05_Godlevska-1.indd UDC 599.4(477) BATS OF CENTRAL UKRAINE: A SYNOPSIS L. Godlevska1*, S. Rebrov2, P. Vorobei1, M. Savchenko1, P. Panchenko2 1Schmalhausen Institute of Zoology NAS of Ukraine, vul. B. Khmelnytskogo, 15, Kyiv, 01030 Ukraine 2Ukrainian Centre for Bat protection *Corrersponding author E-mail: lgodlevska@izan.kiev.ua L. Godlevska (https://orcid.org/0000-0001-6792-6543) S. Rebrov (https://orcid.org/0000-0002-6780-5386) P. Vorobei (https://orcid.org/0000-0002-8230-6374) M. Savchenko (https://orcid.org/0000-0001-5929-5024) P. Panchenko (https://orcid.org/0000-0003-3602-0831) Bats of Central Ukraine: a Synopsis. Godlevska, L., Rebrov, S., Vorobei, P., Savchenko, M., Panchenko, P. — Here, we represent the fi rst review of the bat fauna of the large territory of Central Ukraine (to the west from Dnipro River). Th e review is based on results of the original survey in 1999– 2021 and data from all available sources (publications, museum collections). Fauna of the region includes 24 bat species : Eptesicus, 2 species; Barbastella, 1; Myotis, 10; Nyctalus, 3; Pipistrellus, 4; Plecotus, 2; Vespertilio, 1; Rhinolophus, 1. For each species there is a short description of its distribution (illustrated by a map), used roosts, seasonality of presence and reproductive status within the study area. Th e bat fauna composition is analysed by three distinguished subregions, from north to south: of mixed forests, the forest-steppe, and the Dniester River. Th e estimation of prevalence and comparative abundance of the species in the region is given. Th e general review of bat roosts, by their types (under- and overground), seasonality of usage, and sheltered species is presented. Th e paper is supplemented with the extensive array of original, earlier unpublished, primary data on bat records in the region. K e y w o r d s : bats, distribution, seasonality, reproductive status, abundance, roosts, Central Ukraine. Introduction All 28 bat species of Ukrainian fauna are legally protected: by the national laws of Ukraine and international treaties (EUROBATS Agreement, CMS, Bern Convention). Vulnerability of bats, tasks of their protection and conservation determine a critical need for clarifying their distribution, abundance and ecological preferences, both at national and regional levels (Fenton, 1997; Frick et al., 2020). Recognised role of bats as bioindicators, challenges for the bat conservation associated with the anthropogenic transformation of the environment and climate changes determine the necessity of conducting long-time monitoring of their ranges, abundance, status, etc. (Jones et al., 2009; Sherwin et al., 2013; Voigt & Kingston, 2016), that all require the primary data for assigning the starting point in observations. By the end of the 20th century, data on bats of the vast territory of Central Ukraine had been quite scanty and patchy. Inside this region, there were only two plots comparatively well studied regarding the bat fauna: Kyiv with its vicinities and the territory in the Middle Dnipro River Region; in the fi rst case, due to the presence of the research node in the city of Kyiv; in the second, as a result of qualitative dissertation research in late 1960, early1970s (see below). Zoodiversity, 56(3): 203–232, 2022 DOI 10.15407/zoo2022.03.203 204 L. Godlevska, S. Rebrov, P. Vorobei, M. Savchenko, P. Panchenko In 1999–2021, we carried out extensive fi eld research in diff erent parts of Central Ukraine, which was supplemented with records done in the course of the work of the bat contact centres (kept by the authors). As a result, a big array of completely new data on bats in the region was gathered. Th e current work aims to describe the bat fauna of Central Ukraine (to the west from the Dnipro River) with a focus on the distribution and status of each species and ecological traits of bats in the region (prevalence, seasonality of occurrence, reproductive status, comparative abundance, and roosts). In addition, one of the tasks of this work is to present the array of original, earlier unpublished, primary data on bat records in the region. Study area Central Ukraine, in the paper, is considered in the meaning of the sociо-economic region, with adjacent territories (Shabliy, 1994). We excluded from the current review the part of Central Ukraine to the east from the Dnipro River as this area was considered in another review (Godlevska & Rebrov, 2018). Th e study area is stretched in the meridian direction along the Dnipro. In the north, it is limited by the administrative border of Ukraine; in the south, by the border of Ukraine and the northern border of the steppe zone; in the east, by the Dnipro riverbed; in the west, roughly by the border of the forest-steppe zone. Th e area falls into eight administrative regions of Ukraine: Kyiv, Cherkasy, Kirovohrad (with Kropyvnytskyi as the regional centre), Mykolayiv, Odesa, Vinnytsya, Khmelnytskyi, and Zhytomyr (fi g. 1). Th e total area of the study region is 123,000 km2. According to the classifi cation of physiographic regions, the study area includes part of the zone of mixed forests (Kyiv and Zhytomyr Polissia) and part of the forest-steppe zone (Marynych & Shyschenko, 2006). Within the part, corresponding to the forest-steppe zone, we distinguish the subregion of the Dniester River, based on its landscape features, climatic conditions, and the presence of a signifi cant number of, mainly limestone, mines (with a length, oft en, > 1 km). Th e high density of mines allows considering this subregion as the “cave” one, in contrast to the other (not-cave) parts of the study area, where underground cavities are presented by man- made structures like cellars, basements, fortifi cations, drainage tunnels, etc. Most of them have a small length (< 0.5 km) and volume; with the exception of the extended complex of drainage-mine systems of Kyiv (Godlevskaya, 2007). Th us, in the borders of the study area, there are three identifi ed subregions, from north to south: 1) mixed forests, ZMF, or Polissia; 2) forest-steppe, ZFS; 3) the Dniester River, DRS (fi g. 1). Fig. 1. Study area and localities (nloc = 265). A — data of other authors, 1848–2020 (nloc = 134); B — own data, 1999–2021 (nloc= 168). Subregions, here and further: 1, mixed forests, or Polissia (ZMF); 2, forest-steppe (ZFS); 3, the Dniester River (DRS); see text. 205Bats of Central Ukraine: a synopsis Material and methods Bat records, considered and analysed in the current review, concern two categories: A) data of other au- thors (publications, museum collections); B) own data, collected by the authors of the paper on their own or with their immediate participation. Some of the own data were already published but an extensive array of them is fi rstly presented by this paper. Annex (https://bit.ly/3ORCzGl) provides full details on earlier unpublished own bat records. (Referring below in the text to Annex corresponds to referring to earlier unpublished original data of the current paper’s authors.) Yet, the Annex contains the complete reference list of sources (publications and museums) providing data about bat records in the study region (in Ukrainian). Th e own data were collected in 1999–2021 in diff erent seasons of the year: 1) during over 55 fi eld expeditions and excursions; 2) during the bat survey in the city of Kyiv; and 3) in the course of the work of the bat contact centres. In total, the own data come from 168 study localities in eight administrative regions of Ukraine (fi g. 1). In the fieldwork, we applied the set of methods and approaches (the detailed description in: Godlevs- ka & Rebrov, 2018). Acoustic surveys were done with bat detectors: Pettersson Electronik D 200, D 240, and Wildlife Acoustics Echo Meter Touch 2 Pro. Bat sound records were analysed using Pettersson Elek- tronik BatSound and Wildlife Acoustics Kaleidoscope Pro. Netting was done mainly with mist-nets (3–12 m long; Chinese; Polish, Ecotone). Nets were set up near bat roosts, at their commuting paths, feeding and watering places. Examination of bats was carried out during the short time after capture. The bats were released at the capture site immediately after their examination. Bat roosts’ search was carried out using various methodical approaches (by the presence of faeces, social vocalisation of bats, swarming, etc.); all accessible underground cavities were examined. Where possible, the exact localisation of a roost and its description were done (allocation, type and structure of a roost, bat species, number of bats, the character of usage, etc.). Collecting faunistic data by calls to the bat contact centres was done by the scheme given in details in: Godlevska, 2012. In