pobliżu miejscowości Qafëmal (K. Sachanowicz).
242
Table 1. External measurements and weight of P. macrobullaris individuals mist-netted in the entrance
of the adit near Qafëmal (N Albania) on 9–10 August 2003 (F – female, M – male, ad. – adult, juv. – juvenile, lact. – lactating)
Tab. 1. Wymiary zewnętrzne oraz ciężar osobników P. macrobullaris odłowionych przy otworze sztolni
w pobliżu Qafëmal (północna Albania) 9–10 sierpnia 2003 (F – samica, M – samiec, ad. – dorosły, juv.
– młody, lact. – karmiąca)
No
sex
age
1
2
3
4
5
6
7
F
F
F
F
M
M
M
ad., lact.
juv.
juv.
ad.
juv.
juv.
ad.
mean±SD
forearm length
[mm]
weight [g]
thumb length
[mm]
claw length
[mm]
41.6
41.0
41.4
41.0
38.5
40.8
39.4
9.0 7.0
7.0
7.5
7.0
7.5
7.0
7.2
7.0
7.1
–
7.0
7.0
7.0
2.6
2.8
1.9
–
2.9
2.7
2.9
40.53±1.14
7.43±0.73
7.05±0.08
2.63±0.38
Morphologically, individuals of P. macrobullaris resembled rather P. auritus than P. aus­tri­a­cus, but by
the overall, contrasting coloration they were more similar to the latter species. Fur on dorsal and ventral
sides was very dense, woolly and unusually long (ca. 10 mm), in which character the species differed
markedly from P. auritus. The base of hairs on both sides was blackish. Dorsal fur was dark greyish-brown, strongly contrasting with almost pure white (whi­tish in some adults) on a ventral side. Juveniles
had more greyish dorsal fur and brighter ventral side. The head and muzzle were greyish-brown, slightly
darker than dorsal pelage. Pro­tu­be­ran­ces above eyes were of the size (ca. 1.0–1.5 mm in diameter) similar
to their equivalents in P. auritus. The triangular pad on a lower lip was obvious in all individuals of P.
macro­bul­la­ris. It was pale, lip coloured, in adults and darker in juveniles (Fig. 2). The base of the tragus
was pale pinkish, while its upper half was dark greyish. The tragus width in two adult females was 5.3
and 5.9 mm. The size of a hind foot was almost the same as in P. auritus; feet were also covered with
well visible hairs. Penis was broad (ca. 2.2 mm), cylindrical and parallel sided at almost whole length
with a pointed tip. A thumb of P. macrobullaris was slightly longer than in two P. auritus (6.3 and 6.0
mm), while a claw was of similar size (2.6 and 2.5 mm, cf. Table 1). External me­a­su­re­ments and weight
of P. macrobullaris are given in Table 1.
comments
P. macrobullaris occurs in mountain regions from central Pyrenees to Bosnia and Her­ze­go­vi­na
and from central Greece to the Caucasus, Syria and Iran, with a single records from Corsica
and Crete (Garin et al. 2003, Juste et al. 2004, Kiefer & von Helversen 2004, Spit­zen­ber­ger et
al. 2006). Astonishingly, in the Balkans the species is known only from the western part of the
Peninsula (Slovenia, Croatia, Bosnia and Herzegovina, Greece), while no localities were found
in its eastern regions e.g. in Bulgaria (Benda & Ivanova 2003). The Albanian record confirms
continuous distribution of P. macrobullaris in the Dinarian Mts, as well as its syntopic occur­
ren­ce with P. auritus. However, the species apparently is not re­s­t­ric­ted to higher (above 800 m)
altitudes, at least in the Dinarian Mts, as it has been suggested (Kiefer & Veith 2002, Kiefer
& von Helversen 2004). In Croatia, it was recorded within altitudinal range from the sea level
up to 1800 m a.s.l., with most localities – like the only Albanian – located below 800 m a. s. 1.
(Pavlinić & Tvrtković 2004). In contrast to other members of the genus, P. macrobullaris in
243
Table 2. Comparison of the forearm length of P. macrobullaris from different parts of its range. Data
source: Caucasus and Turkey – Spitzenberger et al. (2003); Greece, Liechtenstein and French Alps (ho­lo­
ty­pe) – Kiefer & Veith (2001), P. Benda, unpubl.; Austria – Spitzenberger et al. (2002); Pyrenees – Garin
et al. (2003), P. Benda, unpubl.; Iran, Syria and Switzerland – P. Benda, unpubl.; Croatia – Tvrtković et
al. (2005); Albania – this paper
Tab. 2. Porównanie długości przedramienia osobników P. macrobullaris z różnych części zasięgu. Źródła
danych: Kaukaz i Turcja – Spitzenberger et al. (2003); Grecja, Liechtenstein i Alpy Francuskie (holotyp)
– Kiefer & Veith (2001), P. Benda, unpubl.; Austria – Spitzenberger et al. (2002); Pireneje – Garin et
al. (2003), P. Benda, unpubl.; Iran, Syria i Szwajcaria – P. Benda, unpubl.; Chor­wa­c­ja – Tvrtković et al.
(2005); Albania – niniejszy artykuł
region
mean
Cau­ca­sus
Iran
Syria
Turkey
Greece
Albania
Croatia
Austria
Liechtenstein
Switzerland
French Alps
Pyrenees
42.77
41.73
41.30
–
39.85
39.57
39.95
40.59
–
41.55
–
42.00
males
SD
min
0.48
1.63
0.96
–
0.21
1.16
1.57
0.59
–
1.20
–
1.13
42.1
39.6
40.2
–
39.7
38.5
37.3
39.6
–
40.7
–
41.3
max
n
mean
43.2 6
43.2 6
42.0 3
40.9 1
40.0 2
40.8 3
42.5 25
41.5 7
–
–
42.4 2
40.5 1
43.3 3
43.10
42.80
42.68
42.67
–
41.25
41.22
41.91
–
40.67
–
42.55
females
SD
min
1.45
1.32
1.44
1.23
–
0.30
1.26
0.86
–
0.60
–
0.66
40.7
41.8
39.8
40.5
–
41.0
39.0
40.5
–
40.1
–
42.1
max
n
44.2 5
45.1 5
44.6 11
44.2 9
–
–
41.6 4
43.5 30
43.5 11
39.7 1
41.7 6
39.6 1
43.5 4
Croatia is restricted to karstic areas, where its coexistence with P. auritus was observed at three
sites (Tvrtković et al. 2005).
The forearm length of Albanian P. macrobullaris includes within the variation range of spe­
ci­mens from the Alps and Croatia (Kiefer & Veith 2001, Spitzenberger et al. 2002, Tvrt­ko­vić
et al. 2005). Individuals from the Pyrenees, Caucasus and the Middle East seem to have longer
fo­re­arms (Spitzenberger et al. 2006, Table 2). From the other hand, bats from Albania have sig­ni­
fi­cant­ly longer thumb than the Pyrenean individuals of P. macrobullaris (Mann-Whitney U-test,
U=0, p<0.005, n=12; data from Garin et al. 2003 and Table 1). Our data, as well as those from
Spain (Garin et al. 2003) seem to contradict that thumb and claw of P. macrobullaris are shorter
than in P. auritus (Spitzenberger et al. 2006). The coloration of the tragus, facial mask as well
as contrasting greyish-brown and white pelage of Albanian bats correspond with characters of
specimens from the Alps and Croatia (Kiefer & Veith 2001, Tvrtković et al. 2005) but differ
from the Pyrenean and other populations (Garin et al. 2003, Spitzenberger et al. 2006). Finally,
me­a­su­re­ments and external characters of Albanian bats may suggest their affiliation to smaller
sub­spe­ci­es P. m. alpinus distributed from the Alps to Bosnia and Herzegovina (Spitzenberger et
al. 2003, 2006, Kiefer & von Helversen 2004). The Pyrenean population shows measurements
and characters more similar to larger P. m. macrobullaris ranging from the north-eastern Italy to
the Caucasus, Syria and Iran (Kiefer & von Helversen 2004, Spitzenberger et al. 2006, Table
2), but the results of genetic analysis have placed that population within the western subclade
(Spit­zen­ber­ger et al. 2006). The­re­fo­re, the zone at least from the eastern Alps to the southern
Dinarian Mts should be inhabited by sympatric western and eastern populations. Moreover, this
244
Fig. 2. Plecotus macrobullaris from the area of Qafëmal, northern Albania (K. Sachanowicz).
Rys. 2. Plecotus macrobullaris z okolic Qafëmal w północnej Albanii (K. Sachanowicz).
may contradict the statement that in P. macrobullaris the forearm length decreases towards the
west (Spitzenberger et al. 2003), suggesting rather different pattern of geographical variation
– with smaller individuals in cen­t­ral part of the range and larger in its marginal parts. However, larger samples for mor­pho­met­ric and molecular studies are required to solve the problem
of external variation and dif­f­e­ren­ces in skull measurements (Spitzenberger et al. 2003, 2006)
among po­pu­la­ti­ons of P. macro­bul­la­ris from different parts of its range.
Streszczenie
Nowy gatunek dla fauny nietoperzy Albanii – Plecotus macrobullaris – został stwierdzony w górach
pó­ł­noc­nej części kraju (region Pukë) 9–10 sierpnia 2003. Na opisanym stanowisku gacek alpejski wspó­łwy­stępo­wał z gackiem brunatnym Plecotus auritus.
Acknowledgements
We thank Petr Benda for unpublished morphological data, comments on the manuscript and help in
obtaining two rare publications.
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Lynx (Praha), n. s., 37: 247–254 (2006).
