A game against natural selection? Hibernation of migratory bat species in their summering range in Eastern Ukraine
A series of new findings of migratory bat species that in recent years formed wintering groups in the region are described. Bats were found in buildings within human settlements in all cases. Descriptions of new autumn and winter records of Nyctalus noctula in Gorlovka and Rubizhne, Vespertilio muri...
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irk-123456789-1643952020-02-10T01:27:40Z A game against natural selection? Hibernation of migratory bat species in their summering range in Eastern Ukraine Zagorodniuk, I. Ссавці в антропогенному середовищі A series of new findings of migratory bat species that in recent years formed wintering groups in the region are described. Bats were found in buildings within human settlements in all cases. Descriptions of new autumn and winter records of Nyctalus noctula in Gorlovka and Rubizhne, Vespertilio murinus in Luhansk, Alchevsk and Donetsk, and Pipistrellus cf. pygmaeus in Mariupol are presented. These findings are considered as further evidence of formation synanthropic bat communities in eastern Ukraine and expand the geographical boundaries of the known wintering range of these migratory species within Ukraine. Similar cases are rare so far, although they indicate new changes that may soon become regular. Cases of wintering are considered as “trial of forces” of a part of the population in the context of trading-off the “cost of migration” for the “cost of hibernation” under conditions of global warming as well as of improving wintering conditions in the region due to various artificial shelters, mainly in cities. Описано серію нових знахідок видів, що є мігрантами в регіоні, проте останніми роками формують зимівельні групи. В усіх випадках кажанів виявлено в населених пунктах, в житлових приміщеннях. Наведено відомості про нові осінні та зимові знахідки Nyctalus noctula в Горлівці та Рубіжному, Vespertilio murinus в Луганську, Алчевську та Донецьку, Pipistrellus cf. pygmaeus в Маріуполі. Ці знахідки розглядаються як свідчення подальшого процесу формування синантропних угруповань кажанів на сході України і розширюють географічні межі відомого зимового ареалу цих видів-мігрантів в межах України. Поки подібні випадки є поодинокими, проте вони є індикаторами нових змін, які дуже скоро можуть стати закономірними. Такі випадки розглядаються як «проба сил» у частини популяції замінити «ціну міграції» на «ціну зимівлі» в умовах глобального потепління клімату і покращення умов зимівлі на місці за рахунок різноманітних штучних сховищ, зокрема й у містах. 2018 Article A game against natural selection? Hibernation of migratory bat species in their summering range in Eastern Ukraine / I. Zagorodniuk // Theriologia Ukrainica. — 2018. — Т. 16. — С. 111–119. — Бібліогр.: 18 назв. — англ. 2616-7379 DOI: http://doi.org/10.15407/pts2018.16.111 http://dspace.nbuv.gov.ua/handle/123456789/164395 599.4:575.89:574.91(477.52/.6) en Theriologia Ukrainica Національний науково-природничий музей НАН України |
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Ссавці в антропогенному середовищі Ссавці в антропогенному середовищі |
| spellingShingle |
Ссавці в антропогенному середовищі Ссавці в антропогенному середовищі Zagorodniuk, I. A game against natural selection? Hibernation of migratory bat species in their summering range in Eastern Ukraine Theriologia Ukrainica |
| description |
A series of new findings of migratory bat species that in recent years formed wintering groups in the region are described. Bats were found in buildings within human settlements in all cases. Descriptions of new autumn and winter records of Nyctalus noctula in Gorlovka and Rubizhne, Vespertilio murinus in Luhansk, Alchevsk and Donetsk, and Pipistrellus cf. pygmaeus in Mariupol are presented. These findings are considered as further evidence of formation synanthropic bat communities in eastern Ukraine and expand the geographical boundaries of the known wintering range of these migratory species within Ukraine. Similar cases are rare so far, although they indicate new changes that may soon become regular. Cases of wintering are considered as “trial of forces” of a part of the population in the context of trading-off the “cost of migration” for the “cost of hibernation” under conditions of global warming as well as of improving wintering conditions in the region due to various artificial shelters, mainly in cities. |
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Zagorodniuk, I. |
