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  • Northern Bats
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Articles published on Barbastella barbastellus

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  • Research Article
  • 10.1016/j.foreco.2026.123634
Habitat preferences of specialist bat are forest-type specific: Western barbastelle in natural temperate woodlands
  • Jun 1, 2026
  • Forest Ecology and Management
  • Fabian Przepióra + 5 more

Habitat preferences of specialist bat are forest-type specific: Western barbastelle in natural temperate woodlands

  • Research Article
  • 10.1002/ece3.72949
Comparative Analysis of Mitochondrial Genomes and Phylogeny of Barbastelle Bats Across China
  • Jan 1, 2026
  • Ecology and Evolution
  • Sen Liu + 6 more

ABSTRACTBarbastelle bats are notable for their nearly square outer ears that converge at the forehead. Knowledge of their mitochondrial genome is limited. This study sequenced the complete mitochondrial genomes of Barbastella beijingensis and B. darjelingensis, the latter being reported for the first time. Subsequently, we performed a comprehensive comparative analysis with the published mitochondrial genome of B. capsica. The mitochondrial genomes measured 16,667 bp and 16,434 bp, respectively, and included 13 protein‐coding genes (PCGs), 22 tRNA genes, 2 rRNA genes, and one D‐loop. Their gene order matched other bat species in the Vespertilionidae and Phyllostomidae families. All three species showed low Ka/Ks ratios in the PCGs, indicating strong purifying selection, especially in cytochrome oxidase respiratory chain complex (COX1, COX2, and COX3), while ATP8 faced relatively relaxed selective pressure. In all three species, tRNAs showed typical cloverleaf structures, except for trnS1, which lacked a D arm. The D‐loop region is divided into ETAS, CD, and CSB domains. Both ETAS domains have ETAS1 and ETAS2 motifs, and the CD domain contains conserved motifs box B–F. While CSB1–CSB3 were conserved in B. capsica, only CSB1 was found in B. beijingensis and B. darjelingensis. Phylogenetic analysis of the 13 PCGs strongly confirmed the three barbastelle species as distinct within Barbastella, closely related to genera Plecotus and Corynorhinus in the Plecotini tribe. This study enhances the molecular data available for the genus Barbastella and offers essential mitochondrial genomic evidence to resolve persistent taxonomic disputes within the Vespertilionidae family.

  • Research Article
  • 10.1111/1365-2664.70205
Managing hedgerows for biodiversity: Disentangling the effects of trimming, structure and connectivity on the use of linear features by bats
  • Nov 25, 2025
  • Journal of Applied Ecology
  • Noé Tarcy + 4 more

Abstract Hedgerows are key semi‐natural elements in European agricultural landscapes supporting diverse wildlife, including bats, that depend on these linear elements for foraging and commuting. Despite replanting initiatives, many hedgerows remain degraded due to intensive management practices such as overtrimming. To develop effective and generalisable conservation guidelines, more research is needed to understand how both management practices and intrinsic hedgerow characteristics influence bat species with different foraging strategies. We conducted passive acoustic monitoring across 448 hedgerows to assess the influence of six characteristics on bat activity: trimming method, height, width, foliage density, connectivity and woody plant diversity. We examined responses for two species associated with hedgerow landscapes ( Barbastella barbastellus , Rhinolophus ferrumequinum ) and three echolocation guilds: short‐, mid‐ and long‐range echolocators (SRE, MRE, LRE), corresponding to clutter, edge and open‐space foragers, respectively. Pollarding consistently increased bat activity, especially for tree‐dwelling species, likely by maintaining mature trees with larger crowns and diversified microhabitats. Coppicing generally reduced activity for SRE, B. barbastellus and R. ferrumequinum , while increasing LRE activity. Taller, tree‐filled hedgerows generally supported higher bat activity, whereas width and foliage density had more limited effects. Finally, uninterrupted hedgerow segments enhanced SRE and MRE activity, highlighting the importance of local habitat continuity in creating effective corridors for bat movement. Woody plant diversity was also positively associated with SRE activity. These results were mostly consistent across simplified open croplands and structurally complex bocage landscapes. Synthesis and applications . Our findings underscore the value of extensive hedgerow management in promoting on‐farm bat activity and support the integration of simple hedgerow quality assessment protocols into agri‐environment schemes. Consistent with previous research, we confirm the importance of tall, tree‐filled hedgerows and emphasise the need to preserve mature trees. We also provide novel insights: hedgerow continuity and low‐intensity trimming methods can further improve habitat suitability, particularly for tree‐dwelling bats. Our management recommendations align with evidence from other taxonomic groups, including birds and invertebrates, and can contribute to broader biodiversity conservation strategies.