ISSN 0024–7774
Supplementary notes on the distribution of Pipistrellus pipistrellus complex
in the Balkans: first records of P. pygmaeus in Albania and in Bosnia and
Herzegovina (Chiroptera: Vespertilionidae)
Uzupełnienia do rozmieszczenia nietoperzy z kompleksu Pipistrellus pipistrellus na
Bałkanach: pierwsze stwierdzenia P. pygmaeus w Albanii oraz w Bośni i Her­ce­gowi­nie
(Chiroptera: Vespertilionidae)
Konrad Sachanowicz1, Mateusz Ciechanowski2 & Alek Rachwald3
1
Department of Animal Ecology, Nicolaus Copernicus University, ul. Gagarina 9, PL–87-100 Toruń,
Poland; [email protected]
2
Department of Vertebrate Ecology and Zoology, University of Gdańsk, al. Legionów 9,
PL–80-441 Gdańsk, Poland; [email protected]
3
Department of Forest Ecology and Wildlife Management, Forest Research Institute,
ul. Bitwy Wars­za­wski­ej 3, PL–00-973 Warszawa, Poland; [email protected]
received on 28 April 2006
Abstract. Two males Pipistrellus pygmaeus were captured in August 2003 near Qafëmal (northern Al­
ba­nia) and identified on the basis of morphological characters. Hunting individuals of this species were
ob­ser­ved and detected also in a riparian forest of the Vjosës valley near Tepelenë (southern Albania) in
April 2004. They represent the first records of this species from Albania. P. pygmaeus is reported for the
first time also from Bosnia and Herzegovina, where echolocation calls of the species (lowest frequencies
52–57 kHz) were recorded near Miljevina and in Dobro Polje.
Introduction
Since the time of recognition of the pygmy (soprano) pipistrelle Pipistrellus pygmaeus (Le­
ach, 1825) within the P. pipistrellus complex (Barratt et al. 1997, Jones & Barratt 1999),
its dis­tri­bu­ti­on has been intensively surveyed across Europe. Soon, it was discovered that the
species ranges widely from Portugal to western part of Russian Federation and the Caucasus,
and from Great Britain, southern Scandinavia and Estonia to Greece and Turkey (Horáček et
al. 2000, Limpens 2000, Mayer & von Helversen 2001, Benda et al. 2003a, Hulva et al. 2004,
Wermundsen & Siivonen 2004, Vierhaus & Krapp 2004).
Although widespread, the range of this species in the Balkans is known only fragmentary due
to the low intensity of bat surveys there in general. Actually, its presence has been con­fir­med
for most of the Balkan countries (Limpens 2000, 2001, Presetnik et al. 2001, Dietz et al. 2002,
Benda et al. 2003b, Decu 2003, Hulva et al. 2004). As P. pygmaeus was found at numerous
localities in Greece (Hanák et al. 2001), including its northern parts, it was ex­pec­ted to occur
also in Albania and in the countries of former Yugoslavia, where it has not been recorded yet.
Hence, taking into account the present state of knowledge, the range of pygmy pipistrelle in
southern Balkans, depicted by von Helversen & Holderied (2003) and reprinted by Vierhaus
247
Fig. 1. Male of Pipistrellus pygmaeus captured on 10 August 2003 near Qafëmal, northern Albania (K.
Sachanowicz).
Rys. 1. Samiec Pipistrellus pygmaeus odłowiony 10 sierpnia 2003 w pobliżu Qafëmal w północnej Albanii
(K. Sachanowicz).
& Krapp (2004) is misleading as giving an impression of its con­fir­med occurrence in southern
half of Albania and in Macedonia. This tentative range also covers a belt of southern Bulgaria
al­thou­ght only one record was known by then from sou­ther­n­most part of the country (Dietz
et al. 2002).
Single old records of bats belonging to small Pipistrellus complex are known from Ma­ce­do­
nia, Montenegro as well as from Bosnia and Herzegovina (e.g. Bolkay 1926, Đulić & Mirić
1967, Kryštufek et al. 1992, Jones 1999). The only record for Albania, published re­cent­ly
without any details – a male captured in a flat in Tirana – has to be assigned to P. pipistrellus
sensu lato (Uhrin et al. 1996).
In addition to the present knowledge of P. pygmaeus distribution, herein we report first data
on the species presence in Albania and in Bosnia and Herzegovina.
Material and methods
Data were collected in the Balkans during the trips in 2003–2004. Hunting bats were detected using a heterodyne detector Pettersson D100 and recorded with broadband Pettersson D980 detector and SONY
248
WM-D6C tape recorder. Recordings were analysed using BatSound 3.1 computer software (Pettersson
Elektronik AB, Sweden). For identification of the time-expanded calls we relied on characters given by
Zingg (1990), Russo & Jones (2002) and Obrist et al. (2004), while for determination of captured in­di­vi­
du­als – we followed Häussler et al. (1999), Sendor et al. (2002), von Helversen & Holderied (2003),
and Dietz & von Helversen (2004).
ReCORDS
P. pygmaeus was recently recorded at two localities in Albania (1–2) and at two sites in Bosnia and Her­
ze­go­vi­na (3–4):
(1) On 10 August 2003, two adult, sexualy active males were mist-netted over a small artificial pool
(42° 06’ N, 20° 06’ E), 731 m a. s. l., in a bed of a mountain stream, east of the Qafëmal village, along
the road Shkodra–Kukës (northern Albania, Pukë district). The habitat around the site comprised a small
settlement, cul­ti­va­ti­ons and overlogged mountain forest of the black pine Pinus nigra and the beech Fagus
sylvatica. More individuals of P. pygmaeus were observed and detected (Pettersson D100) at dusk when
hunting over the stream and sparse vegetation covering surrounding mountain slopes.
Measurements of two males were as follow: forearm length – 31.0 and 29.6 mm, fifth finger length
– 39.0 and 38.1 mm and weight – 5.5 and 5.0 g, respectively. Their pelage was light brown with yellowish
tinge on the back and with weak contrast between dorsal and ventral side, which was whitish-grey with
yellowish tinge. The muzzle and ears (7.9 mm) were relatively short (Fig. 1), light brown but slightly
Fig. 2. The sonogram and power spectrum graph of time-expanded echolocation calls of P. pygmaeus
recorded near Tepelenë, southern Albania, on 19 April 2004.
Rys. 2. Sonogram i wykres widma mocy sygnałów echolokacyjnych P. pygmaeus nagranych w okolicy
Tepeleny w południowej Albanii, 19 kwietnia 2004.
249
darker than the pelage; the skin was lighter, pinkish particularly around the eyes and in the basal part of
the ears. Both individuals had pronounced small ridge between the nostrils. Penis was yellowish without
paler medial stripe. Glandular bumps inside of mouth were of similar yellowish colour. Basal part of the
uropatagium was densely covered with fur on dorsal side, a condition usually seen in P. nathusii (Key­ser­
ling et Blasius, 1839). After being identified, sexed, measured and photographed the bats were released.
(2) P. pygmaeus was also recorded in the Vjosës valley, ca 1 km south of Tepelenë (40° 17’ N, 20° 02’ E),
114 m a. s. l. Soon before the dusk (19.10 p.m.), on 19 April 2004, several individuals were observed and
detected with Pettersson D980 (Fig. 2), while hunting above a small branch of the river and near the tree
crowns in an old stand of oriental planes Platanus orientalis and poplars Populus sp. Apparently, these
bats were shortly after leaving their roost, which could be located in a tree hole or crevice, since no other
type of shelters was available nearby.
The mean parameters of recorded calls were: frequency of maximum intensity: 57.2 kHz (range 55–60 kHz, n=12), highest frequency: 78.8 kHz (range 67–99 kHz, n=8), lowest frequency:
55.4 kHz (range 54–57 kHz, n=9), pulse length: 5.5 ms (range 4–7 ms, n=8), interval length: 74.4 ms
(range 61–78 ms, n=10).
Fig. 3. The sequence of time-expanded search calls of P. pygmaeus (upper row of signals), recorded near
Miljevina, Bosnia and Herzegovina, on 2 May 2004. Frequency of maximum intensity shown in a power
spectrum graph is marked with a cross. Lower row of signals illustrates echolocation calls of Hypsugo
savii recorded simultaneously. High background noise level caused by turbulent water.
Rys. 3. Sekwencja sygnałów echolokacyjnych P. pygmaeus, nagranych w okolicach Miljewiny, w Bośni
i Hercegowinie, 2 maja 2004 (górny szereg). Krzyżyk na wykresie widma mocy wskazuje częstotliwość
maksymalnego natężenia dźwięku. Poniżej, na sonogramie, nagrane jednocześnie sygnały Hypsugo savii.
Wysoki poziom szumów tła powodowany był przez płynącą wodę.
250
(3) Miljevina (43° 31’ N, 18° 39’ E), 599 m a. s. l., approximately 2.5 km south of the village, in a rocky
canyon of Bistrica river (100–150 m in depth) on the Dinarian Mts, eastern Bosnia. The site was sur­roun­
ded by almost vertical limestone walls and steep slopes, partially overgrown with sparse mixed forest
(spruces and beeches). At sunset on 2 May 2004, a search calls of hunting individuals of P. pygmaeus
were detected and recorded with Pettersson D980 bat detector and the tape recorder.
The signals were FM-qcf calls (Fig. 3) of the following mean parameters: frequency of maximum in­
ten­si­ty: 57.2 kHz (range 55–62 kHz, n=12), highest frequency: 72.4 kHz (range 64–85 kHz, n=9), lowest
fre­quen­cy: 54.2 kHz (range 52–57 kHz, n=9), pulse length: 5.8 ms (range 5–7 ms, n=9), interval length:
76 ms (range 70–90 ms, n=11).
(4) Later, in the same night as above, echolocation and social calls of P. pygmaeus (composed of three
elements at frequency of 18 kHz) were recorded near Dobro Polje (43° 35’ N, 18° 31’ E), 1008 m a. s. l.,
ca. 0.5 km west of the village, at a bridge over Bistrica river, eastern Bosnia.
Discussion
Individuals captured near Qafëmal fitted well into the range of morphological variation of P.
pygmaeus, given by the most recent authors (Häussler et al. 1999, von Helversen & Hol­de­
ri­ed 2003, Dietz & von Helversen 2004), thus their identification, in this case also supported
by detector observation, appeared unambiguous. More attention should be paid to the species
determination based on recorded echolocation calls. Although in the northern Europe, acous­tic
identification of P. pygmaeus is one of the easiest among vespertilionid bats, it is much more
complicated in the Meditterranean zone, where the species occurs sympatrically with Mi­ni­op­te­rus schreibersii (Kuhl, 1817) and both these species often share the same hunting habitats
(Russo & Jones 2003). Noteworthy, although M. schreibersii is taxonomically distant and be­longs
to a different family (Miniopteridae), it reveals echolocation calls sup­ri­sin­gly similar to those
emitted by P. pygmaeus. Its structure (FM-qcf) strongly resembles ty­pi­cal pipistrelle signals.