| author_facet |
Zagorodniuk, I. |
| author_sort |
Zagorodniuk, I. |
| title |
A game against natural selection? Hibernation of migratory bat species in their summering range in Eastern Ukraine |
| title_short |
A game against natural selection? Hibernation of migratory bat species in their summering range in Eastern Ukraine |
| title_full |
A game against natural selection? Hibernation of migratory bat species in their summering range in Eastern Ukraine |
| title_fullStr |
A game against natural selection? Hibernation of migratory bat species in their summering range in Eastern Ukraine |
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A game against natural selection? Hibernation of migratory bat species in their summering range in Eastern Ukraine |
| title_sort |
game against natural selection? hibernation of migratory bat species in their summering range in eastern ukraine |
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Національний науково-природничий музей НАН України |
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2018 |
| topic_facet |
Ссавці в антропогенному середовищі |
| url |
http://dspace.nbuv.gov.ua/handle/123456789/164395 |
| citation_txt |
A game against natural selection? Hibernation of migratory bat species in their summering range in Eastern Ukraine / I. Zagorodniuk // Theriologia Ukrainica. — 2018. — Т. 16. — С. 111–119. — Бібліогр.: 18 назв. — англ. |
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Theriologia Ukrainica |
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2025-07-14T16:57:50Z |
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Theriologia Ukrainica, 16: 111–119 (2018)
http://doi.org/10.15407/pts2018.16.111
UDC 599.4:575.89:574.91(477.52/.6)
A GAME AGAINST NATURAL SELECTION? HIBERNATION OF MIGRATORY
BAT SPECIES IN THEIR SUMMERING RANGE IN EASTERN UKRAINE
Igor Zagorodniuk
National Museum of Natural History, NAS of Ukraine (Kyiv, Ukraine)
Correspondence to: I. Zagorodniuk; National Museum of Natural History, NAS of Ukraine;
Bohdan Khmelnytsky St. 15, Kyiv, 01030 Ukraine; e-mail: zoozag@ukr.net
A game against natural selection? Hibernation of migratory bat species in their summering range in
Eastern Ukraine. — I. Zagorodniuk. — A series of new findings of migratory bat species that in recent
years formed wintering groups in the region are described. Bats were found in buildings within human set-
tlements in all cases. Descriptions of new autumn and winter records of Nyctalus noctula in Gorlovka and
Rubizhne, Vespertilio murinus in Luhansk, Alchevsk and Donetsk, and Pipistrellus cf. pygmaeus in Mariupol
are presented. These findings are considered as further evidence of formation synanthropic bat communities
in eastern Ukraine and expand the geographical boundaries of the known wintering range of these migratory
species within Ukraine. Similar cases are rare so far, although they indicate new changes that may soon be-
come regular. Cases of wintering are considered as “trial of forces” of a part of the population in the context
of trading-off the “cost of migration” for the “cost of hibernation” under conditions of global warming as well
as of improving wintering conditions in the region due to various artificial shelters, mainly in cities.
Key words: bats, Vespertilionini, hibernation, Eastern Ukraine, migration, evolution.
Гра проти природного добору? Випадки зимівлі видів-мігрантів кажанів в їхньому ареалі літу-
вання на сході України. — І. Загороднюк. — Описано серію нових знахідок видів, що є мігрантами
в регіоні, проте останніми роками формують зимівельні групи. В усіх випадках кажанів виявлено в
населених пунктах, в житлових приміщеннях. Наведено відомості про нові осінні та зимові знахідки
Nyctalus noctula в Горлівці та Рубіжному, Vespertilio murinus в Луганську, Алчевську та Донецьку,
Pipistrellus cf. pygmaeus в Маріуполі. Ці знахідки розглядаються як свідчення подальшого процесу фо-
рмування синантропних угруповань кажанів на сході України і розширюють географічні межі відомо-
го зимового ареалу цих видів-мігрантів в межах України. Поки подібні випадки є поодинокими, проте
вони є індикаторами нових змін, які дуже скоро можуть стати закономірними. Такі випадки розгляда-
ються як «проба сил» у частини популяції замінити «ціну міграції» на «ціну зимівлі» в умовах глоба-
льного потепління клімату і покращення умов зимівлі на місці за рахунок різноманітних штучних
сховищ, зокрема й у містах.
Ключові слова : кажани, Vespertilionini, зимівля, східна Україна, міграція, еволюція.
