Leaping on urban islands: further summer and winter range expansion of European bat species


  • Anton Vlaschenko a:1:{s:5:"en_US";s:35:"Ukrainian Bat Rehabilitation Center";}
  • Vitalii Hukov
  • Olha Timofieieva
  • Marharyta Moiseienko
  • Anastasia Domanska
  • Oleksandr Zinenko
  • Alona Prylutska




bats, Hypsugo savii, Plecotus austriacus, Pipistrellus nathusii, Pipistrellus pygmaeus, urbanization, range expansion


Cities or urban areas are the new types of landscapes that have rapidly developed in the Anthropocene and generally mimic mountains and rock habitats. Such areas attract different vertebrate species that naturally prefer rocky habitats, for example, bats, which are common animal inhabitants of the cities in the Northern Hemisphere. Here we review records of four bat species (Hypsugo savii, Plecotus austriacus, Pipistrellus nathusii and P. pygmaeus) inhabiting human settlements in Ukraine, encompassing the period from 2011 to 2022. Over the last 20 to 30 years, the winter range of P. nathusii has shifted 200-300 km north, and now covers all Black Sea coast steppe regions of continental Ukraine. The Pl. austiacus range most likely covers the whole territory of Ukraine. We documented the first factual records of H. savii in continental Ukraine and the first winter records of P. pygmaeus for the country. Our observations clearly demonstrate colonization of newly formed urban landscapes by bats species from different ecological groups. Therefore, bats, same as some other mammalian species, can be considered beneficiaries of urbanization and urban heat islands. 


Abelentsev, V.I. & Pidoplichko, I.G. (1956) The Order of Insectivores - Insectivora. Fauna of Ukraine. Volume 1: Mammals. Number 1. The Academy of Sciences of URSR Press, Kyiv. [in Ukrainian].

Ancillotto, L., Tomassini, A. & Russo, D. (2015) The fancy city life: Kuhl’s pipistrelle, Pipistrellus kuhlii, benefits from urbanisation. Wildlife Research. 42, 598-606. DOI: https://doi.org/10.1071/WR15003

Alcalde, J.T., Jiménez, M., Brila, I., Vintulis, V., Voigt, C.C., Petersons, G. (2020) Transcontinental 2200 km migration of a Nathusius’ pipistrelle (Pipistrellus nathusii) across Europe. Mammalia. 85, 161–163. DOI: https://doi.org/10.1515/mammalia-2020-0019

Barlow, K.E. & Jones, G. (1999) Roosts, echolocation calls and wing morphology of two phonic types of Pipistrellus pipistrellus. Zeitschrift fur Sdugetierkunde. 64, 257–268.

Bartonicka, T. Benda, P. & Juda, J. (2017) First findings and phenology of Hypsugo savii in the Děčín District, Czech Republic (Chiroptera: Vespertilionidae). Lynx, n. s. (Praha). 48, 5–14. [in Check with English summary]. DOI: https://doi.org/10.2478/lynx-2017-0001

Bashta, A.-T. (2010) Characteristics of bat community of the city of Lviv (Ukraine): species composition, spatial and seasonal distribution. Studia Biologica. 3, 109–124. [in Ukrainian with English summary].

Bilushenko, A.A. (2009) The first record of the grey long-eared bat, Plecotus austriacus (Chiroptera, Vespertilionidae), in Cherkasy region. Vestnik Zoologii 43, 120. [In Russian]

Bilushenko, A.A. (2013) The current status of Kuhl’s Pipistrelle, Pipistrellus kuhlii (Chiroptera, Vespertilionidae), in the central forest-steppe of Ukraine. Vestnik Zoologii. 47, 343–349. DOI: https://doi.org/10.2478/vzoo-2013-0036

Blomberg, A.S., Vasko, V., Salonen, S., Pētersons, G., Lilley, T.M. (2020) First record of a Nathusius’ pipistrelle (Pipistrellus nathusii) overwintering at a latitude above 60°N. Mammalia. 85, 74-78. DOI: https://doi.org/10.1515/mammalia-2020-0019

Bogdarina, S. & Strelkov, P. (2003) Distribution of bats (Chiroptera) in the north of European Russia. Plecotus et al. 6, 7–28. [In Russian with English summary].