  • Research Article
  • 10.1038/s41598-025-14315-2
Increasing canopy cover elevates vehicle collision risk for barbastelle bats (Barbastella barbastellus) at roads
  • Oct 23, 2025
  • Scientific Reports
  • Kieran D O’Malley + 5 more

Roads can form barriers to movement and act as mortality sinks for bats. However, understanding flight behaviour over roads has been limited by reliance on human observers to characterise flight patterns. Moreover, both species-specific behaviour and the local environment may alter bat-road interactions. Using paired near-infrared cameras with 3-D tracking analysis, we reconstructed the flight trajectories of barbastelle bats (Barbastella barbastellus) at nine road sites in West Sussex, England. Of the 400 flight paths, 56% occurred at heights that placed bats at risk of vehicle collision, and this risk increased as bats flew closer to canopy cover. Increasing canopy cover was associated with bats flying along rather than across roads, and with faster flight speeds (by 0.14 ms− 1 per 10% increase in canopy cover). Flight behaviour was not influenced by traffic density. These findings suggest that hop-overs, in which trees are planted to provide continuous canopy cover, may create an ecological trap by encouraging B. barbastellus to cross roads at unsafe heights. Our results also highlight the value of 3-D tracking to understand bat behaviour, an approach that could be applied to predict better the impact of road developments on bats and evaluate the effectiveness of mitigation in reducing collision risk.Supplementary InformationThe online version contains supplementary material available at 10.1038/s41598-025-14315-2.

  • Research Article
  • 10.1016/j.vetmic.2025.110573
Monitoring of Bartonella spp. in Iberian bats from Spain.
  • Aug 1, 2025
  • Veterinary microbiology
  • Clara Muñoz-Hernández + 7 more

Monitoring of Bartonella spp. in Iberian bats from Spain.

  • Research Article
  • Cite Count Icon 1
  • 10.3390/f16071189
Tree- and Stand-Scale Roost Selection and Partitioning by Bats Barbastella barbastellus Schreber, 1774 and Pipistrellus pygmaeus Leach, 1825 in a European Lowland Forest
  • Jul 19, 2025
  • Forests
  • Alek Rachwald + 7 more

Our research focused on the roost preferences of two bat species in a forest environment. Throughout the Anthropocene, people have heavily altered the landscape. Forested habitation has declined, with remaining forests becoming fragmented and often deprived of old trees that provide shelter opportunities for bats. TReMs (tree-related microhabitats) are essential for forest-dwelling bat species as they provide an opportunity to roost and shelter. Following an infestation of Ips typographus L., the Białowieża Forest is saturated with dead spruce trees. We investigated roost selection in two forest-dwelling species, Barbastella barbastellus and Pipistrellus pygmaeus. To examine similarities or differences in roost selection between species, we radio-tracked 24 barbastelles and 13 soprano pipistrelles over three breeding seasons. We located a total of 48 barbastelle roosts and 15 pipistrelle roosts, together with the characteristics of the surroundings. We found that barbastelles select roosts almost exclusively in dead spruce trees (43/48), while pipistrelles selected roosts predominantly in live (n = 8) and dead (n = 5) broadleaved trees. Our results show that both bat species have clear differences in roost tree preference. In our study area, with an abundance of exfoliating bark, barbastelles showed a preference for roosting under flaking bark despite the availability of crevices within broadleaved trees. Our findings provide useful insight into forestry practices, highlighting the importance of standing dead trees.