Introduction
The current state of the fauna of Ukraine is characterized by significant dynamics of zonal fau-
nal complexes and also by significant changes in geographical ranges and seasonal activity of a
number of species that before had a rather limited presence in space and time. During the last cen-
tury, the territory of Ukraine has changed by 4–26 %, depending on the region, and one of the most
transformed regions is the east of Ukraine (Zagorodniuk, 2006). This region was and still remains an
arena of powerful transformations of nature, which is related to both the strong industrialization in
the first half of the 20th century, and to de-industrialization, which began in the early 21st century
and now reinforced by military actions. A number of synanthropic processes of the eastern fauna
was revealed earlier (Zagorodniuk, 2012).
Among mammals, bats demonstrate distinct tendencies for synanthropy and, respectively, for
noticeable changes in their status, distribution, and abundance (Zagorodniuk, Tkach, 1998). In fact,
Igor Zagorodniuk 112
today bats are one of the most successful mammalian taxonomic groups showing clear synanthropy.
The number records of bats in cities each year reveals distinct positive dynamics (Godlevska, 2012;
Rebrov, 2014).
The present paper considers a group of typical migratory species that are represented in the re-
gion’s fauna by species of the tribe Vespertilionini (Nyctalus, Vespertilio, and Pipistrellus species).
Among them, winter records are known for two species, Nyctalus noctula and Vespertilio murinus,
which are the main subject of this study.
The aim of this article is to summarize the data on findings of migratory species in eastern
Ukraine beyond the breeding period, when normally populations should be located in wintering ar-
eas in other regions.
Material and methods
Data on bat records were amassed during 2005–2014 and used regularly in studies on the com-
position and changes of the fauna (e.g., Zagorodniuk, Korobchenko, 2008; Godlevska, 2012; Re-
brov, 2014; Godlevska, 2015).
The two main data sources in the autumn–winter period were: 1) examination of bat shelters
and, if necessary, extraction of bats for identification and rehabilitation, 2) analysis of residents’
reports on findings and descriptions of bats in buildings and, if possible, photo-documentation of
such records. Both sources were reliable due to the network of students and colleagues of the Faculty
of Natural Sciences Faculty of the University of Luhansk and other educational institutions.
The network remained effective after 2013 (2014–2017) and we continue to receive phone calls
and e-mails asking for advice. Some of these messages are valuable regarding formation on settled
populations of migratory species. All cases are related to the wintering period, which lasts from au-
tumn to early spring seasonal migrations, i.e. from October to late March (see Zagorodniuk, 2002).
Records of Nyctalus noctula
Currently more than 10 records during the non-reproductive period of this species are described
in the region, all related to synanthropic locations, especially the multi-storey residential areas of
large cities (Zagorodniuk, Rebrov, 2014). Earlier described locations (loc. cit.) generally coincided
with the pattern of the summer range of the species in the region. They were confined to the settle-
ments located along the valley of the Siverskyi Donets and its largest tributary, the main corridors
for migratory species (Fig. 1).
New findings are from both the formerly known sites of this species’ records (Siverodonetsk,
Rubizhne, Konstiantynivka, Luhansk) and new locations. During all our research on bats in Luhansk
Oblast (2000–2014), Nyctalus noctula was detected in post-reproductive period only in 7 of 15 years
(2007–2008, 2010–2014) (Zagorodniuk, Rebrov, 2014).
Most of the records (about 10) are from Luhansk, mostly from the central part of the city, and
two other records from Siverodonetsk. In particular, in Siverodonetsk one migration cluster was
discovered in an empty apartment on 11 August 2013 (Zagorodniuk, Rebrov, 2014) and two autumn-
winter findings on the ground near four-storey buildings: 1 specimen on 25 February 2011 and
5 specimens on 10 November 2011 (Godlevska, 2012).
Among the latest records, two important findings of this bat are from Yenakiieve, which is sig-
nificantly distant from other findings, as well as one atypical finding of the species in Rubizhne.