Mean value of its frequency of maximum intensity (53.9 kHz) is slight­ly lower than that recorded
in P. pygmaeus (56.2 kHz), similarly as the lowest frequency (47.4 and 51.5 kHz, re­spe­cti­ve­ly
– Obrist et al. 2004). However, ranges of these parameters overlapp (frequency of ma­xi­mum
intensity: M. schreibersii 49–62 kHz, P. pygmaeus 53–63 kHz; lowest frequency: 47–55 kHz
and 53–60 kHz, respectively – Russo & Jones 2002), thus large portion of their calls from sou­
thern Europe can be determined only with multivariate statistical methods, e.g. discriminant
function analysis (Russo & Jones 2002) or synergetic pattern recognition (Obrist et al. 2004).
Identification of pygmy pipistrelle with narrowband heterodyne detectors is widely practised
in Europe (e.g. Limpens 2000, 2001, Wermundsen & Siivonen 2004), but in the Mediterranean
it might be reliable only when supported with visual determination (flight and hunting style,
size of the bat and its wings shape – short and moderately broad in P. pygmaeus vs. long and
narrow in M. schreibersii). The lowest fre­quen­ci­es of some P. pygmaeus signals recorded in
Albania and in Bosnia were higher than the upper range of this parameter in M. schreibersii
(see also Zingg 1990), so the determination may be regarded as unambiguous. In the second
Bosnian locality (Dobro Polje), the pygmy pi­pis­t­rel­le was also confirmed by recorded social
calls with their typical, three-element structure (Barlow & Jones 1997).
The range of P. pygmaeus covers most likely the whole territory of the Balkan Peninsula. Its
relatively numerous localities are known only from Slovenia (Presetnik et al. 2001, Kryštu­fek
& Donev 2005) and from Greece (Hanák et al. 2001, Mayer & von Helversen 2001), that both
belong to fairly well studied countries. For other Balkan states, mostly single re­cords are avai­
lable, mainly due to the insufficient recording and low intensity of acoustic surveys. Therefore,
251
several localities, scattered over southern and western regions, were pu­b­lis­hed for Romania
(Limpens 2000, Decu 2003, Hulva et al. 2004). Single records are known also for Croatia and
Serbia (Limpens 2001), Bulgaria (Dietz et al. 2002, 2005, Benda et al. 2003b) and for the European part of Turkey (Benda et al. 2003a). Supplementing the picture of P. pygmaeus distribution in this part of Europe, we found the species in the least surveyed countries: Albania and
Bosnia and Herzegovina (this paper). Recently, the species was recorded also in Montenegro,
where two foraging bats were observed and detected on 10 July 2004 near the old fortress of
Budva, located on the Adriatic coast (C. Dietz – pers. comm.). At present, only Macedonia has
no presence of this species confirmed on its ter­ri­to­ry, for which only records of P. pipistrellus
(Schreber, 1774) s. str. are known (Kryštufek et al. 1992, B. Kryštufek – pers. comm.).
Data on habitat use by the species in the Balkans are also scarce. In Slovenia, P. pygmaeus was
observed hunting mainly in riparian habitats, over ponds, lakes and rivers, below 650 m a. s. l.
(Presetnik et al. 2001). In all cases described above, P. pygmaeus was recorded in the vicinity
of water, over small pool and in forested river valleys, what is in accordance with its habitat
preferences in other parts of Europe (Russo & Jones 2003, Bartonička & Řehák 2004).
Finally, the presence of both small pipistrelles has been confirmed for Albania, as P. pi­pis­t­
rel­lus s. str. was recorded recently for the first time at several localities (authors’ unpub­lis­hed
data). It is obvious that both these Pipistrellus species live in sympatry also in other Balkan
countries, as it was reported for Slovenia, Croatia, Serbia, Romania, Bulgaria and for Greece (Limpens 2000, 2001, Hanák et al. 2001, Mayer & von Helversen 2001, Presetnik et al.
2001, Benda et al. 2003, Decu 2003, Kryštufek & Donev 2005). However, relevant data on
P. pygmaeus abundance are available only for Slovenia, where it appears less frequent (44 localities) than P. pipistrellus (99 sites) (Kryštufek & Donev 2005). On the other hand, Hanák
et al. (2001) suggest that in Greece the former species is more abundant, but this conclusion is
based on small number of records (15 and 6, respectively), hence it may be misleading due to
the insufficient recording.
Streszczenie
Dwa samce Pipistrellus pygmaeus odłowiono w sierpniu 2003 w pobliżu miejscowości Qafëmal (pó­ł­noc­na
Albania). Żerujące osobniki tego gatunku obserwowano w nadrzecznym łęgu w dolinie Vjosy w oko­li­
cach Tepeleny (południowa Albania) w kwietniu 2004. Są to pierwsze stwierdzenia karlika drobnego w
Albanii. P. pygmaeus został także po raz pierwszy stwierdzony w Bośni i Hercegowinie, gdzie w maju
2004, w okolicach Miljewiny i Dobrego Polja nagrano sygnały echolokacyjne tego gatunku.
Acknowledgements
Authors express cordial thanks to Boris Kryštufek for valuable information on P. pipistrellus distribution
in Macedonia, giving an access to some hardly available publications and comments on the manuscript,
while to Christian Dietz for his own unpublished observation from Montenegro.
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Lynx (Praha), n. s., 37: 255–262 (2006).
ISSN 0024–7774
Small mammal fauna of the Kraków metropolitan area (southern Poland)
– problem of synurbisation (Insectivora, Chiroptera, Rodentia)
Drobne ssaki dawnego województwa krakowskiego – problem synurbizacji
(Insectivora, Chiroptera, Rodentia)
Katarzyna Stanik & Bronisław W. Wołoszyn
Institute of Systematics and Evolution of Animals, Polish Academy of Sciences, ul. Sławkowska 17,
PL–31-016 Kraków, Poland; [email protected]; [email protected]
received on 6 June 2006
Abstract. Current knowledge regarding the occurrence of small mammal species in the Kraków agglomeration and adjacent areas is presented. Special attention is given to urbanised species and their population
characteristics.
INTRODUCTION
Studies in urban ecology have been carried out in Poland for at least 30 years. Their aim is an
assessment of the current state and future prospects regarding the natural environment in urban
and suburban areas, and its role in creating optimal living conditions for the urban population of
people. Therefore the research is focused on three major aspects: soil science, flora and fauna
(Karolewski 1981).
The occurrence, abundance and structure of animal assemblages in urbicenosis are not intentionally shaped by man, in contrast to floristic ones (Andrzejewski 1975). However, such factors
as climate, technical infrastructure, flora, human population and environmental pollution have
a deep impact – termed “urbanisation pressure” – on urban fauna and its distribution (Markowski
1997). Observation of its effects has led to creation of new terms in urban ecology. One of these
is “antropopressure” defined as a mode of human impact on natural environment. Another term
is “synanthropisation”, covering a whole range of changes in flora and fauna. Organisms that
are well adapted to new conditions are named “synanthrophic species” (Pietraszewska 1999).
“Synurbisation” is defined as an adaptation to populate core urban areas. Synurbisation requires a
wide spectrum of ecological tolerance (eurytopic and polytopic species) and considerable species
adaptability. From the ecological perspective, the enrichment of relatively homogeneous urban
green areas with new species is favourable, because it makes the ecosystem more complicated
and thus more stable and self-sufficient (Gliwicz 1980).
The first studies of small mammal fauna in Kraków were carried out already in the 1940s
(Kowalski 1950). Nevertheless, the state of knowledge regarding the urban fauna is still insufficient (Karolewski 1981). So far, neither a detailed assessment of changes in the fauna
structure caused by urban anthropogenic factors, nor a holistic description of the current state
of urban fauna have been made.
255
Table 1. Climate parameters of Kraków
Tab. 1. Parametry klimatyczne Krakowa
month
January
February
March
April
May
June
July
August
September
October
November
December
temperature [°C]
precipitation [mm]
–2.9
–1.4
2.6
8.6
14.1
17.5
19.3
18.4
14.4
8.8
3.8
–0.2
34
34
35
42
57
86
95
83
56
46
42
34
annual average temperature 8.6
annual precipitation sum 644
This paper is aimed at the description of species composition of small mammal assemblages
in natural and anthropogenic biocenoses of the Kraków metropolitan area. The topographic and
climatic differentiation of neighbouring areas has a great impact on the urban environment,
which makes it an interesting object to study mammal habitat preferences. Despite the fact that
synurbisation of fauna is a dynamic and intensifying process, it is rarely investigated. Therefore,
the presented data may be helpful in planning similar studies in other areas.
MATERIAL AND METHODS
Area under study
The Kraków agglomeration lies at the place of contact between several physiographic units: Małopolska
Uppland in the north, Krakowsko-Wieluńska Uppland in the northwest (from where the Atlantic air masses
flow in), Beskid Zachodni Mts. in the south, and Sandomierz Lowlands in the east (where continental
climate dominates, Table 1). Boundaries of the area overlap with the boundaries of the former Kraków
voivodship that existed between 1975 and 1998.
Faunal data sources
Information on assemblages of small mammal species in the area comes from a large number of publications listed in Table 2. The Jaccard’s index (Q), describing similarity between assemblages, was calculated
according to the formula: Q = (c / a+b–c)×100, where Q = Jaccard’s index, a – number of species in
assemblage 1 (here: centre of the Kraków agglomeration), b – number of species in assemblage 2 (here:
natural areas and semi-natural suburban areas), c – number of species common for both assemblages.
RESULTS
56 species of small mammals occur in Poland, which constitutes 62% of the total number of
mammal species (90) of the country. In the study area, 43 small mammal species have been
256
Table 2. List of small mammal species recorded in the former Kraków voivodship.
Tab. 2. Lista gatunków Micromammalia zasiedlających województwo krakowskie.