  • Research Article
  • 10.53452/tu2905
The Parcani limestone mine—a new bat (Mammalia: Chiroptera) roosting site of major importance in the Republic of Moldova
  • Jun 30, 2025
  • Theriologia Ukrainica
  • Victoria Nistreanu + 5 more

Limestone mines are the most important underground roosts for bat species in the Republic of Moldova. The aim of the paper was to first study the bat fauna of a newly registered roosting site—the limestone mine near Parcani, Soldanesti district, in the north of Moldova, in order to elucidate the diversity of bats in various phenological periods and to highlight the importance of this roost in preserving chiropteran diversity. Field studies were carried out in 2023–2024 in different seasons, according to standard methods of bat monitoring in underground roosts, in order to reveal the changes in the bat community structure thorough the year. The species diversity was relatively high, with 10 bat species recorded during the study period: Rhinolophus hipposideros, Myotis blythii, M. nattereri, M. daubentonii, M. dasycneme, M. bechsteinii, M. mystacinus, Barbastella barba¬stellus, Plecotus auritus, and Plecotus austriacus. The structure of the bat community was different depending on the season. The highest diversity and largest number of individuals were recorded in the hibernation period, when Myotis blythii was the dominant species representing over 80% of the whole bat community. During the summer season, the diversity and number were the lowest, the dominant species was again M. blythii (55.1%), followed by M. daubentonii (20.4%) and M. dasy¬cneme (14.3%). During the breeding season, the diversity and number increased significantly; M. daubentonii and M. blythii were dominant representing more than 90% of the bat community. The Parcani mine is an important roosting site for many bat species, especially in the breeding and hibernation periods. The registered hibernating colony of M. blythii was the largest among all the known colonies in underground roosts of Moldova. The critically endangered species in Moldova—M. nattereri, M. bech-steinii, and B. arbastellus—were registered in the cold period. For M. nattereri and B. barbastellus, it is the second record locality in the territory of the republic. All species recorded in the Parcani mine are protected at national level in Moldova and in neighbouring countries, while the species Myotis blythii, M. dasycneme, M. bechsteinii, Barbastella barbastellus, and Plecotus austriacus are protected at a European level.

  • Research Article
  • 10.53452/nt1705
Зимові агрегації кажанів у сховищах печерного типу Тернопільського Придністер’я: обліки 2021–2024 років
  • May 15, 2025
  • Novitates Theriologicae
  • Oleksandr Vikyrchak + 3 more

Data on species composition and abundance of bats during hibernation surveyed in the winter of 2024 in 12 cave-type shelters are presented. During these studies, 7 species of bats were found in hibernation. As in previous years, the most numerous species was the lesser horseshoe bat. Another 3 species, the greater mouse-eared bat, Daubenton's bat, and the brown long-eared bat were found in several shelters in small numbers. Three species (Myotis nattereri, M. mystacinus, and Plecotus austriacus) were found in one individual each. Two species (Barbastella barbastellus and Eptesicus serotinus), which were found during the 2021 survey, were not found in 2024. A comparison of the 2021 and 2024 survey data shows a stable abundance of winter aggregations of the thermophilic lesser horseshoe bat and a reduction in the number of cold-loving species, which is likely caused by a stable trend of increasing winter temperatures.

  • Research Article
  • 10.32800/amz.2025.23.0069
Estudi bioacústic sobre la riquesa i l’abundància d’espècies de ratpenats a l'Espai d’Interès Natural de les Obagues del Corb
  • Apr 2, 2025
  • Arxius de Miscel·lània Zoològica
  • S Mañosa

Bio-acoustic study on the richness and abundance of bats in the Natural Interest Site Les Obagues del Corb In summer 2021, ultrasound recordings were carried out at 16 points in the Natural Interest Site (EIN) Les Obagues del Corb (Catalonia) in order to determine the composition of chiropteran communities in this protected natural space. We recorded a total of 11 species or phonic groups, with an average of 5.9 species per point and an average abundance of 20%. The most widespread species were the common pipistrelle Pipistrellus pipistrellus and Kulh's pipistrelle Pipistrellus kuhlii, both almost ubiquitous and locally abundant, followed at some distance by the Soprano pipistrelle/Schreiber's bent-winged bat phonic pair Pipistrellus pygmaeus/Miniopterus schreibersii and the serotine Eptesicus serotinus. The greater horseshoe bat Rhinolophus ferrumequinum and the Bbrbastelle Barbastella barbastellus were also relatively well distributed although their abundance was lower. This work reveals the presence of a rich community of chiropterans in the Les Obagues del Corb, and indicates the need for studies and protection from possible impacts in the future.