Buchko, V. Vlaschenko, A., Kravchenko, K., Sudakova, M., Gukasova, A., Kusnezh, O. (2011) Contribution to bat fauna (Chiroptera) of Galytskiy National Nature Park (Ivano-Frankivsk region). Visnuk of Lviv University. Series: biology. 55, 146-159. [In Ukrainian with English summary].

Clergeau, P., Savard, J.P.L., Mennechez, G., Falardeau, G. (1998) Bird abundance and diversity along an urbanrural gradient: A comparative study between two cities on different continents. Condor. 100, 413-425. DOI: https://doi.org/10.2307/1369707

Dietz, C. & Kiefer, A. (2014). Die Fledermäuse Europas: kennen, bestimmen, schützen. Kosmos, Stuttgart.

Dombrovski, V., Fenchuk, V. & Zhurauliou, D. (2017) The bat fauna (Mammalia, Chiroptera) of Stary Zhaden zakaznik, southern Belarus. Proceedings of the Theriological School. 15, 3–9. DOI: https://doi.org/10.15407/ptt2017.15.003

Gashchak, S.P., Vlaschenko, A.S., Naglov, A.V., Kravchenko, K.A., Prylutska, A.S (2013) Bat fauna of the Exclusion Zone in concern of assessment of environmental value of its areas. Problems of Chernobyl Exclusion Zone. 11, 56–78. [in Russian with English summary].

Godlevska, L. (2012a) Expansion of Plecotus austriacus (Chiroptera, Vespertilionidae) Range: First Records in the Kyiv Oblast (Ukraine). Vestnik zoologii. 46, 88. DOI: https://doi.org/10.15407/pts2018.16.051

Godlevska, L. (2012b) Results of the work of the bat contact-centre (Ukraine). Scientific Notes of Taurida V.I. Vernadsky National University. Series: Biology, chemistry. 25, 12-20. [in Russian].

Godlevska, L.V. (2013) New Vespertilio murinus (Chiroptera) Winter Records. An Indication of Expansion of the Species’ Winter Range? Vestnik Zoologii. 47, 239-244. DOI: https://doi.org/10.2478/vzoo-2013-0023

Godlevska, L. (2014) Northward expansion of the winter range of Nyctalus noctula (Chiroptera: Vespertilionidae) in Eastern Europe. Mammalia. 79, 315-324. DOI: https://doi.org /10.1515/mammalia-2013-0178

Godlevska, L. (2015) Results of the Work of the Kyiv Bat Contact-Centre in 2012–2015. Proceedings of the Theriological School. 13, 11–19. [in Ukrainian] DOI: https://doi.org/10.15407/ptt2015.13.011

Godlevska, L. & Shpak, A. (2020) Bats in the collection of the Zoological museum of Belarusian State University. Proceedings of the National Academy of Sciences of Belarus. Biological series. 65, 412–420. [in Russian with English abstract]. DOI: https://doi.org/10.29235/1029-8940-2020-65-4-412-420

Godlevskaya, E.V., Ghazali, M.A. & Postawa, T. (2009) A Current State of Cave Dwelling Bat Species (Mammalia, Chiroptera) of the Crimea. Vestnik zoologii. 43, 253–265. [In Russian with English summary].

Godlevska, L., Ghazali, M. & Tyshchenko, V. (2012) Results of the census of cave-dwelling bats in Podolia and Middle Dniester River region (Ukraine) in 2010–2011. Proceedings of the Theriological School. 11, 89–97. [In Ukrainian]. DOI: https://doi.org/10.15407/ptt2012.11.089

Godlevska, L., Buzunko, P., Rebrov, S., Ghazali, M. (2016a) Underground bat sites of “not-cave” region of Ukraine, on results of 2002–2015. Visnyk of the Lviv University. Series Biology. 71, 178–189. [In Ukrainian].

Godlevska, L.V., Rebrov, S.V. & Panchenko, P.S. (2016b) New data on bat fauna of Rivne oblast (Ukraine). Zapovidna sprava. 22, 72–77. [In Ukrainian].