A – agglomeration / aglomeracija; V – voivodship / województwo
species \ occurrence in Kraków:
A
V
reference
Talpa europaea (Linnaeus, 1758)
+
+
Sorex araneus (Linnaeus, 1758)
+
+
Sorex minutus (Linnaeus, 1766)
–
+
Neomys fodiens (Pendant, 1771)
–
+
Neomys anomalus (Cabrera, 1907)
–
+
Crocidura leucodon (Hermann, 1780)
–
+
Crocidura suaveolens (Pallas, 1811)
+
+
Rhinolophus hipposideros (Bechstein, 1800)
+
+
Rhinolophus ferrumequinum (Schreber, 1774)
–
+
Myotis myotis (Borkhausen, 1797)
+
+
Myotis nattereri (Kuhl, 1817)
–
+
Myotis emarginatus (Geoffroy, 1806)
+
+
Myotis mystacinus (Kuhl, 1817)
–
+
Myotis brandtii (Eversmann, 1845)
–
+
Myotis dasycneme (Boie, 1825)
–
+
Myotis daubentonii (Kuhl, 1817)
–
+
Vespertilio murinus (Linnaeus, 1758)
+
+
Eptesicus nilssonii (Keyserling et Blasius, 1839) –
+
Eptesicus serotinus (Schreber, 1774)
–
+
Pipistrellus pipistrellus (Schreber, 1774)
–
+
Nyctalus noctula (Schreber, 1774)
–
+
Nyctalus leisleri (Kuhl, 1818)
–
+
Plecotus auritus (Linnaeus, 1758)
–
+
Plecotus austriacus (Fischer, 1829)
+
+
Barbastella barbastellus (Schreber, 1774)
+
+
Sciurus vulgaris (Linnaeus, 1758)
+
+
Cricetus cricetus (Linnaeus, 1758)
+
+
Clethrionomys glareolus (Schreber, 1780)
+
+
Arvicola terrestris (Linnaeus, 1758)
+
+
Pitymys subterraneus +
+
(de Sélys-Longchamps, 1836)
Microtus oeconomus (Pallas, 1776)
–
+
Microtus agrestis (Linnaeus, 1761)
+
+
Microtus arvalis (Pallas, 1779)
+
+
Mus musculus (Linnaeus, 1758)
+
+
Rattus norvegicus (Berkenhout, 1769)
+
+
Micromys minutus (Pallas, 1771)
–
+
Apodemus agrarius (Pallas, 1771)
+
+
Apodemus microps (Kratochvíl et Rosický, 1952) –
+
Apodemus sylvaticus (Linnaeus, 1758)
+
+
Apodemus flavicollis (Melchior, 1834)
+
+
Dryomys nitedula (Pallas, 1779)
–
+
Glis glis (Linnaeus, 1766)
–
+
Muscardinus avellanarius (Linnaeus, 1758)
–
+
Kowalski 1950, Rzebik-Kowalska 1972
Skiba 2005, Haitlinger & Szyszka 1977
Grodziński et al. 1958
Ruprecht 1976
Skuratowicz & Warchalewski 1954
Sałata-Piłacińska 1977
Simm 1952, Ruprecht 1976
Harmata 2000
Nowak et al. 2001
Harmata 1994
Harmata 1960, 1962
Harmata 1969, Harmata & Wojtusiak 1963
Ruprecht 1974
Ruprecht 1974
Biała cave
Kowalski 1953
Kowalski 1957
Ojców
Kowalski 1953, Harmata 1960
Harmata 1960
Harmata 1962, Markowski & Suskiewicz 1981
Harmata 1960
Harmata 1969
Ruprecht 1971, Harmata 1969
Kowalski et al. 1957
Kowalski 1950, Udziela 1925
Kowalski 1950, Skiba 2005
Skiba 2005,
Skuratowicz & Warchalewski 1954
Skiba 2005, Kowalski 1960
Skiba 2005, Haitlinger & Szyszka 1977
Skiba 2005, Haitlinger & Szyszka 1977
Skiba 2005, Haitlinger & Szyszka 1977
Skuratowicz & Warchalewski 1954
Skuratowicz & Warchalewski 1954
Skiba 2005, Haitlinger & Szyszka 1977
Skiba 2005, Haitlinger & Szyszka 1977
Skiba 2005, Haitlinger & Szyszka 1977
Grodziński 1959
Kowalski 1950
Kowalski 1950
257
recorded in the last 50 years or so. Most of them represent 4 families: Soricidae, Vespertilionidae, Arvicolidae, and Muridae. Four of the recorded species are considered synurbic: Sciurus
vulgaris, Apodemus agrarius, Mus musculus, and Rattus norvegicus.
Among Insectivora, three out of seven species are common for both compared areas, within
Rodentia the proportion of common species is ⅔ of the total species number and in Chiroptera
only ⅓. In total, 21 out of 43 small mammal species inhabit both subareas: city centre and its
surroundings.
The Jaccard’s index, reflecting a degree of similarity of small mammal assemblages between
(a) core urban and (b) suburban and natural areas, equals 48.84.
DISCUSSION
Studies on structural changes in small mammal assemblages in urban areas have two practical
aspects. Firstly, mammals as animals living in human settlements are a potential source of
zoonoses. Secondly, small mammals may be useful as specific bioindicators of anthropogenic
environmental pollution. Another interesting problem is species composition of fauna inhabiting
buildings and various urban facilities, dominated by such species as Mus musculus, Rattus rattus
and Rattus norvegicus (Andrzejewski 1977).
The urban fauna is dominated by species of wide geographic ranges. Xerophilous and thermophilous species, mostly those characteristic of warmer climatic zones, occur inside buildings
(Markowski 1997). For the majority of city dwelling species communal waste is an abundant
food source. Urban areas offer free ecological niches but also impose many threats, e.g. in-
Figs. 1, 2. Percentage of small mammal species of the former Kraków voivodship in the whole small
mammal fauna of Poland (3, left) and the whole mammal fauna of Poland (4, right).
Rys. 1, 2. Liczba gatunków Micromammalia dawnego województwa krakowskiego na tle ogółu krajowej fauny drobnych ssaków (3, nalevo) i na tle ogółu krajowej fauny ssaków (4, napravo), wyrażona
w procentach.
258
Table 3. Number of small mammal species in the Kraków centre and in the remaining area of the former
Kraków voivodship
Tab. 3. Liczba gatunków Micromammalia w centrum Krakowa i na obszarze całego województwa krakowskiego
order
insectivores
rodents
bats
total / ogółem
Kraków agglomeriation
Kraków voivodship
Σ
3
12
6
21
7
18
18
43
7
18
18
43
creased predation risk (presence of cats, dogs and birds of prey) and many barriers limiting
migration (highways, walls, fences). Also a high level of pollution reduces species richness of
urban biocenoses (Pisarski & Trojan 1976).
The Kraków agglomeration and adjacent areas are situated in the Wisła river valley. Its relatively warm climate is under a strong influence of adjacent physiographic regions. This karstic
region is characterised by high faunistic diversity, enriched with several mountain species whose
northern distribution limits reach the area. The Wisła river and its tributaries flowing through the
Kraków agglomeration contribute to relatively high air humidity and provide feeding grounds
and shelters in riparian habitats. The Jaccard’s index (Q) for the city centre and suburbs is 48.84,
reflecting high faunal similarity of the two areas, probably caused by the presence of a narrow
belt of parkland surrounding the old town. Observations in the Las Wolski, a wood situated
on the western periphery of Kraków, have brought interesting information on many mammal
species from the following families: Arvicolidae, Muridae, Soricidae, and Gliridae (Glis glis,
Muscardinus avellanarius), and a very rare bat species Nyctalus leisleri (Harmata 1960).
It is known that synanthropic species show cyclical changes in population. In the years of
abundance the range of population is wedge-shaped, narrowing towards the city centre. Currently, Apodemus agrarius is regarded as a synurbic species and it is one of the most abundant
species in the Kraków centre. Despite the fact that the small mammal assemblage in the suburbs is relatively rich in species, very often, only Apodemus agrarius is able to populate the
core urban area (Gliwicz 1980). In contrast, Arvicolidae species have been observed more and
more rarely in the urban agglomeration, which is due to the small area of open habitats inside
the city (Kasprzak & Banaszak 1978). One may assume that the Kraków common grounds
(Błonia Krakowskie) and banks of the Wisła river are densely populated by Arvicolidae voles,
however, no data is available. Studies carried out so far have shown that the number of species
connected with forest, shrub and open habitats has been increasing in the urban areas where
they enter a mixture of ‘ecotone biocenoses’ (Ziomek 1998).
Preference of small mammals towards urban areas depends on season. In spring synanthropic
rodent species (Mus musculus, Rattus rattus and Rattus norvegicus) migrate from the city into
surrounding areas and in autumn they are forced to come back due to adverse weather conditions and lack of food. Also Microtus arvalis migrates into agrocenoses in spring (Chudoba
& Humiński 1963). Contrary to them, some Chiroptera hibernating outside the urban area (in
caves or tree hollows) tend to establish summer colonies in human settlements (in attics of old
houses and churches, crevices in buildings etc.) (Wołoszyn 1981).
259
STRESZCZENIE
Składem gatunkowym drobnych ssaków Krakowa zainteresowano się już w latach czterdziestych ubiegłego
stulecia (Kowalski 1950). Brak jak dotąd nie tylko dokładnej oceny zmian w strukturze fauny pod wpływem
miejskich czynników antropogenicznych, ale także całościowego obrazu stanu fauny terenów zurbanizowanych (Kasprzak & Banaszak 1978). Celem niniejszej pracy było określenie składu zespołów drobnych
ssaków w zbiorowiskach naturalnych, oraz antropogenicznych obszaru metropolitalnego Krakowa. Teren
badań wybrano ze względu na zróżnicowanie topograficzno – klimatyczne obszarów z nim graniczących
i mających wpływ na tutejsze środowisko i czyni go dobrym do analizowania preferencji siedliskowych
gatunków. Proces synurbizacji fauny jest zjawiskiem dynamicznym i nasilającym się, a mimo to relatywnie
rzadko badanym. Drobne ssaki w liczbie 56 gatunków stanowią 62,2% spośród 90 gatunków krajowej
teriofauny. W ciągu ostatnich około 50 lat na badanym obszarze zanotowano obecność 43 gatunków
drobnych ssaków. Wśród nich dominują gatunki, należące do czterech Rodzin: Soricidae, Vespertilionidae, Arvicolidae i Muridae. Cztery spośród zaobserwowanych gatunków, są uważane za synurbijne. Są
to Sciurus vulgaris (Linnaeus, 1758), Apodemus agrarius (Pallas, 1771), Mus musculus (Linnaeus, 1758)
i Rattus norvegicus (Berkenhout, 1769). Ogółem 21 spośród 43 gatunków drobnych ssaków zasiedlających
badany obszar, zasiedla zarówno centrum aglomeracji, jak i tereny otaczające.