the paper, particularly for assessing distribution and prevalence of species, we use the term “study local- ity”, which is the conditional territorial unit which covers record(s’) point(s) located inside or at some distance to a certain settlement. In some cases, when the bat survey was done very remotely of settlements, additional localities were distinguished. Bat records inside the built-up districts of Kyiv, the western part of which lies in the borders of the study area, are not considered in detail. In the analysis of data, e. g. species prevalence, Kyiv is given as one study locality except for a few large nature protected areas inside the city administrative borders; they are considered as separate localities. Year periods, in regard to bat life-cycle, were accepted as: the breeding period, from 15th May to 15th August; the warm period of the year, from 16th March to 20th October; the winter period, from 21st October to 15th March. In the preparation of the current review, the own bat database was used (Godlevska, 2018). To outline the contribution of the original data in the general review’s data set, we used the diff erent symbols at species maps. Th e diagram of fi g. 2 indicates the origins of bat observations considered in the review. In the assessment of species prevalence, their comparative abundance, and used roosts the original data are marked accordingly. Abbreviations of bat species names, alphabetically: BBAR — Barbastella barbastellus; ENIL — Eptesicus nilssonii; ESER — E.  serotinus; MBEC — Myotis  bechsteinii; MBLY — M.  blythii; MDAS — M.  dasycneme; MDAU — M.  daubentonii; MMYO — M.  myotis, MMYS_gr — species of M.  mystacinus morphogroup, MNAT — M. nattereri; NLAS — Nyctalus lasiopterus; NLEI — N. leisleri; NNOC — N. noctula; PAUR — Ple- cotus auritus; PAUS — P. austriacus; PKUH — Pipistrellus kuhlii; PNAT — P. nathusii; PPIP — P. pipistrellus; PPYG — P. pygmaeus; RHIP — Rhinolophus hipposideros; VMUR — Vespertilio murinus. Th e history of the bat fauna survey in Central Ukraine Th e fi rst data about the bat fauna of the region refer to the middle of the 19th century (Kessler, 1851). Th e over half-century period aft er, in the history of the bat survey of Central Ukraine, is presented only by some collected specimens in museums. Further, in 1910–1941, Kyiv researchers fi rst collected the signifi cant data set on bats in Central Ukraine (Charlemagne, 1914, 1915, 1933; Popov, 1936, 1939; Abelentsev & Popov, 1956; see also in: Godlevska, 2013 b). First of all, the new data concerned Kyiv and vicinities, but also, records in some other terrains of the region were done. Among others, during these years, the researchers applied the method of fauna inventory by owl pellets’ survey. It enabled to widen data on the fauna of Ukraine, including those on bats of the region (e. g. Izotiv, 1932; Popov, 1932; Pidoplichka, 1932, 1937). Data from a few decades aft er World War II are mainly based on labelled specimens in museum collections. At this time, the work on bat banding, initiated in 1939 by B. Popov, was continued as well, partly at the territory of the study region (Abelentsev et al., 1968; 1969; 1970). Th e signifi cant array of bat data was collected on the Middle Dnipro Region (Cherkasy and Kirovohrad Regions) in the late 1960s and early 1970s by Sologor (1973), during her dissertation research. Data on bat records in this region in the following years are available in publications by Ruzhilenko, Tsvelykh (Ruzhilenko & Tsvelykh, 1992; Ruzhilenko et al., 1998). Recently, the bat survey here was carried out by Bilushenko (e. g.: 2009; 2013; 2014). Results of the bat survey in Kyiv and Kyiv Region in the late 1980s are available in the publication by Likhotop et al. (1990). Bat records in this and other regions of the study area were published by Zagorodniuk 206 L. Godlevska, S. Rebrov, P. Vorobei, M. Savchenko, P. Panchenko and colleagues (e.  g. Zagorodniuk & Tyschenko-Tyshkovets, 2001; Zagorodniuk & Godlevska, 2003; Zagorodniuk & Kalinichenko, 2008). In 1999, Godlevska initiated the systematic survey of bats in Kyiv and, in particularly, in underground structures of Kyiv and suburbs (Godlevskaya, 2007). In 1999, at the Kyiv Zoo, the bat rehabilitation centre was established by V. Tyshchenko and colleagues. Among other things, in the course of the centre’s work, P. kuhlii was fi rst recorded in Kyiv, a regular wintering of V. murinus and N. noctula was also revealed in Kyiv (Godlevsky et al., 2000; Tyshchenko, Godlevska, 2008). Recently, comprehensive bat surveys were done in two Kyiv protected areas: central and northern parts of the Holosiyivskyi National Nature Park (Vlaschenko et al., 2012; Vorobei et al., 2021), and Feofania Park (Bilushenko, 2016). Much attention was devoted to inventory of bat underground sites in the region. In addition to the mentioned bat survey in underground structures of Kyiv Region, in 2006, Godlevska and colleagues started the fi rst large-scale focused survey of bats’ underground sites in Podolian Upland and the Middle Dniester River region, which partly refer to the territory of Central Ukraine (Godlevska, 2010; 2012). Later, the inventory of underground shelters of bats in the so-called not-cave regions of Ukraine (without natural caves or mines, including the study region) was conducted (Godlevska et al., 2016; Annex). In 2009–2015, in the course of the work of the bat contact centre kept by Godlevska (2012; 2015) a lot of data on occurrence of bats all over Ukraine and, among others, in the study region were collected. Aft er 2015, two contact centres were kept with the participation of all authors of the review, in Kyiv and Odesa Regions. Th e intensive bat survey had been carried out in the Chernobyl Exclusion Zone (Gashchak et al., 2009; 2013). Data about bats of the western part of the study region are available in papers by Tyshchenko with colleagues (Tyshchenko, 2002; Tyshchenko et al., 2005) and Drebet with colleagues (Drebet, 2018; Drebet et al., 2020). Some more data on bat observations are also available in other papers of mentioned authors and, as well, of other researchers (Kryshtal, 1947; Golub, 1996; Vasiliev & Andreev, 1998; Zhyla et al., 2001; Smirnov & Smirnov, 2007; Zykov, 2011; Gaschak, 2018; and others: see Reference section in Annex). In 2001, Godlevska started the bat fi eld research aiming to obtain data on the bat fauna from all parts of the study region (besides Kyiv and suburbs). Further, in 2014–2019, all authors of the current review jointly carried out the bat survey in all eight administrative regions of the study area. Most of the obtained data fi rstly presented by this paper (Annex). Th e chronology of getting the faunistic bat data in the study region is illustrated by fi g. 2. In total, by 1999 (the starting point of the authors’ research), the number of bat species (according to their current taxonomic status) known in the region was 13: Barbastella, 1; Eptesicus, 1; Myotis, 3; Nyctalus, 3; Plecotus, 1; Pipistrellus, 2; Vespertilio, 1; Rhinolophus, 1 (by publications and museum collections). Species accounts At present, there are data on the occurrence of 24 bat species (of two families: Vespertilionidae and Rhinolophidae) in the region: Barbastella, 1 species; Eptesicus, 2; Myotis, 10; Nyctalus, 3; Plecotus, 2; Pipistrellus, 4; Vespertilio, 1; Rhinolophus, 1 (fi g. 2). We recorded 23 of them (all except Nyctalus lasiopterus). 18 48 18 58 18 68 18 78 18 88 18 98 19 08 19 18 19 28 19 38 19 48 19 58 19 68 19 78 19 88 19 98 20 08 20 18 A B N um be r o f r ec or ds 1999: 13 species 2021: 24 species Years Fig. 2. Number of bat records considered in the current review by years. A — data of other authors; B — own data. 207Bats of Central Ukraine: a synopsis Barbastella barbastellus (Schreber, 1774) Within the study region, the species has quite patchy distribution (fi g. 3). At present, it is rather frequently found in the northwestern part of the region and in the Dniester subregion. Th e Dnipro distribution patch of the species is relatively isolated from its other record localities. In Kyiv and outskirts, the species was regularly found in the middle of the 20th century (Abelentsev & Popov, 1956; Abelentsev et al., 1969; Godlevska, 2013 b). However, during the last two decades, the species was recorded only in two localities in the vicinities of Kyiv (Annex). Th e breeding was recently fi rst revealed in nine study localities (Annex). Maternity roosts are not known. During the breeding period, males were netted at entrances and inside abandoned banked military structures, up to 20 individuals per object (Annex). Th at allows considering such structures as roosts for the barbastelle males in the warm season of the year. In winter, the species was recorded hibernating in underground cavities: mines, military objects, cellars, drainage tunnels; earlier, in monastery artificial caves (e.  g.  