1. Yenakiieve, 02.02.2016 (time of hibernation). The bat was found near a private ground- and
one-storey building, but next to it was the 3-storey building of the hospital, which could serve as
wintering shelter. The discovered specimen is a male, Fa = 50 mm (Fig. 2 a). The animal was found
on the street during the thaw, in daytime, on the asphalt. The bat was weakened hence it was reha-
bilitated (water, calm) and left to hibernate in a cardboard box. After 50 days, on 20.03.2016 the
animal was found dead, mummified (O. Pronin, pers. comm. and photo). This finding is the most
distant from all of the winter records of the species in the region and demonstrates the expansion of
A game against natural selection: cases of hibernation of migratory bat species in their summering range ... 113
its wintering range. In Donetsk Oblast, this is the third described wintering location, after Konstanti-
novka (05.12.2011: Godlevskaya, 2012)1 and Kramatorsk (winter; Zagorodniuk, Rebrov, 2014)2.
2. Rubizhne, 15.08.2016 (end of autumn migration). The bat was found resting in an apartment
on the 3th floor of a four-storey building in Rubizhne, in a collection of cacti completely stuck in the
hooked thorns of Mammillaria bocasana (H. Olievska, photo and pers. comm.). The species identifi-
cation was complicated because of the rather general photo of the animal, but eventually it was iden-
tified as Nyctalus noctula (L. Godlevska, pers. comm.). Among other things, this finding indicates
that synanthropic bats in unnatural habitats face a new danger: the barbed protective covers of alien
plants, including indoor succulents (Fig. 2 b).
3. Pokrovske, before 02.2018 (probably end of autumn migration). The species of the bat was
identified according to its accurately disembowelled skin that was found on the beam in the attic of
the Palace of Culture. The skin was parchment-like and the limbs were dried. Numerous scraps of
pigeons (Columba livia) were found nearby. In the author’s opinion, the bat probably was a victim of
Martes foina. The skin has no traces of moths or skin beetles, which affirms the late autumn or win-
ter appearance of this specimen; this assumption is also supported by the typical resting pose of the
mummy with bended limbs, i.e. the attack might happened in state of deepest hibernation of the bat
(material obtained from A. Rott).
a (▲) b (▼)
Fig. 1. Records of Nyctalus noctula in eastern Ukraine (Zagorod-
niuk, Rebrov, 2014, with additions). Winter records are indicated
with numbers (w16 and w17 are new records from Yenakiieve and
Pokrovske); the shaded area shows the summering range (loc. cit.).
Рис. 1. Знахідки Nyctalus noctula на сході України (за: Загород-
нюк, Ребров, 2014, з доповненнями). Зимові знахідки подано з
номерами (w16 та w17 — нові знахідки в Єнакієвому та По-
кровському); заливка — літній ареал (loc. cit.).
Fig. 2. General view of the noctule bats found
in February 2016 in Yenakiieve (a, by A. Pro-
nin) and in August 2016 in Rubizhne (b, by
I. Oliyevska).
Рис. 2. Загальний вигляд вечірниць, знайде-
них у лютому 2016 р. в Єнакієвому (a, автор
О. Пронін) та в серпні 2016 р. в Рубіжному
(b, автор І. Олієвська).
1 Subsequently, in 2015, this locality was reissued as a new one (Godlevska, 2015).
2 Detailed information was lost: cited article was finished in state of our evacuation from Luhansk occupied by the
Russian invaders. Probably one of the sources was the “Red Book” of Kramatorsk Park (Solomashenko, 2010).
Igor Zagorodniuk 114
Records of Vespertilio murinus
A plenty of new data mainly based on sound signals suggests the presence of this species in the
region during the autumn-winter period. Data continue to be reported by colleagues who also par-
ticipated in the bat census in late autumn in Luhansk in 2005–2014. Particular attention is paid here
to the findings of the bats themselves, which were passed on by locals concerned about these ani-
mals.
Based on the distribution of known records in space and time, V. murinus in the east of Ukraine
seems to be not typical for summer bat communities and appears here mainly during seasonal migra-
tions from more northern regions (Zagorodniuk, Korobchenko, 2008).
Periods of vocalization. Previously, a number of records of this species have been described
based on the registration of lekking males (Zagorodniuk, 2006; Zagorodniuk, Korobchenko, 2008;
Zagorodniuk, Zaika, 2009). Analysis of these data allows suggesting certain regularities. Relatively
long periods of time fall out during the four years of observation (mainly in Luhansk): 1) during
2005, in Luhansk, lekking was recorded on the extreme dates 29.10.2015 and 26.11.2015, i.e. lek-
king lasted a month (4 weeks); 2) during 2006, in Luhansk, the extreme dates were 10.11.2006 and
28.11.2006 (i.e., lekking lasted 3 weeks), 3) during 2006, in Lysychansk, — from 10.11.2006 to
02.12.2006, i.e., lekking lasted not less than 3 weeks; 4) during 2009, in Luhansk, for three dates
with a span of at least 3 consecutive weeks between them: 13.11.2009, 19.11.2009, and 04.12.2009.