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Lynx (Praha), n. s., 37: 263–273 (2006).
ISSN 0024–7774
Population ecology of Apodemus flavicollis in two lowland forest habitats
(Rodentia: Mu­ri­dae)
Populačná ekológia Apodemus flavicollis v dvoch typoch nížinného lesa
(Rodentia: Muridae)
Kristína Trubenová, Dávid Žiak & Peter Miklós
Department of Zoology, Faculty of Natural Sciences, Comenius University, Mlynská dolina B-1,
SK–842 15 Bratislava, Slovakia; [email protected], [email protected], [email protected]
received on 12 June 2006
Abstract. Several population characteristics were studied and compared between populations of Apo­de­
mus flavicollis inhabiting two different lowland forest habitats. Small mammals were live-trapped during
2000–2002 on two 1.44 ha plots in alder forest and oak-elm forest (Nature Reserve Jurský Šúr, Slovakia).
The density of A. flavicollis was 0.0–25.3 individuals per ha in alder forest and 3.1–28.0 individuals per
ha in oak-elm forest. Throughout the study, A. flavicollis was the most numerous species in oak-elm forest
(72–100%), while in alder forest its proportion was much lower (0–50%). Higher reproductive activity
in alder forest was probably caused by higher food and shelter availability in this area during breeding
season. Ani­mals of both sexes were heavier in oak-elm forest which suggests a better fitness of individuals
in this habitat. While the degree of male residency was significantly higher in oak-elm forest, we did not
find any difference in resident rate of females. Different occurrence and residency of individuals on the
study plots are probably caused by higher heterogeneity and larger total area of alder forest together with
temporary winter insufficiency of resources on the local study plot.
Introduction
The structure and dynamics of a population represent the outcome of the interactions between
life histories of its individuals and the spatial and temporal variations in its environment (Zeng &
Brown 1987). Local populations of widely distributed species living in different en­vi­ron­men­tal
conditions may exhibit different demographic parameters in relation to ecological cha­ra­c­te­ris­tics
of habitats (Montgomery & Gurnell 1985). The yellow-necked mouse, Apo­de­mus fla­vi­col­lis
(Melchior, 1834), as one of the most ubiquitous and abundant small mam­mals in the western
Palaearctic, utilises a variety of habitat types throughout its geographical range. It is primarily
a woodland rodent preferring mature successional stages of deciduous forests with an open
ground layer. However, it can also inhabit field-forest ecotones, grassy areas and brushwood sets
(Gurnell 1985, Jensen 1984). Its occurrence in coniferous fo­rests, subalpine and alpine zone
is mainly seasonal (Kratochvíl & Rosický 1952, Zima et al. 1984), depending on local seed
supply and immigration from nearby stable populations with higher densities (Martínková et al.
2001). Populations of A. flavicollis are able to persist in different habitat types because of their
broad flexibility in demography and habitat selection as well as high adaptability to different
environmental conditions (Douglass et al. 1992, Montgomery & Gurnell 1985).
263
In the present study we compared several demographic parameters (abundance, residency,
age composition breeding activity and body mass) between two populations of A. flavicollis
inhabiting different lowland forest habitats in National Nature Reserve Jurský šúr.
MATERIAL AND METHODS
Study area
The study was carried out in the Jurský Šúr National Nature Reserve, 12 km north-east of Bratislava
(128–132 m a. s. l., DFS 7769c). Study sites were situated in two different forest habitat types, an alder
forest (study plot A) and an oak-elm forest (plot B).
The alder forest forms an isolated complex of swampy boggy forest in the centre of wet meadows. It
spreads over an area of 334.7 ha and forms the greatest part of the Jurský Šúr National Nature Reserve.
The alder forest belongs phytocoenologically to the Carici elongatae-Alnetum medioeuropaeum as­so­ci­a­ti­
on. The tree layer of study plot A is formed by the following species: Alnus glutinosa, Quercus robur and
Ulmus laevis. The shrub layer develops mainly in the sites with loose canopy and is formed by the species
Alnus glutinosa, Salix cinerea, Prunus spinosa and Rosa spp. The herb layer is formed by the species Iris
pseu­da­co­rus, Urtica kiovenensis, Alisma plantago-aquatica and Carex spp. This plot is characterised by
the pre­sen­ce of surface floods (XI–II/III) when the water level reaches 20–50 cm (Kupcová 1980).
The oak-elm forest is a remnant of old thermophilous mixed forests of the Pannonian basin, partly
it keeps its forest-steppe character. It spreads over an area of 42.38 ha. The tree layer of study plot B is
represented by the species Quercus robur, Q. cerris, Ulmus laevis, Acer campestre, and Carpinus betulus.
Shrub layer is formed by the species Prunus spinosa, Rosa canina and Crataegus spp. The herb layer is
formed mainly by grasses (Poales). Besides them, the species Viola sp., Geum urbanum, Alliaria petiolata,
Allium ursinum, Lamium maculatum, and Galium aparine occur most frequently (Kupcová1980, Miklos
1999). The study sites were separated by approximately 1 km wide corn field.
Methods
Small mammals were live-trapped every six weeks between February 2000 and November 2002 on two
1.44 ha plots. Live-traps were set 15 m apart in a 9×9 matrix on each grid. A single Chmela-type live trap
baited with oats was placed at each station. Traps were set during six consecutive nights and checked twice
a day, mornings and evenings. All captured small mammals were individually marked by toe-clipping
and released at trapping points immediately after data collection. Upon capture the following parameters
were noted for each individual animal: capture point co-ordinates, species, individual number, sex, weight
and status of external sexual characters to assess reproductive individuals (males: scrotal testes; females:
enlarged nipples, per­fo­ra­ted vagina or pregnancy). Monthly population abundance was estimated by direct
enu­me­ra­ti­on of mice captured on the study plot, and population densities were obtained by dividing abundance by the trapping area, increased by one trap distance per side. Abundance variation was compared
between habitats using the chi-square goodness of fit test. All the individuals caught were divided into
three groups according to their residential status. The first group (long time residents) comprised animals
present throughout at least two series of trapping, the second group (short time residents) included individuals captured several times during the sampling session. The third group (transients) encompassed
those individuals that were caught only once and no more. Resident rate was expressed as the proportion
of long time residents excluding mice captured in the last trapping period of a year. The age of each individual was estimated on the basis of body weight according to Gliwicz (1988) and all individuals were
divided into adult (m>25 g) and young (subadult and juvenile; m<25 g) age classes. The following cha­
ra­c­te­ris­tics were analysed for each trapping session, in­di­vi­dual years and also jointly for the three years
of study: sex ratio, age structure (proportion of young), breeding intensity (proportion of reproductively
active fe­ma­les) and resident rate. These characteristics were com­pa­red between habitats using chi-square
analysis of con­tin­gen­cy tables. The body weights of sexually active and adult individuals of both sexes
264
were compared between habitats using the Mann-Whitney test. We tested the spatial distribution of either
sex as well as that of both sexes together to find possible departures from random (Poisson) distribution
using the χ2-test. When we found non-random spatial distribution, we used Lloyd (1967) index to decide
whether the distribution was clumped or even.
RESULTS
The total material consisted of 1,927 captures of 617 individuals of Apodemus flavicollis (Mel­
chi­or, 1834), comprising 914 captures of 301 individuals in alder forest and 1,013 cap­tu­res of
316 individuals in oak-elm forest. Besides that, the following species were recorded: in the alder
forest Clethrionomys glareolus (Schreber, 1780), Sorex araneus Linnaeus, 1758, Neomys fodiens
(Pennant, 1771), Micromys minutus (Pallas, 1771) and Mustela nivalis Lin­nae­us, 1766, and in
the oak-elm forest Clethrionomys glareolus, Sorex araneus, Crocidura leucodon (Hermann,
1780), Micromys minutus and Mustela nivalis.
The density of A. flavicollis was 0–25.33 individuals per ha on plot A and 3.11–28 in­di­vi­du­als
per ha on plot B. During the reproduction season (III/IV–IX/X) we did not find any dif­f­e­ren­ces
in abundance variation between the study sites. Statistically significant difference (χ2=105.8139,
p<0.01, df=19) in variation of abundance found, when comparing the whole study period, was
caused by much lower minimum numbers and even absence of individuals on plot A during
winter months (II–IV 2000, II 2001, III 2002).
Throughout the study, the proportion of A. flavicollis in small mammal community was dif­f­
e­rent between the two study sites. In the oak-elm forest, A. flavicollis was the most nu­me­rous
Fig. 1. Resident rate of males in alder forest (pooled data for 2-months intervals).
Obr. 1. Rezidencia samcov v jelšovom lese (zhrnuté dáta pre dvojmesačné obdobia).
265
Fig. 2. Resident rate of males in oak-elm forest (pooled data for 2-months intervals).
Obr. 2. Rezidencia samcov v dubovo-brestovom lese (zhrnuté dáta pre dvojmesačné obdobia).
species, its proportion in small mammal community reaching 72 to 100%. In the alder forest, the
proportion of A. flavicollis ranged from 0 to 50%. Small mammal abundance was always higher
in the alder forest (on the average 3.5-times higher, during peak abundance 2.8–4.3 times).
We found higher resident rate on plot B when comparing pooled data for the whole study
period for all individuals (χ2=8.37, p<0.05) and for males (χ2=13.27, p<0.05) (Figs. 1, 2).
Higher resident rate of males on plot B was statistically significant also for pooled data for 2000
(χ2=6.42, p<0.05), 2001 (χ2=4.01, p<0.05), and in 2002 it was close to the limit of statistical
significance (χ2=3.73, p=0.054). A similar trend can be observed also in the in­di­vi­dual trapping
series, but the difference was statistically significant only in IX 2000 (χ2=8.31, p<0.05), VIII
2001 (χ2=4.82, p<0.05) and VIII 2002 (χ2=9.72, p<0.05). We did not find any difference in
resident rate of females.