Abelentsev & Popov, 1956; Sologor, 1973; Godlevska et al., 2010; 2012; 2016; Annex). The exception is two winter records of single individuals in overground parts of buildings: in a cavity of a stable ceiling and in a greenhouse (Sologor, 1973; Ruzhilenko et al., 1998). Th e number of hibernating individuals per one object is, usually, up to 10 individuals. Th ere are only few hibernacula, where more barbastelles were recorded, with a maximum of 194 individuals (Annex). Eptesicus nilssonii (Keyserling & Blasius, 1939) Within the study region, the species is rare. We recorded it just in two localities in the northwestern part of the region in 2015 and 2017 (fi g. 4). Th e only record of the species in the winter season in Kyiv (Zykov, 2011) may be considered as accidental. Th e breeding was not recorded. No roosts are known. Fig. 3. Record localities of Barbastella barbastellus. Here and further, if no other description is given: A — data of other authors, 1848–2020; B — own data, 1999–2021. 208 L. Godlevska, S. Rebrov, P. Vorobei, M. Savchenko, P. Panchenko Eptesicus serotinus (Schreber, 1774) Widely distributed and one of the most common, all year round occurring species in the region (fi g. 5 ). Summer roosts are overground sections of man-made structures and, rarer, underground cavities. Th e study area entirely falls within the species breeding range. In total, breeding was found in 28 localities; we recorded it in 26 study localities in diff erent parts of the study region. Fig. 4. Record localities of Eptesicus nilssonii. Fig. 5. Record localities of Eptesicus serotinus. 209Bats of Central Ukraine: a synopsis Maternity roosts have been localised in attic cavities of public (used) buildings, cavities in overground sections of abandoned buildings (Sologor, 1973; Annex). In winter time, the species is regularly recorded in buildings (in inner rooms, at balconies, between window frames, etc.) and at the ground within built-up areas of settlements (Godlevska, 2012; 2015 b; Annex). Notably, old winter records of the species in the region (the earliest is dated by 1928) came from the territory of settlements; and the places of revealing of serotine bats were the same as those at present (collected specimens in the NMNH NASU). Th e species is also recorded hibernating in underground shelters (Abelentsev & Popov, 1956; Godlevska et al., 2010; 2012; 2016). Usually, its number per one underground hibernaculum does not exceed few individuals. However, there is a known big winter aggregation of the species (> 100 individuals) in one of the limestone mines in the Dniester subregion (Godlevska et al., 2010). Myotis bechsteinii (Kuhl, 1817) Th e species is rare. It was revealed recently in seven localities in the southern part of the study region (fi g. 6); fi rst, in 2006 (Godlevska et al., 2010). Th e extreme eastern records (southeast of the Podolian Upland, in the north of Odesa Region) correspond to the eastern boundary of species distribution range in Ukraine and in Europe as a whole (Wright et al., 2018). Th e species was recorded hibernating in limestone mines (with the maximum number of 3 individuals per site) and during autumn swarming at the same underground objects. Th e breeding was fi rst recently confi rmed in the region, in two eastern points (Annex). Myotis blythii (Tomes, 1857) Like the previous species, M. blythii was revealed only within the Dniester subregion (fi g.  7) in six study localities; fi rst, in 2006 (Godlevska et al., 2010). Th e species was recorded both in winter and in the warm period of the year. Th e maximum number during hibernation per one site was 27 individuals. Maternity colonies are not known. However, in one locality (the north of Odesa Region), a single breeding female was netted at the mine entrance (Godlevska et al., 2010). Fig. 6. Record localities of Myotis bechsteinii. 210 L. Godlevska, S. Rebrov, P. Vorobei, M. Savchenko, P. Panchenko Myotis dasycneme (Boie, 1825) Although the species was found in all three subregions, its distribution there is highly patchy (fi g. 8). Th e species occurs year-round. During our survey, we fi rst confi rmed the breeding of the species in the region, in four study localities (Vorobei et al., 2021; Annex). Between 1999 and 2021, in the study region, only one maternity roost was found: a ceiling crevice Fig. 7. Record localities of Myotis blythii (A) and Myotis myotis (B); original data. Fig. 8. Record localities of Myotis dasycneme. 211Bats of Central Ukraine: a synopsis in the overground section of an abandoned building in Kyiv Region (Vorobei et al., 2021). Another potential maternity roost is in an abandoned building in Khmelnytskyi Region, where breeding females (together with adult males) were netted (Annex). In summer, M. dasycneme was recorded in tree cavities as well (Charlemagne, 1915; Abelentsev & Popov, 1956; museum specimens in: Zagorodniuk & Godlevska, 2001), and was netted at underground sites during late summer swarming (Godlevskaya, 2007; Godlevska et al., 2010; 2016). Winter roosts are underground; the maximum counted number of pond bats per hibernaculum was 12 individuals, but, in most cases, 1–4 individuals. Among animals netted during the breeding period, the percentage of pond bats was only 0.7 % (fi g.  24). During the autumn swarming, pond bats were netted at entrances to underground sites; however, their quantity was low; e.  g. in Kyiv, 0.2 % in 2003–2005 (Godlevskaya, 2007). All together enables to conclude that the population size of the species in the region is small. Myotis daubentonii (Kuhl, 1817) It is one of the most regularly occurring and common species in the study region. Here it was recorded all year round, in many study localities (fi g. 9). Summer roosts: bridges, overground sections of abandoned buildings, tree cavities, underground cavities. Th e breeding range includes, obviously, the whole territory of the study region. In total, breeding was confi rmed in three subregions in 24 study localities. We fi rst revealed maternity roosts of the species in the region: fi ve, in abandoned buildings; one, in a tree hollow (Annex). Winter shelters are various underground cavities (e. g. Abelentsev & Popov, 1956; Sologor, 1973; Likhotop et al., 1990; Godlevska et al., 2010; 2012; 2016; Annex). In the not- cave part of the study region (ZMF- and ZFS-subregions), the species dominates by number of individuals over other species in underground hibernacula; in the Dniester subregion, codominates (fi g.  25). Th e maximum number of M.  daubentonii per one underground hibernaculum in the region was 353 individuals (Kyiv, DMS  47-P1-2: winter 2016/2017; Annex). Fig. 9. Record localities of Myotis daubentonii. 212 L. Godlevska, S. Rebrov, P. Vorobei, M. Savchenko, P. Panchenko Myotis myotis (Borkhausen, 1797) Th e species was revealed only in western part of the Dniester subregion, in two study localities (fi g. 7); fi rst, in 2010 (Godlevska et al., 2012). Record localities are underground sites. Th ere, single specimens were revealed, both in the warm and winter seasons. Th e breeding was not confi rmed. Myotis mystacinus morphogroup Among European bat species, the group of whiskered bats is one of the most complicated both for systematics and fi eld identifi cation (e. g. Benda & Tsytsulina, 2000; Çoraman et al., 2020). We distinguished species of this group by morphological characters (using description from: Dietz & von Helversen, 2004). Part of specimens was identifi ed only to the group of four or three species, in particular those found during winter counts in hibernacula. In the study region, we identifi ed the occurrence of four bat species from the M. mystacinus morphogroup: M. brandtii, M. mystacinus, M. aurascens, M. alcathoe (fi g. 10, 11). All four species were fi rst recorded in the study region only in the last decades. In the ZMF- and ZFS-subregions, records of species of the group are, generally, very rare (Zagorodniuk & Godlevska, 2003; Godlevskaya, 2007; Gashchak et al., 2009; 2013; Annex). In the DRS-subregion, whiskered bats are recorded regularly; here, we fi rst identifi ed all four species. In general, clarifi cation of the distribution and status of M. mystacinus morphogroup species in the study region, like in other regions of Ukraine, requires further research with applying genetic methods. — Myotis brandtii (Eversmann, 1845). In the ZMF- and ZFS-subregions, it is very rare. Single individuals of this species were found there only in six study localities (fi g. 10). In the DRS-subregion, its abundance, at present, cannot be estimated correctly: most records of whiskered bats refer to the winter season and are identifi ed only to the group M. brandtii / M. mystacinus s. l. Fig. 10. Record localities of Myotis brandtii (A, B), Myotis alcathoe (C), and not identidied specimens of Myotis mystacinus morphogroup (D). 