Generally, turns out that the lekking period lasts about 3–4 weeks. At the same time, the number of
records in a decade increases gradually from the beginning of the period of the transaction to its end
and then it breaks down. We can assume that the growth of lekking (reproductive) activity, although
stimulated by the first strong cooling and the first thaw, grows until stable colds and, actually, the
cooling stops it.
Actual findings are much rarer than voice records. In addition to the already described findings
(Zagorodniuk, Korobchenko, 2008), we have three new ones:
1. Luhansk, 22.11.2012 (the beginning of hibernation). The species was found in the northern
outskirts of Luhansk (Quarter of the 50th Anniversary of October) at the end of October 2012, i.e. in
the period when autumnal lekking of males occur in this city (Zagorodniuk, Korobchenko, 2008).
A bat was found near a four-storey building, on the 4th floor of which replacement of windows was
in progress (O. Uzun, pers. comm., photo and video of the bat; Fig. 3 b). On the same day, 22 No-
vember 2012, after the photo was taken, the bat escaped and lived in a room in cavities under the
false ceiling showing activity every midnight. Within a few days (until 02.12.12) it was caught and
released. The sex of the animal was not checked, while the length of its forearm (after reconstruction
from the photo) is ca. 45–50 mm. Evidently, the record of V. murinus from Luhansk on 22.11.2012
published without details (Godlevska, 2013) refers to the same specimen that is described here.
2. Alchevsk, 16.02.2014 (strong frosts). The bat was found on the asphalt on 5–7 February
2014, it was warmed up in a box and handed over to rehab “in the state of hibernation”, but actually
it was dead. The specimen was an adult female, FA = 44.9 mm, collection number Z-1002 (working
collection of the author). A series of photos was taken, both of general view and details (tragus,
postcalcar lobe). This is one of the very first records of V. murinus females during late winter in the
east of Ukraine, although there is a late autumn record from Donetsk (31.10.2016).
3. Donetsk, 10.04.2014 (the end of hibernation). A message about the discovery came from a
student hostel; bat was found in a room on 10 April 2014 (Fig. 3 c). The correspondent (student
Zhenia) reported that the bat was found by accident, in a corner with unfinished speeches, however,
most likely, it was not wintering there (it was a rather a dry place), but the animal found a hiding
place there during the recent thaw, in warm spring evenings. Following our advice, the bat was fed
and released. No information about sex and forearm size was recorded.
4. Donetsk, 31.10.2016 (the beginning of hibernation). A bat was found in the late October
2016 in a yard among flats, apparently attacked western jackdaws (Corvus monedula). The animal
was hissing trying to scare the birds; drops of blood were on its wings.
A game against natural selection: cases of hibernation of migratory bat species in their summering range ... 115
a b
c
Fig. 3. General view of Vespertilio murinus discovered in wintering time in synanthropic sites in eastern Ukraine:
a — a bat found on 03.11.2016, Donetsk; b — a bat found in a state of hibernation on 22.11.2012 near a five-storey
building in Luhansk (quarter 50 Years of October); c — a bat found in the economic corner of a dormitory in Donetsk
10.04.2014.
Рис. 3. Загальний вигляд лилика двоколірного (Vespertilio murinus), виявленого в період зимівлі в синантроп-
них місцезнаходженнях сходу України: a — кажан, виявлений 3.11.2016 р. в Донецьку; b — кажан, виявлений
22.11.2012 біля багатоповерхівки в м. Луганськ (кв. 50-річчя Жовтня); c — кажан, виявлений у господарсько-
му кутку студентського гуртожитку 10.04.2014 в м. Донецьк.