We did not find any difference in timing of reproduction (sexually active individuals and
juveniles were recorded during the same trapping series on both plots). Reproduction period
started in III/IV and ended in IX/X.
In the alder forest we found a higher proportion of sexually active females, this difference
being statistically significant for the pooled data for the whole study period (χ2=5.7, p<0.05),
and for the pooled data for the year 2001 (χ2=8.14, p<0.05) (Fig. 3a, b).
In alder forest we found also a higher proportion of young (subadult and juvenile) in­di­vi­du­
als, this difference being statistically significant for the pooled data for the whole study pe­ri­od
(χ2=4.17, p<0.05) (Fig. 4a, b). A higher proportion of young individuals in the alder forest was
observed also in individual trapping series, the difference being statistically significant in VIII
2001 (χ2=4.37, p<0.05) and X 2002 (χ2=5.2, p<0.05).
266
Table 1. Mean body weights and standard deviations for sexually active and adult individuals of Apode­
mus flavicollis
Tab. 1. Priemerné hmotnosti pohlavne aktívnych a adultných jedincov Apodemus flavicollis
females
males
alder forest (m±SD) [g]
active
adult
29.98±3.68
31.96±5.00
28.78±4.68
30.60±6.23
oak-elm forest (m±SD) [g]
active
adult
31.45±4.21
32.25±4.99
31.79±4.59
33.32±5.95
Average weights of sexually active and adult individuals were higher in the oak-elm forest,
this differences being statistically significant only for females (sexually active females: w=5,340,
p<0.01, adult females: w=9,433, p<0.01) (Table 1).
Spatial distribution of long-time residents on plot A was random in most trapping series,
with the exception for females in November 2000 (χ2=6.50, p<0.05, L=1.22) and August 2001
(χ2=6.10, p<0.05, L=2.24) and for both sexes together in October 2001 (χ2=9.63, p<0.01,
L=1.68), when the spatial distribution was aggregated. Spatial distribution of males was random
in all trapping series. On plot B we found aggregated spatial distribution for ma­les in February
(χ2=20.01, p<0.001, L=3.83), April (χ2=18.12, p<0.001, L=4.13), June 2001 (χ2=12.66, p<0.01,
L=2.88) and in August 2002 (χ2=6.57, p<0.05, L=2.88). For both sexes together we found aggregated dis­tri­bu­ti­on in September (χ2=53.59, p<0.001, L=1.97) and November 2000 (χ2=10.44,
Fig. 3a. Proportion of sexually active females in alder forest (pooled data for 2-months intervals).
Obr. 3a. Zastúpenie pohlavne aktívnych samíc v jelšovom lese (zhrnuté dáta pre dvojmesačné obdobia).
267
Fig. 3b. Proportion of sexually active females in oak-elm forest (pooled data for 2-months intervals).
Obr. 3b. Zastúpenie pohlavne aktívnych samíc v dubovo-brestovom lese (zhrnuté dáta pre dvojmesačné
obdobia).
Fig. 4a. Proportion of young individuals in alder forest (pooled data for 2-months intervals).
Obr. 4a. Zastúpenie mladých jedincov v jelšovom lese (zhrnuté dáta pre dvojmesačné obdobia).
268
p<0.05, L=1.84), February (χ2=6.05, p<0.05, L=2.43), April (χ2=8.05, p<0.05, L=2.45) and July
2001 (χ2=15.22, p<0.01, L=1.79) and February (χ2=11.69, p<0.001, L=3.31) and August 2002
(χ2=9.32, p<0.01, L=2.80). Spatial distribution of fe­ma­les was random in all trapping series.
DISCUSSION
Populations under study revealed typical seasonal trend of abundance fluctuation known for
this species (Flowerdew 1985, Gurnell 1985, Montgomery 1979, Pucek et al. 1993) with peak
numbers in late summer/early autumn and decrease during late autumn and winter months.
Maximal densities achieved on both plots are consistent with published data for A. flavicollis
in Central Europe (Mazurkiewicz & Rajska-Jurgiel 1998, Wojcik & Wolk 1985).
The most part of the study plot A was flooded and frozen during winter (XI–II/III), so as
regards abundance and availability of food, which is the most important factor governing local
distribution and abundance of A. flavicollis (Angelstam et al. 1987, Montgomery 1979), as
well as regarding the availability of shelters, it represents an unfavourable environment for this
species (Montgomery 1978). The animals retreat from this study plot during winter. However,
after a temporary absence during winter we repeatedly captured the same in­di­vi­du­als on plot
A as in the previous year, and in quite high numbers (Trubenová et al. 2004). The animals inhabiting study plot A are able to survive winter, despite the fact that they are forced to seek out
more favourable refugees for overwintering, and probably the refugees inhabited during winter
are within easy reach for this species. In contrast to study plot in alder forest, the continuous
Fig. 4b. Proportion of young individuals in oak-elm forest (pooled data for 2-months intervals).
Obr. 4b. Zastúpenie mladých jedincov v dubovo-brestovom lese (zhrnuté dáta pre dvojmesačné obdobia).
269
occurrence of individuals of A. flavicollis on the study plot in oak-elm forest sug­gests that this
habitat is suitable for A. flavicollis all year over, also during winter months (Trubenová et al.
2004). This seems consistent with the findings of Castien & Gosalbez (1994) and Montgomery
(1979), according to whom A. flavicollis is quite rare in many forest types during winter and
the populations are restricted to one habitat type characterised by the presence of a number of
seed-bearing tree species and by a dense en­tan­g­le­ment of branches above ground level. This
way could be characterised the study plot in the oak-elm forest as well as the oak-elm forest
as a whole.
Distinct difference in the proportion of A. flavicollis in the small mammal community between
these two habitats is caused by the much higher numbers of small mammals in the alder forest
(especially Clethrionomys glareolus), which suggests higher productivity and/or higher he­te­ro­
ge­ne­i­ty of this habitat (M’Closkey 1976, Van Horne 1983). Almost mo­no­spe­ci­fic occupancy
of plot B by A. flavicollis is in contrast with the situation in alder forest, where the numbers
of A. flavicollis were similar or lower than the numbers of C. glareolus. However, we do not
suppose that the presence of C. glareolus limits the numbers achieved by A. flavicollis population. Where these two species occur together, A. flavicollis is generally considered dominant
(Andrzejewski & Olszewski 1963, Bergstedt 1965, Grüm & Bujalska 2000), affecting po­pu­
la­ti­on dynamics, spatial distribution and/or demographic variables of C. glareolus population,
but not conversely.
The different residency of males and females is caused by different reproductive strategies
between sexes. Females are characterised by lower mobility, higher residency and selection of
places providing secure cover for rearing the young, while males are distributed with re­spect
to breeding opportunity (Ostfield 1985). As we have found difference only in the resident rate
of males, while the resident rate of females did not differ between the plots, we can suppose
that higher residency of males in oak-elm forest was not caused only by higher quality of this
habitat. In oak-elm forest we have found also longer residency time of in­di­vi­du­als of both
sexes, sig­ni­fi­cant­ly for males (Trubenová et al. 2004). Differences in male residency could be
associated with more opportunity to disperse in alder forest, and also with higher heterogeneity
of this habitat (Trubenová et al. 2004). The alder forest seems to be a highly seasonal environment in which different habitat patches may be preferred at different seasons. Spatial activity
and residency of Apodemus spp. are affected not only by the wo­od­land habitat itself but also
by the surrounding landscape, distances to neighbouring woodlands and connectivity between
them (Kozakiewicz 1993, Ylönen et al. 1991). In smaller woodlands, where the surrounding
area of ancient woodland tends to be lower, there is less opportunity to disperse (Marsh &
Harris 2000). Brown (1966) concluded that a sub­stan­tial proportion of male Apodemus have
different areas of activity at different times. She reported that the individuals concerned were
the dominant males having very large home-ranges and visiting various parts of these ranges
on a regular rotation basis. So the lifetime home ranges of A. flavicollis males in oak-elm forest
are probably smaller, the animals have fewer seasonal centres of activity, and they stay longer
in the same place. Aggregated dis­tri­bu­ti­on of males in oak-elm forest can be connected with
their higher residency and restricted spatial activity in this habitat.
Balanced sex ratio found in both populations (Trubenová et al. 2004) is a characteristic
feature of stable populations living in optimal habitats. Although we have found no difference
in timing and duration of reproduction period, the higher proportion of sexually active fe­ma­les
and young individuals in alder forest indicate a higher intensity of reproduction in this habitat.
As the numbers of females active sexually may be a good indicator of the quality of a given
270
habitat (Mazurkiewicz 1991, 1994), the alder forest seems more suitable for A. flavicollis during
breeding season. Aggregated spatial distribution of females in alder forest can be con­nec­ted
with locally and temporally more favourable conditions, which cause con­cen­tra­ti­on of females
in this habitat during the time it provides suitable conditions for we­a­ning the litter.
Higher numbers of sexually active females in alder forest could have also another reason. In
oak-elm forest the individuals remain all year over, as they find suitable conditions for sur­vi­ving
the winter there. So the structure of the local population is stable, with well-defined dominance
hierarchy. Social interactions may prevent young animals from entering the re­pro­du­cing part of
population (Mazurkiewicz & Rajska-Jurgiel 1998). In contrast, the study plot in alder forest is
not populated by A. flavicollis individuals during winter and the population in spring consists
mainly of immigrants. In such environment, where there is no stable structure of the population and where at that time there is no social interaction factor, every spring and early-summer
immigrant may mature and breed (Mazurkiewicz & Rajska-Jurgiel 1998, Van Horne 1983).
The higher body weight of individuals in the oak-elm forest found, when comparing all in­
di­vi­du­als (Trubenová et al. 2004), can be partly caused by the higher proportion of adults in
this habitat. However, higher body weights were found in this habitat also when comparing
sexually active individuals (although significantly only for females), and also when com­pa­ring
adult individuals (significantly for females), which suggest that individuals living in the oak-elm
forest are heavier. It is used to present body weight as one of the measures of fitness, which
provides information about habitat quality for the studied species, as animals in higher-quality
habitats tend to be heavier (Van Horne 1983). Higher body weight of animals of both sexes in
oak-elm forest suggests better fitness of individuals in this habitat, which is probably caused
by more suitable year-long conditions here in comparison to the alder forest, where the fitness
of individuals may be lowered by the cost of winter migration.