213Bats of Central Ukraine: a synopsis Th e breeding of the species in the study region was confi rmed only in one locality, in Zhytomyr Region, where one juvenile female was netted in the middle of July (Annex). As for now, there is no information about summer roosts. Only one winter shelter of this species was verifi ed: in a drainage mine in Kyiv, where a single individual was found (Annex). — Myotis alcathoe von Helversen & Heller, 2001. One juvenile specimen of a small whiskered bat (ra = 31.5 mm), identifi ed as M. alcathoe, was netted in the DRS-subregion, in Odesa Region, in 2014 (Annex). Any other records are so far absent (fi g. 10). — Myotis aurascens Kuzyakin, 1935. Bats identifi ed as M.  aurascens were recorded in the Dniester subregion, in three localities (fi g. 11): in two of them, in underground sites during the autumn swarming season; in one, by skeleton remains, found in a rock crevice (Annex). — Myotis mystacinus (Kuhl, 1817) s. s. was recorded in two localities. We identifi ed M.  mystacinus s.  s. in one underground site in the Dniester subregion during autumn swarming (Annex). In the ZMF-subregion, in summer 2009, an adult male, identifi ed as M. mystacinus s. s. was netted at the territory of the Chernobyl Exclusion Zone (Gash- chak et al., 2009). Th e latter record, however, stands alone among other records of whis- kered bats in this part of Ukraine. All other records of identifi ed to species whiskered bats here are presented by M.  brandtii (Godlevska et al., 2016 b; 2020; this paper). In adjacent territories of southern Belarus, only M.  brandtii was recorded (Shpak, 2010; Dombrovski, 2018). Number of whiskered bats (M. bra. + M. mys. s. l.) comprised ca. 6 % of all counted bat individuals in underground hibernacula of the DRS-subregion (fi g. 25, B). However, during winter counts, when no contact examination of bats is possible, the determination of species of this group was not done. Nevertheless, taking into account that all four species were identifi ed in the Dniester subregion, and that whiskered bats in Europe are considered to be sedentary (Hutterer et al., 2005), we presume that all these species winter in the DRS- subregion. Fig. 11. Record localities of Myotis mystacinus morphogroup species: M. mystacinus s. s. (A, B), Myotis aura- scens (C). 214 L. Godlevska, S. Rebrov, P. Vorobei, M. Savchenko, P. Panchenko Myotis nattereri (Kuhl, 1817) Th e species was revealed in three subregions. However, its distribution is irregular. Points of records may be combined in three rather separate patches: southern, in the Dniester subregion and adjacent territories; eastern, in the Central Dnipro zone; and northern, in Polissia (fi g. 12). Th e species was found in all seasons of the year. Th e breeding was revealed in three subregions in seven localities (Sologor, 1973; Biletskaya et al, 1990; Godlevska et al., 2012; Annex). In the course of the authors’ study, no maternity roosts were found. During the breeding season, juvenile individuals were netted in an abandoned building and at an underground site entrance (Annex); however, animals’ shelters were not localised. Earlier, breeding females with juveniles were found in a tree cavity and in a bird box, within the eastern distribution plot of the species in the study region (Sologor, 1973; Biletskaya et al., 1990). Solitary males and non-breeding females were found or netted inside overground sections of abandoned buildings, and in underground or semi-underground objects (Annex). Known winter roosts are only underground. Th e counted number per one hibernaculum was, commonly, 1–6 individuals. Bigger aggregations were found only in four sites. Th e maximum counted number of M.  nattereri in a hibernaculum was 68 ind.; in the ZMF- subregion (Godlevska et al., 2016 a). Nyctalus lasiopterus (Schreber, 1780) Th e species is extremely rare. It was recorded in the warm period of the year in eight localities with only 10 records in total (fi g.  13). Most records refer to the middle of the 20th century (Abelentsev & Popov, 1956; Zagorodniuk & Godlevska, 2001; Godlevska, 2013 b). Recently it was found only twice, at the territory of the Chernobyl Exclusion Zone (Gashchak et al., 2009; 2013). During our study, the species was not recorded. Two recent records in 2009 and 2013 (in the Chernobyl Exclusion Zone) were presented by an immature male and a female, both netted in the 3rd decade of July (Gashchak et al., 2009; 2013). Th ese individuals comprise only 0.06 % of the general number of bats, examined by Gaschak with colleagues in 2007–2013. However, the records may prove the species breeding in the ZMF-subregion. Fig. 12. Record localities of Myotis nattereri. 215Bats of Central Ukraine: a synopsis Nyctalus leisleri (Kuhl, 1817) Th e species was recorded in diff erent parts of the study region during the warm period of the year (fi g. 14). Records mostly come from forest and park patches. Th e region seems to be entirely within the breeding zone of the species: the breeding was recorded in all three subregions, in 24 localities (during our survey, in 21 study Fig. 13. Record localities of Nyctalus lasiopterus. Fig. 14. Record localities of Nyctalus leisleri. 216 L. Godlevska, S. Rebrov, P. Vorobei, M. Savchenko, P. Panchenko localities). Known roosts were located in hollow trees and one, in a bird box, in which the maternity colony of 46 individuals, including juveniles was found (Biletskaya et al., 1990). Winter records are not known. Nyctalus noctula (Schreber, 1774) Th e species is common and widespread in the region (fi g. 15). We recorded it in the majority of study localities. Th e breeding zone includes all the territory of the study region. In total, the breeding was marked in all three subregions, in 51 study localities (we’ve found in 43). Known earlier and recently found summer roosts were almost exceptionally in hollow trees (tens of roosts). Maternity colonies were found only in hollow trees. Known records in buildings during summer are not numerous and concern the migration season; thus, such roosts are, obviously, transit. E. g. we recorded a colony of common noctules, in which all four sex-age groups were presented (MMad, MMjuv, FFad, FFjuv) in an abandoned concrete hangar in Cherkasy Region in late August, 2018 (Annex). At present, Central Ukraine is entirely within the species winter range due to its recent expansion northward (Godlevska, 2015 a). Th e fi rst winter records of the species in the region came from Kyiv in the early 2000s. Since then, it is regularly found there in winter. All known records in winter refer to the territory of settlements, and all localised by us winter roosts were in buildings, mostly multi-storey. Identifi ed roosts were: diff erent structural cavities, ventilation channels, cavities inside balconies’ facing, etc. Th e number of individuals in one winter colony is estimated to reach a few thousand (Godlevska, 2015 a; L. Godlevska, comm.). Recently, Bilushenko (2015) reported his observation of N. noctula colonies in hollow trees in winter in Cherkasy and Kyiv. Pipistrellus kuhlii (Kuhl, 1817) During the last half a century, the species expanded into many regions of Europe (e. g. Strelkov, 2004; Ancillotto et al., 2016; Lučan et al., 2020). For Central Ukraine, it is new as well. Here, its occurrence was fi rst recorded in Kyiv in 1999 (Godlevsky et al., 2000). At Fig. 15. Record localities of Nyctalus noctula. 217Bats of Central Ukraine: a synopsis present, the species is distributed throughout the study region, although in the northwest it occurs rarer than in other parts (fi g. 16). Th e species is recorded year-round. All known roosts, including maternity ones, were found out only in overground sections of buildings (in diff erent cavities). Th ree biggest maternity colonies counted were from ca. 50 to over 100 individuals, with juveniles (Bilushenko, 2013; Annex). Fig. 16. Record localities of Pipistrellus kuhlii. Fig. 17. Record localities of Pipistrellus nathusii. 