Characters of this individual: weight 14 g, FA = 4 cm, female (Fig. 3 a). The bat was given wa-
ter to drink, on the third day fed with zophobas and subsequently kept in captivity for 9 days, until
09.11.2016, when warmer evenings arrived, and then released (pers. com. and photos from O. Pak-
ter). The value of this finding is that it is a female, well fed and ready for wintering; usually in the
autumn only males of the species are found (see also the above finding No. 2). In addition, this is the
first actual record of this species in the winter in Donetsk.
Records of Pipistrellus cf. pygmaeus
The status of pipistrelles in the region is quite specific. Previously, the common pipistrelle and
Nathusius’s pipistrelle were described in this region. However, in fact, only the soprano pipistrelle
(P. pygmaeus) and the Kuhl’s pipistrelle (P. kuhlii) are known to author, and in some locations
P. nathusii as well, one of the rarest bat species in the region3. Both “common” species (kuhlii and
pygmaeus) are alien in the regional fauna and are unknown in natural shelters: P. pygmaeus takes
refuge in a variety of “small architectural forms” (forest houses, cabins, houses, water towers), while
P. kuhlii occurs constantly4 in buildings of different types (kindergartens, multistorey buildings,
various window openings). Survey of records of pipistrelles in synanthropic shelters was published
by S. Rebrov (2012). Thus, only one settled species of the genus Pipistrellus is relatively abundant
in fauna of the region, as well as one migratory species — the soprano pipistrelle (P. pygmaeus), but
it has all signs of an alien species.
Mariupol, 11.12.2012. The soprano pipistrelle (Pipistrellus cf. pygmaeus) was discovered dur-
ing the processing of data on the finding of a wintering group of bats in Mariupol city. During the
dismantling of window blocks on the 2nd floor of the Mariupol Regional Museum, a group of bats
was discovered. The analysis of photographs and descriptions (O. Shakula, pers. com. and photos)
showed that that group consists of about 50 specimens of P. kuhlii. Among them, one specimen was
significantly smaller than P. kuhlii and clearly dark coloured (which is clearly visible on the photo of
3 One record (♀) was in Kreminna near the Klishnia lake (30.06.2001), another one (♀) — in the Rih tract of Prydint-
sivska Floodplain Reserve (05.07.2001) (Petrushenko et al., 2002), and a third one (♂) at the Novo-Illienko Biosta-
tion, 04.07.2009. For comparison, during 10 years of working in eastern Ukraine the author of this paper examined
about 500–700 specimens of P. pygmaeus, 150–250 of P. kuhlii, and only 3 of P. nathusii, but not any P. pipistrellus.
In the NNPM collection, there are several samples of P. pipistrellus, which findings were repeatedly published (e.g.,
Kondratenko, 2001), but subsequently re-identified as P. pygmaeus (including Novo-Illienko).
4 This pipistrelle is not a seasonal migrant, although most of the records are concerned with this very species. One
report differs from the others and it is related to the location where the species had not been recorded earlier hence we
present it with details.
Igor Zagorodniuk 116
the entire group placed into a cardboard box). Apparently, it was a different species: given the possi-
ble set of species known in the region it could be only P. pygmaeus. This finding clearly refers to the
wintering period. The appearance of soprano pipistrelle (and in general the pipistrelle from the group
pipistrellus-pygmaeus-nathusii) in hibernated colony of Kuhl's pipistrelle is a unique fact.
Discussion
Several important aspects of our knowledge on winter bat records may affect the value of spe-
cies for further research, as well as for the development of several scientific hypotheses. In particu-
lar, for the hypothesis of regular attempts of migratory bats to remain in places of summering,
which, under favorable conditions, can lead to the formation of a settled population. Obviously, due
to the respondents, we record only a very small portion of hibernating individuals, probably less than
1 ‰, and there are incomparably more such cases.
1. Is there an expansion of bat ranges or our knowledge?
Until 2000, special studies on bats were not conducted in this region. In fact, research was
started only in the late 1990s by a research group, in which the author also participated. Therefore, it
is possible that new findings only confirm the presence of species that actually existed here for a
long time. However, the reconstruction, including the comparison of descriptions of the fauna of
Luhansk region in the review of I. Sakhno (1963) with modern data (Zagorodniuk, 2012 b), suggests
that certain species, including bats (in particular Eptesicus serotinus s. l. and Pipistrellus kuhlii) have
appeared in the region recently. The same alien status should be recognized also for P. pygmaeus,
which is known only from anthropogenic shelters.