SÚHRN
V práci sú vyhodnotené a porovnané demografické parametre populácií Apodemus flavicollis, obý­va­j­ú­
cich dva typy nížinného lesa. Terénny výskum prebiehal v rokoch 2000–2002 v jelšovom lese a dubovo-brestovom lese (NPR Šúr, SR) na odchytových kvadrátoch veľkosti 1,44 ha s použitím CMR metódy.
Počas obdobia výskumu bolo v jelšovom lese zaznamenaných 914 od­chy­tov 301 jedincov a v dubovo-brestovom lese 1013 odchytov 316 jedincov A. flavicollis. Denzita druhu dosahovala hodnoty 0,0–25,3
jedincov na hektár v jel­šo­vom lese a 3,1–28,0 je­din­cov na hektár v dubovo-brestovom lese. Zatiaľ čo
v dubovo-brestovom lese bol A. flavicollis najpočetnejším druhom počas celého obdobia (72–100 %),
v jelšovom lese bolo jeho zastúpenie výrazne nižšie (0–50 %). Obdobie rozmnožovania, stanovené na
základe odchytávania pohlavne aktívnych a juvenilných jedincov trvalo v jednotlivých rokoch v obidvoch
prostrediach rov­na­ko. Vyšší podiel pohlavne aktívnych samíc, ako aj juvenilných jedincov v jelšovom
lese po­u­ka­zu­je na vyššiu reprodukčnú aktivitu v tomto prostredí. V dubovo-brestovom lese bol zistený
vyšší podiel dlhodobo rezidentných samcov, rezidencia samíc sa medzi sledovanými pro­stre­di­a­mi nelíšila.
U oboch pohlaví bola zistená vyššia hmotnosť jedincov v dubovo-brestovom lese, čo naznačuje lepšiu
kondíciu jedincov v tomto prostredí. Rozdiely v rezidencii je­din­cov medzi sledovanými lokalitami boli
pravdepodobne spôsobené väčšou rozlohou a heterogenitou jel­šo­vé­ho lesa, súčasne s jeho nevhodnými
podmienkami počas zimného obdobia.
ACKNOWLEDGEMENTS
This work was carried out with the financial support of the grants by VEGA No. 1/7197/20 and 1/0017/03.
We also thank Lenka Trebatická for her help with the field-work.
271
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Lynx (Praha), n. s., 37: 275–284 (2006).
ISSN 0024–7774
Karyotype analysis of Murina suilla and Phoniscus atrox from Malaysia
(Chiroptera: Murininae, Kerivoulinae)
Karyotypová analysa dvou netopýrů z Malajsie: trubkonosa vepřího (Murina suilla)
a vlnouška vroubkozubého (Phoniscus atrox) (Chiroptera: Murininae, Kerivoulinae)
Marianne Volleth
Department of Human Genetics, Leipziger Str. 44, Otto-von-Guericke University, D–39120 Magdeburg,
Germany; [email protected]
received on 26 May 2006
Abstract. For the first time, chromosomal data are presented for Murina suilla (2n=44, FN=58) and
Pho­ni­s­cus atrox (2n=40). The karyotype of Murina is distinguished from that of Myotis by the presence
of euchromatic short arms on chromosomes 12, 13 and 15. The chromosomal complement of Phoniscus
atrox is characterized by extensive addition of heterochromatic short arms and a paracentric inversion in
chro­mo­so­me 8. The reduction in the diploid chromosome number is due to Robertsonian fusion of chro­
mo­so­mal arms 8+13 and 11+14, respectively.
Introduction
In recent years, progress in molecular genetic methods yielded fascinating new insights into bat
phylogeny (for references see Eick et al. 2005, Teeling et al. 2005) and let, together with mor­
pho­lo­gi­cal and call frequency differences, to the discovery of cryptic species (e.g. Pi­pis­t­rel­lus
pygmaeus, Barrat et al. 1997, Mayer & von Helversen 2001; Plecotus macro­bul­la­ris, Kiefer
& Veith 2001, Kiefer et al. 2002; Myotis alcathoe von Helversen et al. 2001, etc.).
The members of the vespertilionid subfamilies Murininae and Kerivoulinae, however, ga­i­ned
less interest than most other vespertilionid genera. This is not only true for molecular genetic
investigations but also for cytogenetic studies.
Up to now, out of 15 recognized species of Murina Gray, 1842 (13 in Nowak 1994 plus two
newly descri­bed species: Maeda & Matsumura 1998, Csorba & Bates 2005), chromosomal
and fundamental numbers have been reported for six species only. Banded karyotypes have
been studied only in Murina ussuriensis Ognev, 1913 and M. hilgendorfi Peters, 1880 (Harada
et al. 1987).
Even less is known from the subfamily Kerivoulinae. Out of the 23 species recognized (Nowak
1994, Vanitharani et al. 2003, Bates et al. 2004), conventionally stained ka­ry­o­typ­es have been
presented only from Kerivoula lanosa (Smith, 1847) (Rautenbach et al. 1993) and K. pa­pil­lo­sa
(Temminck, 1840) (McBee et al. 1986).
In this paper we present detailed karyological information of Murina suilla and Phoniscus
atrox from Malaysia together with a literature overview.
275
Material and methods
Specimens examined
Murina suilla, 1 male, 1 female, Templer Park near Kuala Lumpur, SMF 69348, 69349, 1 male, Ulu
Gombak Field Studies Centre, Selangor, Malaysia, SMF 69350.
Phoniscus atrox, 1 female, Ulu Gombak Field Studies Centre, Selangor, Malaysia, SMF 69316.
(SMF = catalogue number of the Senckenberg Museum, Frankfurt am Main, Germany)
Chromosome preparations and analysis
Cell culture and cytogenetic procedures were done as described in Volleth (1987) and Volleth et al.
(2001). Chromosomal arms were numbered according to Bickham’s scheme for American Myotis species
(Bickham 1979). Karyotypes were compared with the basic karyotype of Vespertilionidae (Volleth &
Hel­ler 1994)
Results
Murina suilla (Temminck, 1840)
The karyotype of Murina suilla shows a diploid chromosome number (2n) of 44 and a fun­da­
men­tal number of autosomal arms (FN) of 58. The G-banded karyotype of a male specimen is
shown in Fig. 1. Chromosomes 7, 12, 13 and 15 show a small G-negative short arm. C-ban­ding
revealed that these short arms consist of euchromatic material (Fig. 2). Therefore these short
arms arose very likely by pericentric inversions from the acrocentric homologues pre­sent in the
basic karyotype of Vespertilionidae (Volleth & Heller 1994). AgNOR-staining detected active
Nucleolus Organizing Regions (NORs) in the distal part of the euchromatic short arms of chro­
mo­so­mes 7, 12 and 15 and proximal to the centromere in minute short arms of chromosomes
8, 9, 14, 18–20, 22 and 24 or 25. These minute short arms are visible only in silver-stained
pre­pa­ra­ti­ons. The differences in the distribution of active NORs in two specimens are shown
in Fig. 3 (35 analyzed cells from specimen 69350 and 20 from spe­ci­men 69349). In many
cells only one ho­mo­lo­gue of a chromosomal pair was shown to bear an active NOR. The X is
a submetacentric chromosome with a heterochromatic segment in the long arm adjacent to the
centromere (see Fig. 2). The subtelocentric Y-chromosome has ap­pro­xi­ma­te­ly the same size as
chromosome 18 and consists largely of heterochromatic ma­te­rial.
Phoniscus atrox Miller, 1905
The karyotype of P. atrox consists of 40 chromosomes and shows a total number of au­to­so­
mal arms of 76. However, quite a large number of autosomal arms consist of heterochromatin
and therefore the fundamental number is presumed to be approximately 50 only. Fig. 4 shows
a comparison of G-banded and C-banded chromosomes of the female studied.
The reduction in the diploid chromosome number from the 2n=44 vespertilionid basic ka­ry­
o­ty­pe to a 2n=40 in Phoniscus atrox is due to two Robertsonian fusion events, i.e. fusion of
chromosomal arms 8+13 and 11+14, respectively. In addition, the karyotype is characterized
by a paracentric inversion in arm 8.
In the P. atrox karyotype, extensive heterochromatin addition was found: first, thin in­ter­s­ti­tial
heterochromatic bands are present in arm 2 (two bands) and in arm 6; second, he­te­ro­chro­ma­tin
276
Fig. 1. G-banded karyotype of Murina suilla.
Obr. 1. Karyotyp trubkonosa vepřího (Murina suilla) barvený G-pruhováním.
277
Fig. 2. C-banded metaphase spread of a Murina suilla male. The arrows point to the euchromatic short
arms of chromosomes 7, 12, 13, and 15.
Obr. 2. Rozptyl C-pruhováním barvené metafase samce Murina suilla. Šipky ukazují euchromatická krátká
ramena chromosomů 7, 12, 13 a 15.
Fig. 3. NOR distribution in two specimens of Murina suilla (SMF catalogue number in parentheses).
Chro­mo­so­mal arms of metacentric chromosomes are marked by x.
Obr. 3. Distribuce oblastí organisujících jadérko (NOR) u dvou jediců Murina suilla (v závorkách jsou
sbírková čísla exemplářů Senckenbergského musea, Frankfurt nad Mohanem, Německo). Chromosomální
ramena metacentrických chromosomů jsou označena x.