218 L. Godlevska, S. Rebrov, P. Vorobei, M. Savchenko, P. Panchenko Known hibernation roosts are in buildings (all localised ones were in cavities beside window frames). Wintering groups and colonies were recorded during repair works (Godlevska, 2015 b; Hukov et al., 2020). Th e maximum colony size in winter for Central Ukraine was 138 individuals (Hukov et al., 2020). Pipistrellus nathusii (Keyserling & Blasius, 1839) Th e species was recorded all over the study region (fi g. 17). Winter records within the study region are absent. Th e earliest date of a record in the warm period is 05.04 (1954); the latest, 10.10 (2013) (Abelentsev & Popov, 1956; Bilushenkо, 2014). In the warm period of the year, it is one of the common bat species (fi g. 24). Roosts are: cavities in buildings, both used and abandoned, hollow trees. Maternity colonies are sometimes mixed with P.  pygmaeus. Th e breeding was revealed in the ZMF- and ZFS- subregions, in 40 localities, in total. Th e biggest of counted maternity colonies was of > = 90 adult individuals (Vorobei et al., 2021). Pipistrellus pipistrellus (Schreber, 1774) N o t e . P. pipistrellus and P. pygmaeus were recognised as separate species in late 1990s (Barratt et al., 1997; Mayer & von Helversen, 2001). Correspondingly, in the publications on bats of Ukraine before 2000, this pair of species was mentioned as P. pipistrellus (fi g. 18, D). We identifi ed species of this pair of pipistrelles by morphological characters (on Dietz & von Helversen, 2004). In some cases the identifi cation was verifi ed by recording echolocation signals during bats’ release. In the case with P. pygmaeus, acoustic observations were taken into consideration (this method is not reliable enough for P. pipistrellus s. s. because of the signifi cant overlapping of its echolocation signals with those in P. nathusii и P. kuhlii). Th e occurrence of P. pipistrellus s. s. was confi rmed recently only in the south of the study area, in the DRS-subregion (fi g. 18). Th ere, the breeding of the species was confi rmed (Annex), and its hibernation (in mines) was recorded (Godlevska et al., 2012). In the rest of the study area, only P. pygmaeus was recorded. Fig. 18. Record localities of Pipistrellus pygmaeus (A, B), P. pipistrellus s. s. (C), and P. pipistrellus s. l. (D). 219Bats of Central Ukraine: a synopsis Pipistrellus pygmaeus (Leach, 1825) Th e species was fi rst mentioned as the species for the region only in 2001 (Zagorodniuk, Tyschenko-Tyshkovets, 2021; see Note to P. pipistrellus). Further, the species was revealed in many localities all over the study region (fi g. 18). Winter records are absent. In total, during the last two decades, the breeding of P. pygmaeus was recorded in all three subregions in 35 study localities. Roosts are hollow trees, diff erent cavities in overground sections of buildings, both abandoned and used (Vlaschenko et al., 2012; Godlevska et al., 2021; Annex). All checked by us colony roosts (9 of 18 found) were maternity. Th e largest counted colony was > = 177 individuals (Annex). Mixed maternity colonies with P. nathusii are known (Vlaschenko et al., 2012; Annex). Taking into account the data on the current distribution of P.  pygmaeus and P. pipisrellus s. s. in the region, we presume that all old records of P. pipistrellus s. l. in the ZMF- and ZFS-subregions (fi g. 18, D) may be reff ered to P. pygmaeus. Plecotus auritus (Linnaeus, 1758) N o t e . Till late 1990s, in publications of Ukrainian zoologists, long-eared bats (P.  auritus and P.  austriacus) were considered as one species (fi g.  19, C). Th erefore, to estimate the distribution of these two species in the past only by publications is not possible. However, in museum collections, there are 34 specimens collected within the study region (in ZMF- and ZFS-subregions) in 1912–1969. All of them are P. auritus s. s. (Zagorodniuk & Godlevska, 2001; Godlevska, 2013 b). P. auritus was recorded in three subregions (fi g. 19), all year round. Th e breeding was confi rmed in three subregions (in total, in 23 localities). Revealed maternity roosts were in underground cavities, n = 3; and in overground sections of (abandoned) buildings, n = 4 (Godlevska et al, 2016 a; Annex). In the breeding roosts, adult males were recorded together with females and juveniles. Th e maximum number of the summer colony, found in a big abandoned factory cellar, was about 60 individuals, with 1M, 7Fad-repr, 1Mjuv examined (Annex). Fig. 19. Record localities of Plecotus auritus (A, B), Plecotus spp. (C). 220 L. Godlevska, S. Rebrov, P. Vorobei, M. Savchenko, P. Panchenko Known hibernacula are underground. Th e maximum hibernation number per site, counted in one of the mines in the Dniester region (VRB-1 mine, Vinnytsya Region), was 55 individuals (Godlevska et al., 2012). However, in most hibernacula, the counted number of P. auritus was 1–12 individuals. Plecotus austriacus (Fischer, 1829) Th e current distribution of the species in the region (fi g.  20) seems to be the consequence of the recent and continued nowadays range expansion in northern, north- eastern and eastern directions. Th e records of the species in the ZMF- and ZFS-subregions were previously absent; see Note to P. auritus. Th e occurrence of the species there was fi rst reported by Bilushenko (2009), in the ZFS-subregion. Until recently, the northernmost record locality of P. austriacus was at the border of the ZFS and ZMF-subregions in Kyiv (Godlevska et al., 2016 a). However, in winter 2020/2021 the species was fi rst recorded in the ZMF-subregion (Annex). Notably, all record points of the species in these subregions concern anthropogenic objects or habitats (mostly in diff erent man-made structures). Th e bat fauna of the DRS-subregion was studied much less than in two other regions; focused bat studies were carried out only recently. Th us, it is not possible to estimate how new the species is for the subregion. During the two last decades, the species in the DRS- subregion was recorded in almost the same number of localities as P.  auritus (14 and 16 correspondingly). However, in comparison with P. auritus, an abundance of P. austriacus seems to be lower. In particular, the total number of P.  austriacus in the underground hibernacula of the DRS-subregion was 32 individuals against 96 in P. auritus (fi g. 25). In the study region, P. austriacus is recorded in all seasons of the year. Breeding females and/or juvenile individuals were found in seven localities of ZFS- and DRS-subregions (Godlevska et al., 2010; 2021; Annex). Revealed maternity roosts were underground (mines) or in overground sections of (abandoned) buildings (ibid). Known hibernation roosts are underground; the number per one hibernacula was up to 9 individuals (Godlevska et al., 2012). Fig. 20. Record localities of Plecotus austriacus. 221Bats of Central Ukraine: a synopsis Vespertilio murinus Linnaeus, 1758 Th e species was recorded in all three subregions (fi g.  21). However, its occurrence within the study region is characterized by evident seasonal dynamics. We found breeding of the species only in two localities in the northwest of the study region with one localised maternity roost in the attic space (Annex). In the northeast of the region, in the Chernobyl Exclusion Zone, Gaschak et al. (2009; 2013) regularly recorded V.  murinus during the breeding season, in 2007–2013: individuals of this species there comprised 5 % of all netted bats. In other parts of the study region, during the breeding season, the species is almost absent. Summer, exactly dated, observations of the species, there, are presented by single specimens with no breeding females or juveniles among them (e. g. Bilushenko, 2016; Ruzhilenko et al., 1998; Godlevska, 2012; 2015). In previous publications, there are mentions of maternity colonies in two localities of the ZFS-subregion: one, in a hollow oak in southern vicinities of Kyiv (Charlemagne, 1933); the second, two colonies in hollow trees in Cherkasy Region (Sologor, 1973). Taking into account the absence of any other records of maternity colonies in the ZFS- and Dniester subregions, both in past and present, we presume that mentioned records might concern colonies of Nyctalus leisleri, the species which is similar by size with V. murinus and which, in Ukraine, typically roosts in trees (e. g. Vlaschenko, 2009; Godlevska & Rebrov, 2018; Annex). Misidentifi cation between these two species occurs even now (L. Godlevska, comm.). Since 1998, V.  murinus is recorded regularly in autumn and winter in large settlements, mostly in districts of multi-storey buildings, where parti-coloured bats are regularly found in inner rooms, at the outside surface of buildings or on the ground, in particular in Kyiv. Inside such districts, the autumn display song of males of V. murinus is recorded as well (Godlevska, 2013 a; 2015; Annex). Th e current regular winter occurrence of V. murinus in the region evidently indicates another case of expansion of the bat winter range (Godlevska, 2013 a). Noteworthy that the described seasonal dynamics of the species occurrence is similar to that at the adjacent territories to the east from the Dnipro River (Godlevska, Rebrov, 2018): the breeding is known only in the northernmost part, summer records are single, and in winter parti-coloured bats are found in settlements. Fig. 21. Record localities of Vespertilio murinus. 