Godlevska, L. & Rebrov, S. (2018) Bats of the Left-Bank Dnipro Region in the northern part of Ukraine. Theriologia Ukrainica. 16, 25–50. [In Ukrainian]. DOI: https://doi.org/10.15407/pts2018.16.025

Godlevska, L., Panchenko, P., Rebrov, S., Savchenko, M., Yakovlev, M., Formanyuk, O., Ghazali, M. (2018) Knowledge of the bats on the territory of the Black Sea coast of Ukraine (AR Krim, Mykolayivska, Odeska, Kherson region) In: Materials up to the 4th edition of the Red-data book of Ukraine. Volume. 1. (pp. 196–203). Kyiv: Schmalhausen Institute of Zoology of National Academy of Sciences of Ukraine. [In Ukrainian].

Grimm, N.B., Foster, D., Groffman, P., Grove, J.M., Hopkinson, C.S., Nadelhoffer, K.J., et al. (2008) The changing landscape: ecosystem responses to urbanization and pollution across climatic and societal gradients. Frontiers in Ecology and the Environment. 6, 264-272. DOI: https://doi.org/10.1890/070147

Gukasova, A., & Vlaschenko, A. (2011) Effectiveness of mist-netting of bats (Chiroptera, Mammalia) during the non-hibernation period in oak forests of Eastern Ukraine. Acta Zoologica Cracoviensia-Series A: Vertebrata. 54, 77-93. DOI: https://doi.org/10.3409/azc.54a_1-2.77-93

Evans, K.L., Chamberlain, D.E., Hatchwell, B.J., Gregory, R.D., Gaston, K.J. (2011) What makes an urban bird? Global Change Biology. 17, 32-44. DOI: https://doi.org/10.1111/j.1365-2486.2010.02247.x

Hukov, V., Timofieieva, O., Prylutska, A., Rodenko, O., Moiseienko, M. Bohodist, V., et al. (2020) Wintering of an urban bat (Pipistrellus kuhlii lepidus) in recently occupied areas. European Journal of Ecology. 6, 102-120. DOI: https://doi.org/10.17161/eurojecol.v6i1.13629

Hutterer, R., Ivanova, T., Meyer-Cords, C., Rodrigues, L. (2005). Bat migrations in Europe: a review of banding data and literature. Bonn: Federal Agency for Nature Conservation.

Jung, K., & Threlfall, C.G. (2016) Urbanization and Its Effects on Bats - A Global Meta-Analysis. In: Voigt, C.C. and Kingston, T. (Eds.). Bats in the Anthropocene: Conservation of Bats in a Changing World (pp. 13-33). Springer, Cham. DOI: https://doi.org/10.1007/978-3-319-25220-9_2

Juste, J. & Paunović, M. (2016) Hypsugo savii. The IUCN Red List of Threatened Species 2016: e.T44856A22072380. Available at: DOI: https://dx.doi.org/10.2305/IUCN.UK.2016-3.RLTS.T44856A22072380.en. (Accessed on 22 January 2022).

Kovalov, V., Hukov, V. & Rodenko, O. (2019) New record of Nyctalus lasiopterus (Schreber, 1780) in Ukraine with a new confirmation of carnivory. North-Western Journal of Zoology. 15, 91-95.

Krauel, J.J. & LeBuhn, G. (2016) Patterns of Bat Distribution and Foraging Activity in a Highly Urbanized Temperate Environment. PLoS ONE. 11, e0168927. DOI: https://doi.org/10.1371/journal.pone.0168927

Kravchenko, K., Vlaschenko, A., Prylutska, A., Rodenko, O., Hukov, V., Shuvaev, V. (2017) Year-round monitoring of bat records in an urban area: Kharkiv (NE Ukraine), 2013, as a case study. Turkish Journal of Zoology 41, 530-548. DOI: https://doi.org/10.3906/zoo-1602-51

Kravchenko, K.A., Vlaschenko, A.S., Lehnert, L.S., Courtiol, A., Voigt, C.C. (2020) Generational shift in the migratory common noctule bat: first-year males lead the way to hibernacula at higher latitudes. Biology Letters. 16, 20200351. DOI: https://doi.org/10.1098/rsbl.2020.0351

Kruskop, S. V., Borisenko, A. V., Ivanova, N. V., Lim, B. K., Eger, J. L. (2007) Using of DNA barcodes to recover phylogeographical splits among East Palaearctic bats. Molecular and genetic basis for preservation of the Holarctic mammal diversity (pp. 115-121). Materials of the conference. Moscow: KMK Scientific Press.