Obviously, facts of formation (or existence) of local wintering ranges in migratory species
should be discussed carefully. It is possible that a part of the population can remain on the ground
year-round. In addition, the migratory status may not be obligatory. However, all cases of wintering
in migrants are associated with synanthropy, although such facts were not discovered and collections
of winter records of these species are absent as well. We can therefore suggest that the absence of
findings evidence the absence of hibernation in cities. Moreover, the number of winter records of
migratory bats has been clearly increased over the past 20 years, as noted in the recent analysis of
winter findings of Nyctalus noctula (Zagorodniuk, Rebrov, 2014). A similar pattern can be seen in
the analysis of the frequency of winter records of migratory bats in Ukraine, as shown for Vespertilio
murinus (Godlevska, 2013) and N. noctula (Godlevska, 2015).
Thus, the process of synantropy and expansion of winter habitats, as well as the increase in fre-
quency of winter finds of bats, which until recently were obligated migrants (or they were distributed
beyond the region), occur objectively in eastern Ukraine. Increase of records occurs especially in
large and medium-sized cities with multi-storey buildings and available warm shelters (balconies,
windowsills, elevator shafts, utility rooms). At the same time, many technological premises in indus-
trial facilities remain unexplored.
2. What is growing: the number of bats or the frequency of registrations?
Obviously, the significant distribution of knowledge about bats and the needs of their protec-
tion, the creation of web pages and booklets, organization of such events as “Year of bats” and
“Bat’s Night”, as well as publications and discussions in the mass media (in which the author ac-
tively participated) significantly affected the growing of interest to these animals and the emergence
of a series of new important messages about findings of bats from amateurs to specialists. Probably
the proliferation of the Internet and mobile communications also contributed. However, bats did not
become more numerous in the region, and their populations were mainly reduced due to destruction
of natural shelters, changes in the food base, etc. Comparison of data from districts that differ in
urbanization levels and data from respondents of different age and social groups suggests that the
frequency of migrant bat records (i.e., the proportion of actual populations) in synanthropic habitats
clearly increases. Moreover, both elderly and younger people are surprised by the new facts of such
more and more frequent registrations.
A game against natural selection: cases of hibernation of migratory bat species in their summering range ... 117
Thus, it can be assumed (a general impression, without statistics) that the increase in the fre-
quency of registration of bats in synanthropic locations is an objective process caused by the specif-
ics of both the development of urban landscape and synanthropic faunas (Zagorodniuk, 2012 a). In
addition, such changes and their effects on the fauna are definitely more powerful and significant
than the processes described in the framework of the concepts of global warming, although it is ob-
vious that warming also has a significant impact.
3. Can trends change due to current deindustrialization?
Military actions that have been taking place in the east of Ukraine since the spring of 2014
caused significant changes not only in nature management (reduction of anthropogenic pressure on
natural complexes, absence of hunting), but also the structure of natural complexes (violation of soil
and vegetation cover, destruction of woodlands, changes of hydroregime). The largest current
changes occur in synanthropic locations, which is due to the termination of a significant number of
industrial objects and often their dismantling, an increase in the portion of objects that lost their eco-
nomic value and maintenance (including cleaning and heating). All these processes lead to essen-
tially unpredictable changes, although along with the increase of shelters available for bats. Build-
ings are attractive as places for rest and reproduction, with minimal anthropogenic pressure and
small influence of predators. In addition, human buildings have an attractive temperature regime.
Therefore, deindustrialization may be considered as a positive factor for further synanthropiza-
tion of populations, of essentially synurbic (i.e., susceptible to artificial urban landscapes and micro-
climate) animals, including bats. On the other hand, obtaining new data on the fauna of the region
since 2014 has been largely limited by the impossibility of conducting field research and the gradual
loss of contacts with residents and of residents’ interest to the objects of nature.
All these three factors can contribute to the development of synanthropic bat communities.