278
is found in the short arms of chromosomes 7, 9, 10, 12, 15, 18–25. In the specimen studied,
the size of these heterochromatic arms differs between the homologues of several pairs. This is
especially evident for chromosomes 9 and 18 (Fig. 4). In the heterochromatin blocks, G-positive
segments are followed by G-negative bands. According to the staining pro­per­ties, three dif­f­e­rent
entities can be distinghuished: C-positive, CMA brightly fluorescing, G-negative bands are found
only adjacent to centromeres of chromosomes 9, 10, 12, 18, 19, 22 and 24. Second, G-positive,
C-positive, and DAPI-positive heterochromatin is found in the proximal parts of the short arms
of chromosomes 10, 15, 18, 22 and 24. Applying Distamycin A/DAPI double staining, these
regions show a weak but clearly positive fluorescence, rarely found in other bat species. The
third class is C-negative non-euchromatin, staining either G-positive or G-negative. It is found
in the distal parts of the short arms of chromosome 7, 10, 12, 19–21, 23–25. Silver staining
revealed active NORs in the short arms of chromosomes 7, 12, 18, 20 and 23. In the case of
chromosomes 12, 18 and 23, however, only one homologue each showed an active NOR. These
NOR-bearing chromosomes were clearly distinguishable from the respective homologues by the
presence of a secondary constriction in the short arm. The X chromosome is a medium-sized
submetacentric with a banding pattern similar to that of Myotis Kaup, 1829.
Karyotype comparison of vespertilionid genera have revealed that some of the chro­mo­so­mal
arms exist in two states (I, II) differing by inversions (Volleth & Heller 1994). In Pho­ni­s­cus,
chromosomal arms 11, 13, 15 and the X, all show state I, as in the genus Myotis. The state of
chromosome 7 is difficult to determine. The short arm is clearly different from that of Myotis due
to the terminal addition of two G-positive bands. According to the banding pattern of the long
arm, however, chromosome 7 of Phoniscus Miller, 1905 evolved very likely from state I.
Discussion
As Myotis, the members of the subfamily Kerivoulinae show the highest tooth number found
in Vespertilionidae, i.e. 38. Apart from this primitive character, a wide range of mor­pho­lo­gi­cal
fe­a­tu­res characterizes this subfamily (see Miller 1907, Koopman 1994). There are 23 recognized species in two genera, Kerivoula Gray, 1842 and Phoniscus. According to Miller (1907),
“the greatly increased size and peculiar shape of the upper canine and the four-cusped inner
mandibular incisors distinguish this genus (Phoniscus) sufficiently from Kerivoula”. To­ge­ther
with Hill (1965) and Corbet & Hill (1992) we follow Miller (1907) in treating Pho­ni­s­cus as
a separate genus and not as a subgenus of Kerivoula as Koopman (1994).
The subfamily Murininae, comprising three genera, Murina, Harpiola Thomas, 1915 (Bhat­
tacharyya 2002, Kuo et al. 2006) and Harpiocephalus Gray, 1842, is characterized by tubular
nostrils. It has been regarded by Miller (1907) “as a specialized offshoot from some low,
Myotis-like Ve­sper­ti­li­o­ni­nae form”.
Up to now, only few molecular genetic studies have dealt with phylogenetic relationships
of Murininae and Kerivoulinae. Consistently all studies revealed a sister relationship between
Myotis and a clade containing Murininae and Kerivoulinae (Van Den Bussche & Hoofer 2001,
Hoofer & Van Den Bussche 2003, Stadelman et al. 2004, Kawai et al. 2002). As a result, Hoofer
& Van Den Bussche (2003) proposed subfamiliar rank for Myotini.
Our cytogenetic results revealed one common feature for Myotis, Murina and probably also
Phoniscus, i.e. an euchromatic short arm on chromosome 7 (see Volleth & Heller 1994). From
karyological reasons, it cannot be decided at the moment whether this character is ancestral or
derived. Remarkably, in the Australian genus Vespadelus Troughton, 1943, belonging to the
279
280

Fig. 4. Karyotype of Phoniscus atrox after G-banding (upper rows) and C-banding (lower rows). Note
size polymorphism of heterochromatic short arms.
Obr. 4. Karyotyp vlnouška vroubkozubého (Phoniscus atrox) po G-pruhovém barvení (horní řady) a C-pruhovém barvení (dolní řady). Nápadný je polymorfismus velikosti heterochromatických krátkých ramen.
Vespertilionini, eu­chro­ma­tic short arms are also present on chromosomes 7 and 13 (Volleth &
Tidemann 1989). As in Myotis and Murina, these short arms occurred probably by pe­ri­cent­ric
inversions. In the case of Vespadelus, however, these characters are clearly derived, as closely
related genera lack euchromatic short arms. In the case of Murina and Vespadelus, we are clearly
faced with a re-use of identical or similar inversion break-points on two chro­mo­so­mes, 7 and 13.
It could be sus­pec­ted that these break-points have occurred within re­la­ti­ve­ly short fragile regions,
being hot-spots of rearrangements. Recently, this model has been proposed to explain break-point
clus­te­ring in chromosome evolution by Pevzner & Tesler (2003). The authors concluded that
“mam­ma­li­an genomes are mosaics of fragile regions with high propensity for rearrangements
and solid regions with low propensity for re­ar­ran­ge­ments”. This break-point clustering in fragile
regions reveals limitations of the widely accepted ran­dom breakage theory.
Comparison with published karyological data
From the subfamily Kerivoulinae, only two out of 23 species have been studied ka­ry­o­lo­gi­cal­ly
up to now. The examined female specimen of Kerivoula papillosa showed a diplod chro­mo­so­
me number of 38 and a fundamental number (including the X chromosomes) of 52 (Mc­Bee et
al. 1986). The karyotype of the second species studied, K. lanosa, consists of 28 chromosomes
Table 1. Chromosomal data for Murina. Abbreviations: M – metacentric; SM – submetacentric; ST
– subtelocentric; A – acrocentric; conv – conventional staining; G – G-banding; C – C-banding; AgNOR
– silver-staining of NORs
Tab. 1. Chromosomální údaje o rodu Murina. Zkratky: M – metacentrický; SM – submetacentrický; ST
– subtelocentrický; A – akrocentrický; conv – konvenční barvení; G – G-pruhování; C – C-pruhování;
AgNOR – barvení oblastí organisujících jadérko (NOR) stříbrem
species
2n
aurata1
cyclotis
hilgendorfi2
hilgendorfi
leucogaster
leucogaster
puta
silvatica
suilla
ussuriensis
44 60
44 50?
44
44 56
44 58
44 50
44 50
44 56
44 58
44 56
FN M-SM ST
5
4
4
4
4
4*
4
3
4
4
4
–
–
3
4
–
–
4
4
3
A
X
Y
method
source
12
17
17
14
13
17
17
14
13
14
SM
SM
SM
SM
SM
SM
M
M
SM
SM
A
–
–
A
A
A
A
A
ST
A
conv
conv
conv
G
conv
conv
conv
conv, AgNOR
G, C, AgNOR
G
Ando et al. 1977
Rickart et al. 1999
Harada 1973
Harada et al. 1987
Ando et al. 1977
McBee et al. 1986
Lin et al. 2002
Ono & Obara 1994
this study
Harada et al 1987
* chromosomes showing different morphology compared to the other species.
1
these Japanese specimens would probably be classified with M. ussuriensis today.
2
the subspecies leucogaster hilgendorfi has been elevated to species rank by Yoshiyuki (1989).
281
with a FN of 50 (Rautenbach et al. 1993). The Phoniscus atrox female pre­sen­ted here is the first
Kerivoulinae species where banding methods have been applied. Com­pa­ring the three species
studied, it becomes clear that the karyological variability is much greater in the subfamily Ke­
ri­vou­li­nae than in the Murininae and Myotinae. Chromosomal data from other Kerivoulinae
spe­ci­es are therefore urgently needed for a karyological cha­ra­c­te­ri­za­ti­on of this subfamily.
More chromosomal data are available from the subfamily Murininae. Conventionally sta­i­ned
karyotypes of 6 Murina species have been published (references see Table 1), all with a diploid
chromosome number of 44. Concerning the number of subtelocentric chromosomal pairs, the
data differ, in some cases even within the same species. To my opinion, this is more likely
a matter of interpretation than of real differences, as the minute short arms are clearly visible only
in high quality metaphases. On the other hand, chromosomal differences could probably point
to hidden species. One example could be the M. leucogaster Milne-Edwards, 1872 specimen
of McBee et al. (1986), showing a clearly different morphology of the metacentric pairs. An
indication for cryptic spe­ci­es in the genus Murina was also revealed by cytochrome b se­quen­ce
comparison of Murina cf. cyclotis Dobson, 1872 specimens from Laos (Stadelman et al. 2004).
From the two other Murininae genera, Harpiola and Harpiocephalus, chromosome description
has only be given for a single female specimen of Harpiocephalus mordax Thomas, 1923 with
a 2n=40 (McBee et al. 1986). According to the 5 subtelocentric chromosomal pairs observed
in this specimen, it could be suspected that there are also some pairs with euchromatic short
arms as in Murina.
From the genus Murina, G-banding patterns of two species, Murina ussuriensis and Mu­ri­na
hilgendorfi, have been published previously (Harada et al. 1987). The results are in good agreement with the data published here. Only the euchromatic short arm on chromosome 7, present
in Murina and the genus Myotis, has not been mentioned by Harada et al. (1987).
The distributional pattern of Nucleolus Organizing Regions has only been studied in M. sil­
vatica (Ono & Obara 1994). As in the genus Myotis and in M. suilla, multiple NORs on minute
short arms of acrocentric chromosomes have been found in this species.
As multiple NORs are also present in Phoniscus atrox, the distributional pattern of NORs
is very similar in Myotinae, Kerivoulinae and Murininae. However, the ancestral location of
the NORs for Vespertilionidae is not yet ascertained and therefore this feature is not suited for
judging phylogenetic relationships amongst these subfamilies.
SOUHRN
Poprvé jsou uvedeny popisy chromosomů u trubkonosa vepřího (Murina suilla) (2n=44, FN=58) a vlnouška vroubkozubého (Phoniscus atrox) (2n=40). Karyotyp rodu Murina se liší od karyotypu rodu Myotis
pří­tom­nos­tí euchromatických krátkých ramen na chromosomech 12, 13 a 15. Chromosomová sada druhu
Pho­ni­s­cus atrox je typická extensivním přídavkem heterochromatických krátkých ramen a pa­ra­cen­t­ric­kou
inversí na chromosomu 8. Redukce diploidního počtu chromosomů je způsobena Robertsonskými fusemi
chro­mo­so­mál­ních ramen 8+13 a 11+14.
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