222 L. Godlevska, S. Rebrov, P. Vorobei, M. Savchenko, P. Panchenko Rhinolophus hipposideros (Borkhausen, 1797) Th e distribution of the species within the region is restricted to its southern part, to the Dniester subregion (fi g. 22), where it was recorded in 10 study localities in diff erent seasons of the year. All found roosts, both winter and summer, were underground. Th e maximum counted number per one hibernaculum was 167 individuals (Godlevska et al., 2010). Maternity colonies were revealed in three underground sites; the largest counted ca. 50 adult females (ibid; Annex). Chiropterofauna of subregions, seasonality of species occurrence, their reproductive status Th e summary on bat species’ occurrence by subregions, seasons, and their reproductive status is presented in table 1. In total, in the ZMF-subregion, 17 bat species were recorded; in the ZFS-subregion, 15. Th e diff erence concerns two species: E. nilssonii and M. mystacinus s. s. Th e former is known by records in few study localities of the ZMF-subregion; the latter was found there only once. P. austriacus is the new species for the ZMF-subregion. N. lasiopterus is an extremely rare species, which was recorded only twice in the ZMF-subregion during the last two decades. In the rest, species composition and reproductive status of species of these subregions are pretty similar. Th e highest number of species (in total, 23) was recorded in the Dniester subregion, which is the smallest by area. Here, so-called cave species (R.  hipposideros, M.  blythii, M. myotis) and M. bechsteinii, species of M. mystacinus morphogroup, P. pipistrellus s. s. were found. Winter occurrence of 17 species in the region was established with direct observations. Occurrence, in winter, of three species of M. mystacinus morphogroup: M. alcathoe, M. aurascens and M. mystacinus s. s. is presumed. Winter records for four, known to be long-distance migrants (Hutterer et al., 2005), species (P. nathusii, P. pygmaeus, N. leisleri, N. lasiopterus) are not known in the study region; it is presumed to be out of their winter range. During the last two decades (and in general), reproduction was confi rmed for 20 bat species in the study region. So far, there are no data on breeding of four species (E.  nilssonii, M.  myotis, M.  aurascens, M.  mystacinus) in the region. In the case of the Fig. 22. Record localities of Rhinolophus hipposideros. 223Bats of Central Ukraine: a synopsis T a b l e 1 . Bat species by three subregions of Central Ukraine: mixed forests (ZMF), forest-steppe (ZFS), and Dniester River (DRS), in the warm period of year (S) and in winter (W) Species ZMF-S ZMF-W ZFS-S ZFS-W DRS-S DRS-W B. barbastellus + repr + + repr + + + E. nilssonii * + + – – – – E. serotinus + repr + + repr + + repr + M. alcathoe * – – – – + repr (+) M. aurascens * – – – – + (+) M. bechs teinii * – – – – + repr + M. blythii * – – – – + repr + M. brandtii * + + + repr – + (+) M. dasycneme + repr + + repr + + + M. daubentonii + repr + + repr + + repr + M. myotis * – – – – + + M. mystacinus * +? – – – + (+) M. nattereri + repr + + [repr] + + repr + N. lasiopterus + repr – [+] – [+] – N. leisleri + repr – + repr – + repr – N. noctula + repr + + repr + + repr – P. auritus + repr + + repr + + repr + P. austriacus * (+) + + repr + + repr + P. kuhlii * + repr + + repr + + repr (+) P. nathusii + repr – + repr – + – P. pipistrellus * – – – – + repr + P. pygmaeus + repr – + repr – + repr – R. hipposideros – – – – + repr + V. murinus + repr + + [repr?] + [+] (+) Total, species 16 (17) 12 15 10 23 12 (18) 17 15 23 * Th e species fi rst found in the region during the last two decades (old records of P. pipistrellus s. l. were assigned to P. pygmaeus); repr — breeding of a species was confi rmed by records of breeding females and / or juvenile individuals in the breeding period; [+] / [repr] — species occurrence / breeding was recorded before 1999; [repr?] — breeding was mentioned in publications, but reliability of species identifi cation is questionable; (+) — species occurrence is presumed. 0 1 3 3 6 7 7 10 17 2 0 28 3 0 37 41 45 60 70 76 7 8 11 4 11 6 1 2 3 3 6 7 25 11 19 23 29 33 57 46 66 79 90 10 9 94 14 7 13 5 NL AS M M YO EN IL PP IP ss M BL Y M BE C VM UR RH IP M M YS _g r M DA S PA US M NA T PK UH BB AR NL EI PP YG PA UR ss PN AT M DA U NN OC ES ER a b n, lo ca lit ie s Fig. 23. Bat species of Central Ukraine by number of localities in 1999–2021, according to: a, own data; b, sum of all available geographically attributed data for the time period. fi rst two species, the region seems to be on the edge of their main distribution ranges. In M. aurascens and M. mystacinus the breeding in the region is presumed; given their sedentary. 224 L. Godlevska, S. Rebrov, P. Vorobei, M. Savchenko, P. Panchenko Distribution of species, their prevalence and abundance In total, by results of 1999–2021 study, the most common, by the number of localities, species are: E. serotinus, N. noctula, M. daubentonii, P. nathusii, P. auritus and P. pygmaeus. Other species were found in a fewer number of localities. Th e species N.  lasiopterus, M.  myotis, E.  nilssonii, P. pipistrellus s. s., M. blythii, M. bechsteinii have the restricted distribution (see Species accounts). Th ey are known from the least (among other species) number of localities (fi g. 23). In general, our own results on the prevalence of species (fi g. 23, a) correspond to the broader array of data (fi g. 23, b). Th e only species with a considerable diff erence in a number of localities (by own vs. sum of all data) is V. murinus. Th e majority of known localities of this species are in the Chernobyl Exclusion Zone. Th ere, the species is regularly found in summer (Gashchak et al., 2009; 2013) in contrast to the biggest part of the study region (see V. murinus species account). Th e most prevalent species have, in general, the highest abundance. In particular, the quantitative representativeness of these species among netted bats during the breeding season was high in all three subregions (fi g. 24). Bat roosts In total, in the course of our study within the region, we revealed and/or examined 108 underground bat shelters; with the length of a few meters to over 10 km each (Godlevskaya, 2007; Godlevska et al., 2010; 2012; 2016; Annex). Data on two underground bat roosts for the same time period are available in publications of other authors (Smirnov & Smirnov, 2007; Bugaichenko, 2019). During 1999–2021, in underground shelters of two not-cave subregions (ZMF and ZFS), we revealed eight bat species: B. barbastellus, E. serotinus, M. brandtii, M. dasycneme, M. daubentonii, M. nattereri, P. auritus, and P. austriacus. Th ey exhaust the full list of bat species been recorded in underground sites of the not-cave (NC) subregions. In summer, in underground sites of this, not-cave, part of the region, we found bats in comparatively low numbers. Maternity colonies in underground structures of the NC subregions were observed only in one species, P. auritus, in two objects (Godlevska et al., 2016; Annex). Also breeding females of M. daubentonii were caught at the entrance to one semiunderground object (Annex). In the NC part of the region, in 1999–2021, we examined 50 underground hibernacula: in the ZMF-subregion, 34; in the ZFS-subregion, 16. Th e maximum bats’ number per one underground hibernaculum in the NC subregions was 356 individuals: in DMS  47- P1-2, Kyiv (Annex). However, the median value of bats’ number per one underground hibernaculum was 4 individuals (Nav = 42 ind.). In winter aggregations in underground hibernacula of the NC subregions, M. daubentonii predominated considerably (fi g. 25, А). Notably, the biggest portions (96 %) of counted hibernating bats in the NC subregions were revealed in less than a half of underground hibernacula (43 %). Th e largest complex of underground hibernacula in the NC subregions is represented by the drainage mine systems (DMS) in Kyiv. Th e general number of bats counted in DMSs (18 objects in diff erent years) amounts ca. 76 % of all bats in underground hibernacula in not-cave subregions. About 20 % of counted bats hibernating underground in the not-cave part of the region were found in four other objects in the ZMF-region (tree, military; one, a