About the growth of the number of registrations
The increase in the number of bats’ registrations in cities that has taken place in recent years is
probably due to the growing interest in research and active distribution of knowledge about bats
among townspeople with requests to report bat records. The activity of “contact centres on bats”, one
of which — the “Donbas”, led by the author of this publication (along with M. Korobchenko,
M. Kolesnikov and S. Rebrov) — was precisely related to the identification of such cases and distri-
bution of knowledge about the needs of the protection of these animals. However, it was not only the
growth of research interest: numerous facts of meetings with bats in buildings also indicate the
growing interest of bats to dwellings. Changes in ecosystems and forms of nature taking place in
recent years suggest a further increase in cases of occurrence of bats in artificial shelters. Nonethe-
less, the interest of people to bats from people in eastern Ukraine has clearly faded in the recent
years, and during 2014–2017 the author received from the east only 20–30 phone calls per year.
This fact explains the decrease in the media coverage of the topic: if in 2005–2014 there were at
least 5–10 different reports on the local (oblast) television and somewhat more in the press, but now
data can be obtained only after an extensive search for information on the Internet, as well as by
preservation of a unique network of respondents and constantly reminding them about bats and other
animals. This approach was also used in data collection for this article.
Is there something Darwinian involved?
Bats are mammals, and therefore, unlike other groups that also form migratory populations, they
have a more complex and plastic behaviour. Bats seem to be able to change their migratory status
faster than other animals. The numerous artificial and very diverse microclimatic conditions, the
degree of comfort in shelters, as well as the general warming of cities are important factors in chang-
ing the migratory status. It is worth remembering that, in addition to global changes, there are proc-
esses of energetic warming of cities. All this creates the preconditions for the appearance of seasonal
synurbic animal population.
Synurbization contributes to the change of migratory behaviour and appearance of stationary
groups further affected by selective pressure upon at least the following 6 positions:
Igor Zagorodniuk 118
1) bats that successfully wintered under synanthropic conditions are the first that occupy the
most suitable summer shelters, while migrants are still passing through migratory routes;
2) such bats do not waste large amount of energy for migration and do not face the risks of
death due to predators or extreme climatic conditions on long migratory routes;
3) since individuals of both sexes of the last year offspring can stay for wintering, bisexual win-
tering groups can be formed and subsequently exist as summer bisexual colonies, unlike typical ma-
ternity ones, which are usually present (Nehoda, 2002) at our latitudes;
4) the next year offspring may not migrate to the traditionally recognized wintering range and
stay with the already settled groups becoming a new generation of a non-migrating colony;
5) subsequently, subpopulations can be formed with some level of genetic isolation between
migratory and settled colonies;
6) since migratory status is a species trait, especially in bats of temperate latitudes, such rapid
and large-scale changes on the level of population groups (and not separate individuals) can be pre-
conditions for more significant microevolutionary events even up to possible speciation.
Today, we have a large number of examples of synanthropy among bats, often seasonal, and es-
pecially winter. Many species also demonstrate their expansion into new geographical regions, espe-
cially through the network of synanthropic locations (the common and Khazar serotines, soprano and
Kuhl’s pipistrelles, parti-coloured bat and grey long-eared bat). The winter finds of migratory bats
described here are a logical manifestation of attempts of migratory species to wintering within their
summering range, without the cost of migration to long distances and without the risks of death dur-
ing migration. Obviously, such attempts of colonies to remain in summering places are a normal
reaction to changes in the macroclimate of cities and in the set of available shelters.
Many researchers, including the author of this publication, when first face similar facts of win-
tering, they consider them as occasional events ending with the death of individuals, i.e. a deviation
from the norm immediately eliminated by natural selection. However, research results presented in
this paper allows suggesting that these processes can involve an essential part of the population and
lead to microevolutionary processes eventually resulting in changes of both of the species and com-
munities. Therefore, this “game” seems to be not against but controlled by selection.
Acknowledgement
I thank to the respondents for valuable information about findings of bats, in particular I. and
H. Oliyevska (Rubizhne), O. Uzun (Luhansk), O. Pronin (Yenakiieve), O. Packter (Donetsk),
O. Shakula (Mariupol), A. Rott (Pokrovske). My thanks to M. Kolesnikov and S. Rebrov (Luhansk
University) for their ongoing support in the examination of locations, trips to places of detection of
bats and participate in creating a database of bats records in the Luhansk region.
My thanks to V. Parkhomenko for editing the photographs and L. Godlevska for helping to
identify of some samples. Thanks to A.-T. Bashta for useful comments, and Z. Barkaszi and
D. Vishnevsky for the discussion of ideas.
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