Manyuk, V. & Lahuta, A. (2018) Plecotus austriacus in the valley of the Kilchen river (left bank of the Dnipro), a new stage of expansion to the East. Theriologia Ukrainica. 16, 149–151. [in Ukrainian]. DOI: https://doi.org/10.15407/pts2018.16.149

McKinney, M.L. (2008) Effects of urbanization on species richness: A review of plants and animals. Urban Ecosystems. 11, 161-176. DOI: https://doi.org/10.1007/s11252-007-0045-4

Mas, M., Flaquer, C., Puig-Montserrat, X., Porres, X., Rebelo, H., López-Baucells, A. (2022) Winter bat activity: The role of wetlands as food and drinking reservoirs under climate change. The Science of the total environment. 828, 154403. DOI: https://doi.org/10.1016/j.scitotenv.2022.154403

Maxinová, E., Kipson, M., Naďo, L., Hradická, P., Uhrin, M. (2016) Foraging Strategy of Kuhl’s Pipistrelle at the Northern Edge of the Species Distribution. Acta Chiropterologica. 18, 215–222. DOI: https://doi.org/10.3161/15081109ACC2016.18.1.012

Panchenko, P., & Godlevska, L. (2018) Data on the bat fauna of the Northern Black Sea Region based on results of the work of bat contact centres. Theriologia Ukrainica. 16, 120-126. [in Ukrainian]. DOI: https://doi.org/10.15407/pts2018.16.120

Paunović, M. & Juste, J. (2016) Pipistrellus nathusii. The IUCN Red List of Threatened Species 2016: e.T17316A22132621. Available at: DOI: https://doi.org/10.2305/IUCN.UK.2016-2.RLTS.T17316A22132621.en. (Accessed on 22 January 2022).

Prylutska, A.S. & Vlaschenko, A.S. (2013) Materials on bat distribution on the base of results of contact-center in Kharkiv (2008–2012). Biological systems. 5, 532-537.

Prylutska, A., Moiseienko, M., Yerofieieva, M., Hukov, V., Vlaschenko, A. (2020) Northern record for Tadarida teniotis (NE Ukraine) far from known species range. Journal of Bat Research & Conservation. 13, 104-108. DOI: https://doi.org/10.14709/BarbJ.13.1.2020.17

Rakhmatulina, I.K. (2005). Bats of Azerbaijan (fauna, ecology, zoogeography). Baku: Institute of Zoology of NAS Azerbaijan.

Robins, J. H., Hingston, M., Matisoo‐Smith, E. Ross, H. A. (2007) Identifying Rattus species using mitochondrial DNA. Molecular Ecology Notes. 7, 717-729. DOI: https://doi.org/10.1111/j.1471-8286.2007.01752.x

Rodenko, E.E., Kravchenko, K.A., Vlaschenko, A.S., Chepizhnaya, A.J., Suvorova, A.D., Prylutskaya A.S. (2014) The record of Nyctalus noctula in winter time in Poltava city (Ukraine). Plecotus et al. 17, 43–51. [in Russian with an abstract in English]

Russo, D. & Ancilotto, L. (2015) Sensitivity of bats to urbanization: a review. Mammalian Biology. 80, 205–212. DOI: https://doi.org/10.1016/j.mambio.2014.10.003

Russo, D., Salinas-Ramos, V.B., Cistrone, L., Smeraldo, S., Bosso, L., Ancillotto, L. (2021) Do We Need to Use Bats as Bioindicators? Biology. 10, 693. DOI: https://doi.org/10.3390/biology10080693