tunnel of unclear purpose). And only 4 % of bats were counted in the rest (56 %) of the examined underground hibernacula of this, not cave, part of the study region. More bat species were revealed in underground sites of the Dniester, cave, subregion, where there are many extended underground objects, diverse by structure and microclimate (Godlevska et al., 2010; 2012). Th ere, R. hipposideros, M. blythii, M. myotis, M. bechsteinii, and species of M. mystacinus morphogroup and P. pipistrellus s. s. are added to the list of species roosting underground (fi g. 25, B). In summer, in examined underground cavities of the Dniester subregion, maternity colonies of R. hipposideros (four sites) and P. austriacus (three sites) were recorded. As well, 225Bats of Central Ukraine: a synopsis breeding females of M.  blythii and P.  auritus were netted at the entrances to mines, that indirectly indicates the using such objects by these species as maternity. In winter, in the DRS-subregion, 26 underground hibernacula were inspected. Th e maximum number of bats per one underground hibernaculum was 371 individuals: in VRB1-mine, Vinnytsya Region (Godlevska et al., 2012). In comparison with the not-cave subregions, the median value of bats’ number per one underground hibernaculum was higher: 30 individuals (with Nav = 68 ind.). Dominant species (by the number of individuals) were R. hipposideros and M. daubentonii. However, the distribution of counted individuals by species in the Dniester region is less sharp than in not-cave subregions (fi g. 25, B). Overground bat roosts in the study region include various cavities in man-made structures, trees, and, probably, in rock and soil outcrops. 0 50 100 150 200 250 C 0 50 100 150 200 250 300 I II B 0 50 100 BB AR EN IL ES ER M BE C M MY S_ gr M DA S M DA U M NA T NL EI NN OC PA UR PA US PK UH PN AT PP IP PP YG VM UR A N , s pe ci m en s N , s pe ci m en s N , s pe ci m en s Fig. 24. Quantitative representativeness of species among netted bats during breeding season in 1999–2021 in subregions: A, Dniester River, DRS (nind = 265); B, forest-steppe, ZFS (nind = 1097); C, mixed forests, ZMF (nind = 1007). I — all netted specimens; II — bats netted remotely of identifi ed roosts. Results of catchings at underground sites in the DRS were not included. Only own data were used. 226 L. Godlevska, S. Rebrov, P. Vorobei, M. Savchenko, P. Panchenko In the warm period of the year, we revealed 162 overground bat roosts (67, in trees; 95, in man-made structures). For 135 of them, at least one species of sheltered bats was identi- fi ed; for 27, the species of sheltered bats was not identifi ed or identifi ed only to a genus (e. g. Nyctalus, P. nathusii/P. kuhlii; etc.). In total, roosts of 14 species were localised (fi g. 26). In general, these species are the all ones with localised summer overground roosts during the last two decades of the bat survey in the region. Th e largest number of such roosts, in the warm period of the year, was found for four species: E. serotinus, N. noctula, P. nathusii, P. pygmaeus, which are among the most fre- quent and abundant species in the region (see above). Roosts of six species (M. daubento- nii, N. noctula, N. leisleri, P. auritus, P. pygmaeus, P. nathusii) were found in tree cavities. N. leisleri was found roosting only in trees; N. noctula, almost solely. In overground sec- tions of various man-made structures, roosts of 13 species were revealed. Among them, eight species were revealed roosting only in the man-made structures (fi g. 26). In winter, we localised overground bat roosts only in buildings (Godlevska, 2015 a; Annex; L.  Godlevska, comm.) where four species (N.  noctula, P.  kuhlii, E.  serotinus, V.  murinus) were regularly found. Th ese four species also predominate by the number of calls to contact centres, both within the study region and Ukraine overall (Godlevska, 2012 b, M M YS _g r M BE C M M YO M M YS _g r M DA S M DA S M DA U M NA T M BL Y PA US PA US M DA U RH IP BB AR BB AR M NA T PA UR ss PA UR ss ES ER ES ER PP IP s 5623 32 737382 96108 115 168 438 478 1545 212 125 59 25 6 6 4 Fig. 25. Quantitative representativeness of bat species in underground hibernacula of the study area (by own data of 1999–2021). A, ZMF- and ZFS-subregions (34 and 16 objects; 1931 and 51 identifi ed ind., correspondingly); B, DRS-subregion (26 objects, 1697 identifi ed ind.). By: Godlevskaya, 2007; Godlevska et al., 2010, 2012; this paper. In case of numerous counts, maximal numbers were taken. 227Bats of Central Ukraine: a synopsis 2015 b; Prylutska & Vlaschenko, 2013; Panchenko & Godlevska, 2018). N. noctula is known as the species forming large hibernation colonies in hundreds of specimens in multi-storey buildings in settlements (Godlevska, 2015 a). P.  kuhlii hibernate in cavities of overground sections of buildings in groups or colonies with a number up to 150 individuals (see P. kuhlii species account). Numerous and regular records of E.  serotinus and V.  murinus in inner rooms of buildings are the evidence of their wintering in cavities of these structures. Of the four species mentioned, three are “new” in winter in the region: N. noctula, V. murinus, due to the expansion of their winter range; P. kuhlii, due to the expansion of the general range. In buildings, in the inner rooms of their overground sections, there were few records of single specimens of P. austriacus and Plecotus sp. (Ruzhilenko et al., 1998; Bilushenko, 2009; Annex). Th at may be considered as the evidence of roosting of long-eared bats in buildings in winter as well. Th ere were also two records of single specimens of B.  barbastellus in overground sections of buildings (see the species account). Few records of solitary bats in hollow trees in the winter period concern, mainly, the autumn (October) or spring (March) seasons (Abelentsev & Popov, 1956; Sologor, 1973). Th ese seasons in certain years may be characterised by rather high temperatures, and, correspondingly, such shelters may be transit or temporal. Th e only mention in publications of bats (N. noctula) in hollow trees during the main winter period in the region, namely in the cities of Kyiv and Cherkasy, concerns the last decade (Bilushenko, 2015). It can not be ruled out that the warming of winters allow bats, at present, to hibernate in trees in the regions (earlier) characterised by low temperatures and long periods of frosts (Babichenko et al., 2003). However, only further research will show whether the wintering of bats, particularly N. noctula, in trees is constant, or such shelters are satellite to those already known in buildings. One more type of conditionally overground roosts in crevices of rock or soil outcrops is not studied enough. In the borders of the study region, there were two records of bats at the surface of outcrops in the late winter period: in the middle of March. In particular, on the 15th of March 2008, few specimens of E. serotinus were observed at ledges of granite rocks (Zagorodniuk & Kalinichenko, 2008). Th e single specimen of B.  barbastellus was found at the loess wall of a ravine on the 7th of March 1992 (Ruzhilenko et al., 1998). Discussion Th e history of the bat fauna survey in Central Ukraine covers the period of over 170 years. However, the biggest portion of the available data on the bat fauna of this region refers to the period of the two last decades (fi g. 2). During 1999–2021, the list of bat species VM UR PP IP M DA S NL EI M NA T PA US BB AR PA UR PK UH M DA U PP YG ES ER NN OC PN AT 0 10 20 40 50 30 Building 1 2 2 0 4 4 7 7 14 13 13 19 1 45 Trees 0 0 0 4 0 0 0 2 0 1 8 3 43 0 Fig. 26. Localised overground bat roosts and their types (natural / anthropogenic) in summer by species, Central Ukraine (by own data, 1999–2021). 