  • Research Article
  • 10.1038/s41598-025-95141-4
Predicted climate-induced range shifts and conservation challenges of the western barbastelle bat (Barbastella barbastellus)
  • Mar 28, 2025
  • Scientific Reports
  • Monika Górska + 2 more

Climate change is driving changes in species distributions, particularly affecting specialists and species with limited adaptability. A notable example is the European bat, Barbastella barbastellus, which hibernates in cold, less insulated shelters, is closely associated with old mixed or deciduous forests, and is adapted to feed on moths. Populations of this species are declining and climate change is recognised as a significant contributing factor. In this study, we used species distribution modelling (SDM) techniques to evaluate current and future potential habitat suitability across Europe, incorporating three Shared Socioeconomic Pathway (SSP) scenarios: SSP126, SSP370, and SSP585. Additionally, we assessed the extent of habitat suitability changes under projected climatic conditions within Natura 2000 sites that are aimed at conserving this species. Our projections indicate a possible northward range shift for B. barbastellus, coupled with fragmentation and habitat loss in southern Europe. Furthermore, areas currently protected by the Natura 2000 network may no longer provide suitable conditions for this species in the future. Our study highlights the urgent need for adaptive conservation strategies within networks such as Natura 2000 to protect species increasingly threatened by climate change, with B. barbastellus serving as an example.

  • Research Article
  • Cite Count Icon 1
  • 10.71424/azb76.4.002806
Which Bat Species are Captured by the Long-eared Owl Asio otus (Linnaeus, 1758) (Aves: Strigiformes) in Poland?
  • Dec 10, 2024
  • Acta zoologica bulgarica
  • Grzegorz Lesiński + 3 more

Twenty-two bat individuals were identified to the species level in published and unpublished data based on diet samples of Asio otus from Poland collected between 1989 and 2022. Of the seven species that made up the owl prey, Myotis nattereri (9 cases) and Plecotus auritus (6 cases) were by far the most numerous. The proportion of bats in the vertebrate prey was very low. In the central and north-eastern parts of the country, it was 0.05% between 1980 and 2002 and 0.04% between 2010 and 2024, although the differences were not statistically significant. The hypothesis that bat species associated with foraging in open areas are more abundant in the diet of this owl species, which hunts mainly in agricultural landscapes, could not be confirmed. The vast majority of prey were species that hunt near trees, in forests or over bodies of water. This indicates that the bats are mostly caught in situations other than typical foraging, e.g. near their roosts or resting places of owls. The fact that typical forest bats (Myotis bechsteinii, Barbastella barbastellus) were found in the winter diet of A. otus suggests that they were caught near underground hibernacula, which may be located in open areas away from forests.

  • Research Article
  • 10.71424/azb76.4.002817
Use of Forest Tracks by Commuting or Foraging Bats in a Forest-lake Landscape: A Case Study in the Wigry National Park, Poland
  • Nov 10, 2024
  • Acta zoologica bulgarica
  • Grzegorz Lesiński + 3 more

Studies on the use of forest tracks by bats were carried out in the forest-lake landscape of the Wigry National Park (NE Poland). Netting was performed in July and August 2023 at 14 sites situated near water bodies and at 14 sites remote from waters. In total, 396 individuals belonging to 11 species were caught. Tracks were most often used by Eptesicus serotinus, Myotis daubentonii and Pipistrellus pygmaeus (55% of all caught bats). Eptesicus serotinus and Barbastella barbastellus were more numerous on tracks remote from waters. The group of species known to feed over waters or in riparian habitats (M. daubentonii, N. noctula and Pipistrellus spp.) was represented more frequently on tracks near waters. Forest tracks in the studied landscape are important feeding grounds and corridors for commuting to feeding grounds for many bat species.

  • Research Article
  • Cite Count Icon 2
  • 10.5324/fn.v19i0.6008
Holocene mammals (Insectivora, Rodentia, Chiroptera) of Dollsteinhola Cave, Western Norway
  • Oct 11, 2024
  • Fauna norvegica
  • Joanna Godawa Stormark

The Dolisteinhola cave comprises a sequence of Holocene deposits dating from the late Atlanticum to the first half of the Subboreal period. Climate-sensitive species were found in layers V (Sciurus vulgaris) and layers II - III, IV, IV - V (Apodemus flavicollis). Two humerus of Barbastella barbastellus were also recognised. The faunal assemblage were grouped in three climatic complexes: species which lived both in cold and warm climates; species connected with warmer phases of the climate and species typical for a cold climate.