Sachanowicz, K., Wower, A. & Bashta, A. (2009) Further range extension of Pipistrellus kuhlii (Kuhl, 1817) in Central and Eastern Europe. Acta Chiropterologica. 8, 543–548. DOI: https://doi.org/10.3161/1733-5329(2006)8[543:FREOPK]2.0.CO;2

Sachanowicz, K., Piskorski, M. & Tereba, A. (2017) Systematics and taxonomy of Pipistrellus kuhlii (Kuhl, 1817) in Central Europe and the Balkans. Zootaxa. 4306, 053–066. DOI: https://doi.org/10.11646/zootaxa.4306.1.2

Sachanowicz, K., Ciechanowski, M., Tryjanowski, P., Kosicki, J.Z. (2018) Wintering range of Pipistrellus nathusii (Chiroptera) in Central Europe: has the species extended to the north-east using urban heat islands? Mammalia. 83, 260-271. DOI: https://doi.org/10.1515/mammalia-2018-0014

Schneider, A., Friedl, M.A. & Potere, D. (2010) Mapping global urban areas using MODIS 500-m data: new methods and datasets based on ‘urban ecoregions. Remote Sensing of Environment. 114, 1733–1746. DOI: https://doi.org/10.1016/j.rse.2010.03.003

Shpak, A. (2018) Hibernating bat species of Belarus: results of the work of the Minsk bat contact centre. Theriologia Ukrainica. 16, 156–162. DOI: https://doi.org/10.15407/pts2018.16.156

Steffens, R., Zöphel, U., Brockmann, D. (2004). 40th Anniversary Bat Marking Centre Dresden — Evaluation of Methods and Overview of Results. Dresden: Saxon State Office for Environment and Geology.

Strelkov, P.P. (1969) Migratory and stationary bats (Chiroptera) of the European part of the Soviet Union. Acta Zoologica Cracoviensia. 14, 393–439.

Strelkov, P.P. (1999) Sex ratio in breeding season in adult individuals of long-distant migrant bats (Chiroptera, Vespertilionidae) of Eastern Europe and adjacent territories. Zoologicheskiy zhurnal. 78, 1441–1454. [in Russian with English summary].

Strelkov, P.P. (2000) Seasonal distribution of migratory bat species (Chiroptera, Vespertilionidae) in eastern Europe and adjacent territories: nursing area. Myotis. 37, 7–25.

Strelkov, P.P. (2002) Materials on wintering of migratory bat species (Chiroptera) on the territory of the former USSR and adjacent regions. Part. 2. Nyctalus noctula. Plecotus et al. 5, 35–56. [In Russian].

Strelkov, P.P. & Il’in, V. Yu. (1990) Chiropterans (Chiroptera, Vespertilionidae) of the Southern Middle and Low Volga Regions. Proceedings of the Zoological Institute of the SSSR. 225, 42–167. [In Russian].

Strelkov, P.P., Unkurova, V.I. & Medvedeva, G.A. (1985) New data on Pipistrellus kuhlii and dynamics of its range in the USSR. Zoologichesky zhurnal. 64, 87-97.

Sumasgutner, P., Nemeth, E., Tebb, G., Krenn, H.W., Gamauf, A. (2014) Hard times in the city attractive nest sites but insufficient food supply lead to low reproduction rates in a bird of prey. Frontiers of Zoology. 11, 13. DOI: https://doi.org/10.1186/1742-9994-11-48

Uhrin, M., Hüttmeir, U., Kipson, M., Estók, P., Sachanowicz K., Bücs, S.L., et al. (2016) Status of Savi's pipistrelle Hypsugo savii (Chiroptera) and range expansion in Central and south-eastern Europe: a review. Mammal Review. 46, 1-16. DOI: https://doi.org/10.1111/mam.12050

United Nations, Department of Economic and Social Affairs, Population Division (2014) World urbanization prospects: the 2014 revision, highlights (ST/ESA/SER.A/352).

Vlaschenko, A.S. (2011) Research history and list of records of bats (Chiroptera) in the Kharkov Region in the XIX and XX centuries. Plecotus et al. 14, 26-54. [in Russian].