228 L. Godlevska, S. Rebrov, P. Vorobei, M. Savchenko, P. Panchenko was signifi cantly clarifi ed, both for the region overall and for its subregions. In particular, the presences of 11 bat species were fi rst identifi ed in the region (table 1). At that, only P. kuhlii is the new species of the regional fauna; others seem to be discovered due to the intensifi cation of the bat research activities. Th e distribution ranges of all species were signifi cantly clarifi ed both for the study region and Ukraine overall. Among other, the distribution of recently separated species (P. pipistrellus and P. pygmaeus, P. auritus and P. austriacus) was determined. Among the subregions, the Dniester subregion stands out by number of species both in winter and warm seasons of the year. Noteworthy that increase in the number of bat species known for the study region (from 13 to 24) occurred, to the large extent, due to observations fi rst carried out in the DRS-region (Godlevska et al., 2010; 2012; this paper). Only here, six of nine “new” species were fi rst recorded: M. aurascens, M. alcathoe, M.  bechsteinii, M. blythii, M.  myotis, and P.  pipistrellus s.  s. Th e high diversity of bat species in this subregion is, obviously, determined by its landscape diversity (Marynych & Shyschenko, 2006), microclimate and availability of the large number of underground shelters (see Study area). Th e total number of bat species of this subregion (namely 23) enables to identify it as one of the bat diversity hotspots in Ukraine; together with Carpathian and Crimean regions. Th e available data allow specifying the seasonality of occurrence and outlining more precisely the breeding ranges of studied species. Most of the data on breeding of bat species in the region were collected just in the last two decades; and most of them were obtained in the course of authors’ fi eld surveys. Availability of appropriate roosts is very important for any bat species. Recently received data enable to assess the distribution and signifi cance of underground bat roosts in the region. In particular, we show an unequal distribution of bats among known underground hibernacula in the not-cave subregions where 96 % of wintering underground bats were counted less in a half of all inspected hibernacula. Moreover, 76 % of these bats were revealed in the complex of drainage mines of Kyiv City. Th e distribution of hibernating bats among underground hibernacula in the Dniester subregion is more equal, in comparison with not-caves ones. As well, the distribution of counted individuals by species in the Dniester region is less sharp than in not-cave subregions (fi g. 25). Th is is, obviously, caused by the diversity of underground sites (including their structure, volumes and microclimate) of the Dniester subregion. With intensifi cation and broadening research activities in future, the number of bats hibernating underground in the region may be defi ned more precisely. However, available data unambiguously indicates the importance of already known large hibernacula for bat conservation, especially in the not-cave part of the region. During our surveys, we had collected a big array of data on overground bat roosts in the warm period of year (n = 162). We suppose that roosts in tree cavities (n = 67) were undercounted. Revealing and examining bat shelters in trees is a more methodologically complicated task, which oft en requires more time and eff ort than for roosts in man-made structures, which are more accessible for direct inspection. At the same time, the considerable number of revealed bat roosts in man-made structures (n = 95) shows the signifi cance of such types of shelters for bats in the region and determines the necessity to draw special attention to their protection in the development and implementation of bat conservation measures, both at regional and national levels. Th e same is highly relevant for winter bat shelters in buildings. Almost all known overground roosts of bats in winter were localised in buildings. In general, the data collected during the last two decades signifi cantly broadened the knowledge about the bat fauna of the region and Ukraine as a whole. In fact, the current review represents the “picture” of the bat fauna for this period. We suppose that it may be used further as a checkpoint both for monitoring certain species, their communities and localities or roosts. 229Bats of Central Ukraine: a synopsis Conclusions Th e bat fauna of Central Ukraine (to the west from the Dnipro River) includes 24 species: Eptesicus, 2 species; Barbastella, 1; Myotis, 10; Nyctalus, 3; Pipistrellus, 4; Plecotus, 2; Vespertilio, 1; and Rhinolophus, 1. During 1999–2021, the breeding was confi rmed in 20 species (all except E.  nilssonii, M. myotis, M. aurascens, M. mystacinus). Winter records of four, known to be long-distance migrants, species (P. nathusii, P. pygmaeus, N. leisleri, N. lasiopterus) are not known in the region. Winter occurrence of three species of M. mystacinus morphogroup is presumed. Species composition (and their reproductive status) in two subregions, of the mixed forests (ZMF) and forest steppe (ZFS), are pretty similar. In the ZMF-subregion, it total, 17 bat species were recorded; in the ZFS-subregion, 15. Th e highest number of species (23) was recorded in the Dniester subregion. By results of 1999–2021 studies, the most common species (by the number of localities) were E. serotinus, N. noctula, M. daubentonii, P. nathusii, P. auritus and P. pygmaeus. Other species were found in a fewer number of localities. Th e species N. lasiopterus, M. myotis, E. nilssonii, P. pipistrellus s. s., M. blythii, M. bechsteinii have the restricted distribution. Th eir records come from the least (among other species) number of localities. Th e most common species have, in general, the highest abundance. Th e quantitative representativeness of these species among netted bats during the breeding season was high in all three subregions. Eight species were recorded in underground sites of the not-cave part of the study area (ZMF- and ZFS-subregions). Maternity colonies in underground structures of the NC subregions were observed only in P. auritus. Th e maximum bats’ number per one underground hibernaculum in the not-cave subregions was 356. In these subregions, the biggest portion of counted hibernating bats (96 %) was revealed in less than a half of inspected underground hibernacula. Th e dominant species among hibernating bats in underground sites of these subregions was M. daubentonii. In the Dniester subregion, 13 bat species were found in underground sites. At least two species (R. hipposideros, P. austriacus) use such sites as maternity roosts. Th e maximum number of bats per one underground hibernaculum was 371. Th e distribution of hibernating bats among underground hibernacula in the Dniester subregion is more even than in those of not-cave subregions (median values of bats’ number per one underground hibernaculum: 30 vs. 4 individuals, correspondingly). Th e dominant species in winter aggregations (by number of counted individuals) were R. hipposideros and M. daubentonii. During the two last decades, in the warm period of the year, overground roosts of 14 species were revealed. During our survey in the warm period of the year, we revealed 162 overground bat roosts, and for 135 of them at least one sheltered bat species was identifi ed. Th e largest numbers of such roosts were found for four species, which are among the most common and abundant in the region: E. serotinus, N. noctula, P. nathusii, and P. pygmaeus. In the warm period of the year, six species were found roosting in tree cavities; and 13, in overground sections of various man-made structures (buildings, hangars, bridges, military structures). In winter, almost all known overground bat roosts are in buildings. Species that are regularly found to hibernate in such shelters are N. noctula, P. kuhlii, E. serotinus, V. murinus. We sincerely thank N.  Atamas, I.  Balashov, A. Bednarchuk, P.  Buzunko, S.  Davydenko, D.  Demchenko, M. Ghazali, P. Gol’din, A. Grushko, A. Haleyeva, L. Kravchuk, O. Krokhmal, A. Larchenko, P. Lina, A. Marchevka, M. Maruschak, O. Mischenko, I. Parnikoza, I. Pobivantseva, T. Postawa, O. Proskura, Ju. Pryadko, M. Storozhuk, V.  Tyshchenko, A.  Plyha, Zh. Zynkevich, all who helped us in diff erent years with the fi eldwork. We thank L.  Kravchuk, E.  Zhuravleva, A.  and  B. Paschenko, M.  Storozhuk, members of the Kyiv Animal Rescue Group, and O. Bubela, L. Bugaichenko, V. Frankova, O. Gulai, Ju. Kutsokon, O. Matviychuk, Ju. Nesin, V. Linkevich, Je. Rudenko, B. Popov, O. Vasyljuk, D. Vishnevskiy, S. Zhyla, for communications about bat fi ndings; R. Ivanenko, O. Kovalenko, A. Plyha, A. Podobailo, S. Zhyla for assistance in organization of the fi eldwork. Part of the fi eldwork was carried out with the fi nancial support of: Stichting Zoogdierenwerkgroep Zuid- Holland, Bat Support Fund for Eastern Europe, Ruff ord Foundation, European Mammal Foundation, Bobryk Village Charity “Development and prosperity”, Fund for the protection of biodiversity of Ukraine, and Fund of Dovkolabotanica’s Friends. 230 L. Godlevska, S. Rebrov, P. Vorobei, M. Savchenko, P. Panchenko We thank P.  Gol’din, for the discussion of the manuscript; two anonymous reviewers, for the valuable critical comments and remarks. References Abelentsev, V. I., Popov, B. M. 1956. Order Chiroptera, or bats. In: Pidoplichko, I. G. Fauna Ukrainy. Vol. 1: Mammals. Vyd-vo AN URSR, Kyiv, 229–446 [In Ukrainian]. 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