  • Research Article
  • Cite Count Icon 12
  • 10.1016/j.scitotenv.2024.175661
Re-establishing historic ecosystem links through targeted species reintroduction: Beaver-mediated wetlands support increased bat activity
  • Aug 21, 2024
  • Science of the Total Environment
  • Jack Hooker + 3 more

Despite the global significance of wetlands, conservation strategies often fall short in preserving these ecosystems due to failures in incorporating processes that sustain the ecosystem functioning, hydrological dynamics, ecological processes, and biodiversity of wetlands. Nature-based solutions, such as the reintroduction of beavers, have emerged as effective tools for promoting wetland restoration. Whilst the impact of beavers on wetland restoration is well known, their broader influence on ecosystem health, particularly in modifying habitats for other species, remains inadequately understood. Here we assess the impact that habitat modification through the reintroduction of beavers has on bat populations. There were significantly greater activity levels within beaver-modified wetland habitats for multiple bat species, including higher activity levels of 393 % for Barbastella barbastellus and 313 % for Plecotus spp.. Additionally, we observed positive effects on bat populations in the woodland habitat surrounding beaver-modified wetland for certain taxa. In the face of escalating challenges posed by climate change and habitat loss, addressing biodiversity loss necessitates a shift toward ecosystem-centric mitigation measures. Our study demonstrates that the reintroduction of keystone species like beavers can re-establish historical facilitative links between aquatic and terrestrial food webs, highlighting the importance of such interventions in fostering the resilience and sustainability of entire ecosystems.

  • Research Article
  • Cite Count Icon 1
  • 10.1016/j.foreco.2024.122204
Traits and trails: Guild-specific effects of forest paths on bat activity
  • Aug 17, 2024
  • Forest Ecology and Management
  • D Saracino + 3 more

Traits and trails: Guild-specific effects of forest paths on bat activity

  • Open Access Icon
  • Research Article
  • Cite Count Icon 3
  • 10.1002/ece3.11571
The devil is in the detail: Environmental variables frequently used for habitat suitability modeling lack information for forest-dwelling bats in Germany.
  • Jun 1, 2024
  • Ecology and evolution
  • Lisa Bald + 4 more

In response to the pressing challenges of the ongoing biodiversity crisis, the protection of endangered species and their habitats, as well as the monitoring of invasive species are crucial. Habitat suitability modeling (HSM) is often treated as the silver bullet to address these challenges, commonly relying on generic variables sourced from widely available datasets. However, for species with high habitat requirements, or for modeling the suitability of habitats within the geographic range of a species, variables at a coarse level of detail may fall short. Consequently, there is potential value in considering the incorporation of more targeted data, which may extend beyond readily available land cover and climate datasets. In this study, we investigate the impact of incorporating targeted land cover variables (specifically tree species composition) and vertical structure information (derived from LiDAR data) on HSM outcomes for three forest specialist bat species (Barbastella barbastellus, Myotis bechsteinii, and Plecotus auritus) in Rhineland-Palatinate, Germany, compared to commonly utilized environmental variables, such as generic land-cover classifications (e.g., Corine Land Cover) and climate variables (e.g., Bioclim). The integration of targeted variables enhanced the performance of habitat suitability models for all three bat species. Furthermore, our results showed a high difference in the distribution maps that resulted from using different levels of detail in environmental variables. This underscores the importance of making the effort to generate the appropriate variables, rather than simply relying on commonly used ones, and the necessity of exercising caution when using habitat models as a tool to inform conservation strategies and spatial planning efforts.

  • Research Article
  • 10.71424/azb76.2.002748
Upward Trends in Some Bat Populations from Central Poland between 2000 and 2023 Revealed in Small Hibernacula
  • Apr 24, 2024
  • Acta zoologica bulgarica
  • Grzegorz Lesiński + 1 more

Results of a long-term study conducted in large bat hibernacula of central Europe showed upward trends in populations of many species over the last three decades. This study was aimed to check if similar changes in numbers could be observed in smaller underground winter roosts of bats. Surveys were made in three military shelters near Warsaw between 2000 and 2023 (once a year in February). These roosts hosted relatively small number of hibernating individuals (up to 38). Six bat species were recorded: Myotis nattereri, Myotis daubentonii, Eptesicus serotinus, Plecotus auritus, P. austriacus and Barbastella barbastellus. The species preferring cold sites for hibernation (P. auritus and B. barbastellus) predominated in the studied roosts, representing together over 86% of the recorded individuals. The numbers of the two most abundant species and of all bats showed statistically significant upward trends.