Vlaschenko, A.S. & Gukasova, A. S. (2010) Bat (Chiroptera) fauna and structure of assemblage of projecting protected area “Yaremovskoe” (Kharkov region). Nature reserves in Ukraine. 16, 44-50. [In Russian].

Vlaschenko, A.S. & Plylutska, A.S. (2018) Das Bat Rehabilitation Center des „Feldman Ecoparks“, Charkiw, Ukraine. Nyctalus (N.F.) Berlin. 19, 158–161. [in German].

Vlaschenko, A., Kravchenko, K., Prylutska, A., Ivancheva, E., Sitnikova, E., Mishin, A. (2016) Structure of summer bat assemblages in forests in European Russia. Turkish Journal of Zoology. 40, 876-893. DOI: https://doi.org/10.3906/zoo-1508-56

Vlaschenko, A., Kovalov, V., Hukov, V., Kravchenko, K., Rodenko, O. (2019) An example of ecological traps for bats in the urban environment. European Journal of Wildlife Research. 65, 20. DOI: https://doi.org/10.1007/s10344-019-1252-z

Vlaschenko, A.S., Prylutska, A.S., Kravchenko, K., Rodenko, O., Hukov, V., Timofieieva, O., et al. (2020) Regional recaptures of bats (Chiroptera, Vespertilionidae) ringed in Eastern Ukraine. Zoodiversity. 54, 53–66. DOI: https://doi.org/10.15407/zoo2020.01.053

Vlaschenko, A., Yatsiuk, Y., Hukov, V., Prylutska, A., Straka, T. M., Kravchenko, K. (2021) Urban forest preserves local bat species diversity, but not forest dweller specialists—renewed study 65 years later (Kharkiv city, Ukraine). Mammal Research. 66, 615-626. DOI: https://doi.org/10.1007/s13364-021-00580-9

Volokh, A., Gorlov, P., Siokhin, V., Polishchuk, I. (2021) Species diversity of bats (Chiroptera) in the Ukrainian Azov Region and features of their residence by seasons. Theriologia Ukrainica. 20, 24–36. [In Ukrainian with English summary]. DOI: https://doi.org/10.15407/TU2104

Wilmer, W.J., & Barratt, E. (1996) A non-lethal method of tissue sampling for genetic studies of chiropterans. Bat Research News. 37, 1–3.

Zagorodniuk, I. (2018) A game against natural selection: cases of hibernation of migrant bat species in their summering range in Eastern Ukraine. Theriologia Ukrainica. 16, 111-119. DOI: https://doi.org/10.15407/pts2018.16.111

Zagorodniuk, I. (2019) Range dynamics in sibling species: facts and reconstructions for the mammal fauna of Eastern Europe. Theriologia Ukrainica. 18, 20–39. DOI: https://doi.org/10.15407/pts2019.18.020

Zahn, A., & Kriner, E. (2014). Winter foraging activity of Central European Vespertilionid bats. Mammalian Biology. 81, 40–45. DOI: https://doi.org/10.1016/j.mambio.2014.10.005

Zuñiga-Palacios, J., Zuria, I., Castellanos, I., Lara, C., Sánchez-Rojas, G. (2021) What do we know (and need to know) about the role of urban habitats as ecological traps? Systematic review and meta-analysis. The Science of the total environment. 780, 146559. DOI: https://doi.org/10.1016/j.scitotenv.2021.146559

Zsebok, S., Estók, P. & Görföl, T. (2012) Acoustic discrimination of Pipistrellus kuhlii and Pipistrellus nathusii and its application to assess changes in species distribution. Acta Zoologica Academiae Scientiarum Hungaricae. 58, 199–209.




How to Cite

Vlaschenko, A., Hukov, V. ., Timofieieva, O. ., Moiseienko, M. ., Domanska, A. ., Zinenko, O. ., & Prylutska, A. . (2023). Leaping on urban islands: further summer and winter range expansion of European bat species. European Journal of Ecology, 9(1). https://doi.org/10.17161/eurojecol.v9i1.18664