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  • Research Article
  • Cite Count Icon 3
  • 10.3390/su16062337
In the Spotlight: Bat Activity Shifts in Response to Intense Lighting of a Large Railway Construction Site
  • Mar 12, 2024
  • Sustainability
  • Fabrizio Gili + 3 more

Artificial light at night (ALAN) is a major form of anthropogenic pollution, disrupting nocturnal wildlife behaviour and ecosystem function. Large construction sites are typically located at the intersection of urban and natural areas, introducing intense lighting into previously dark natural habitats. This study examines the responses of bats to intense nighttime lighting at a major construction site associated with a linear transport infrastructure (LTI) project. We used passive acoustic monitoring to record bat activity and species richness at the construction site and in adjacent urban and natural areas with different lighting levels. Generalist species, such as Pipistrellus kuhlii and Nyctalus leisleri, were attracted to illuminated areas, likely due to increased prey availability. Conversely, sensitive species, such as those from the Myotis and Plecotus genera, along with Barbastella barbastellus, avoided illuminated areas, particularly the construction site. Species richness was significantly lower at the construction site compared to other environments, reflecting ALAN’s barrier effect on movements and habitat accessibility for more sensitive species. The findings highlight the ecological impacts of introducing ALAN to previously unlit natural environments, underscoring the urgent need for implementing ALAN mitigation strategies in urban planning and construction practices to protect biodiversity in urbanising areas.

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  • Research Article
  • Cite Count Icon 1
  • 10.3390/app14052125
Bat Hibernation: In Groups or Individually?
  • Mar 4, 2024
  • Applied Sciences
  • Grzegorz Kłys + 2 more

This study focuses on the hibernation behavior of the western barbastelle bat (Barbastella barbastellus) in underground systems in Poland from 2006 to 2011, specifically during the peak hibernation months of December to February. The impact of climate parameters, namely temperature (T), humidity (Rh), and air flow velocity (v), on the clustering behavior of bats during hibernation was investigated. The climate parameters varied within specific ranges: T fluctuated between 6.0 and 12.4 °C, Rh ranged from 56.4 to 91.8%, and v varied from 0.01 to 1.17 m/s. The quantile linear regression method for statistical analysis of the results was employed. This study found that certain combinations of climate parameters influenced the grouping behavior of bats during hibernation. The model structural parameters revealed the following relationships: 1. An increase in the product of T and v led to an increase in the bats’ group size. 2. For pairs of variables such as T and Rh, and Rh and v, an increase in their product resulted in a decrease in the bats’ group size. 3. When considering the product of T, Rh, and v, a decrease in the bats’ group size was also observed.

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  • Research Article
  • Cite Count Icon 8
  • 10.3390/f15020337
Exceptionally Uniform Bat Assemblages across Different Forest Habitats Are Dominated by Single Hyperabundant Generalist Species
  • Feb 8, 2024
  • Forests
  • Mateusz Ciechanowski + 6 more

Woodland bat assemblages are usually structured in a space according to the distance from the ground, water, and obstacles, features that often define chiropteran hunting tactics. Consequently, the bat species composition differs strongly among various habitats, even within the same forest patch. However, when conducting local bat surveys in Wolin National Park (WPN), we revealed an unexpected uniformity in the qualitative and quantitative structure of bat assemblages, based on mist netting and ultrasound recording. In total, 10 vespertilionid species were detected. Across all methods and sampled habitats, a single species, Pipistrellus pygmaeus, predominated, while no Barbastella barbastellus, an old forest specialist, were detected, despite the abundance of their preferred daily roosts. We also reviewed the literature for mist-netted bat samples in four different habitats in lowland Polish forests. The samples usually clustered based on habitats, and the same habitat classes often clustered very closely despite representing geographically distant forests. The exception was WPN, where all four habitat classes formed a tightly packed cluster. We hypothesize that P. pygmaeus might act as a hyperabundant native species, a successful generalist that reduces the contribution of more specialized taxa in the assemblage. It probably benefits from both forest renaturation and anthropogenic cross-boundary subsidy.

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