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In late summer, migratory bats of the temperate zone face the challenge of accomplishing two energy-demanding tasks almost at the same time: migration and mating. Both require information and involve search efforts, such as localizing prey or finding potential mates. In non-migrating bat species, playback studies showed that listening to vocalizations of other bats, both con-and heterospecifics, may help a recipient bat to find foraging patches and mating sites. However, we are still unaware of the degree to which migrating bats depend on con-or heterospecific vocalizations for identifying potential feeding or mating opportunities during nightly transit flights. Here, we investigated the vocal responses of Nathusius’ pipistrelle bats, Pipistrellus nathusii, to simulated feeding and courtship aggregations at a coastal migration corridor. We presented migrating bats either feeding buzzes or courtship calls of their own or a heterospecific migratory species, the common noctule, Nyctalus noctula. We expected that during migratory transit flights, simulated feeding opportunities would be particularly attractive to bats, as well as simulated mating opportunities which may indicate suitable roosts for a stopover. However, we found that when compared to the natural silence of both pre-and post-playback phases, bats called indifferently during the playback of conspecific feeding sounds, whereas P. nathusii echolocation call activity increased during simulated feeding of N. noctula. In contrast, the call activity of P. nathusii decreased during the playback of conspecific courtship calls, while no response could be detected when heterospecific call types were broadcasted. Our results suggest that while on migratory transits, P. nathusii circumnavigate conspecific mating aggregations, possibly to save time or to reduce the risks associated with social interactions where aggression due to territoriality might be expected. This avoidance behavior could be a result of optimization strategies by P. nathusii when performing long-distance migratory flights, and it could also explain the lack of a response to simulated conspecific feeding. However, the observed increase of activity in response to simulated feeding of N. noctula, suggests that P. nathusii individuals may be eavesdropping on other aerial hawking insectivorous species during migration, especially if these occupy a slightly different foraging niche., Zweitveröffentlichungen der Universität Potsdam : Mathematisch-Naturwissenschaftliche Reihe; 1306

Molecular analyses of feces are widely used to study the feeding ecology of bats. However, little is known about how detectability of prey DNA from bat feces is influenced by gut retention time, mass of prey eaten, or prey identity, which hampers the interpretation of field data. Here, we address these knowledge gaps by conducting a feeding experiment in which different insect species and meal sizes were offered to two bat species, Myotis daubentonii and Pipistrellus nathusii. The feeding regime consisted of three main and three side prey species fed in different masses, whereupon feces were collected over 72 h and examined for prey DNA using a species‐specific multiplex PCR system. DNA of all three main prey species was detectable from feces produced within 1 h after consumption of a meal. Prey detectability decreased over the following 10–20 h, and after 40 h only spurious detections occurred. For both prey and bats, species identity affected detectability: for example, mealworms were detected for a longer time post‐feeding compared to wax moths/house flies, and the latter were detected for a shorter time in M. daubentonii than in P. nathusii. While DNA from all three side prey species was detectable, signals were weak and detections limited to the first 20 h after consumption. These findings indicate that detectability of prey DNA from bat feces is foremost affected by the mass of prey consumed and time post‐feeding, whereas it does not strongly differ between prey species. To increase the detection of prey ingested in low mass, we suggest collecting fecal samples soon after bats return from foraging, as detectability is highest right after feeding occurs, and detections largely depict the previous foraging bout (with detection windows usually

The results of fauna study of National Park Narochansky (Minsk region) during the field seasons 2019-2020 are presented. We performed bat captures using mist nets and also surveyed settlements with an ultrasonic heterodyne detector. 177 specimens of 9 species were captured, specifically: Eptesicus nilssonii (Keyserling et Blasius, 1839), Myotis brandtii (Eversmann, 1845), M. dasycneme (Boie, 1825), M. daubentonii (Kuhl, 1817), Nyctalus noctula (Schreber, 1774), Plecotus auritus (Linnaeus, 1758), Pipistrellus nathusii (Keyserling et Blasius, 1839), P. pygmaeus (Leach, 1825), Vespertilio murinus (Linnaeus, 1758). 11 settlements of bats were discovered and mapped within the study area. According to the mist net captures, the dominant species is P. nathusii, with subdominant M. daubentoniid; andM. brandtii with P. auritus as the rarest species. Other species had represented ratios of 2.7 to 9.5 % in the nets captures. Species E. nilssonii andM. brandtii are only represented by male specimens. We also report the first registered discovery of Myotis brandtii on the territory of National Park Narochansky, which is also the most northern registration of that species in Belarus. In comparison to the previous studies, we have confirmed the presence of several species with previously questionable local status: P. auritus, M. brandtii, M. daubentonii and V. murinus.

Being diverse and widely distributed globally, bats are a known reservoir of a series of emerging zoonotic viruses. We studied fecal viromes of twenty-six bats captured in 2015 in the Moscow Region and found 13 of 26 (50%) samples to be coronavirus positive. Of P. nathusii (the Nathusius’ pipistrelle), 3 of 6 samples were carriers of a novel MERS-related betacoronavirus. We sequenced and assembled the complete genome of this betacoronavirus and named it MOW-BatCoV strain 15-22. Whole genome phylogenetic analysis suggests that MOW-BatCoV/15-22 falls into a distinct subclade closely related to human and camel MERS-CoV. Unexpectedly, the phylogenetic analysis of the novel MOW-BatCoV/15-22 spike gene showed the closest similarity to CoVs from Erinaceus europaeus (European hedgehog). We suppose MOW-BatCoV could have arisen as a result of recombination between ancestral viruses of bats and hedgehogs. Molecular docking analysis of MOW-BatCoV/15-22 spike glycoprotein binding to DPP4 receptors of different mammals predicted the highest binding ability with DPP4 of the Myotis brandtii bat (docking score −320.15) and the E. europaeus (docking score –294.51). Hedgehogs are widely kept as pets and are commonly found in areas of human habitation. As this novel bat-CoV is likely capable of infecting hedgehogs, we suggest hedgehogs can act as intermediate hosts between bats and humans for other bat-CoVs.

Migration is an essential process in animals’ lives, which they use to avoid adverse weather conditions and resource scarcity. Migratory animals are considered particularly vulnerable to anthropogenic changes to their environment because they require a specific temporal and spatial progression of appropriate habitats for foraging and reproduction, and a refuge from harsh environmental conditions. Anthropogenic activities that change the environment such as habitat loss, habitat fragmentation, and climate change have had a global effect on migratory species resulting in widespread population declines. Climate change is a particularly serious threat for migratory species as it can have impacts across all the habitats within which migratory animals must move. Efforts to mitigate climate change may also have adverse effects on migratory species. The trade-off between renewable energy use and wildlife conservation is not always straightforward. Changes in energy systems have the potential to alter the functioning of ecosystems and wildlife populations by affecting species' access to resources, habitat availability, and connectivity. The use of wind energy poses a particular threat for many migratory species. Studies have shown that wind energy farms pose a mortality risk to flying migratory individuals both inland and on the coast, and new concerns are being raised around offshore establishments. Conservation of migratory species in the face of such threats is of vital importance and will likely require the protection of the multiple habitats that are used during the migratory journey. However, there is often limited knowledge about the migratory pathways and stopover sites used by migratory species, posing a major challenge for effective conservation. Stable isotope analysis can be used to trace terrestrial migratory routes and identify migratory origins. The development of such techniques to determine the corridors and pathways migratory animals are using to move between habitats has assisted us in further honing conservation efforts. Stable isotope analysis allows the use of a small number of samples, which can be collected in one sampling event, to detect environmental tracers that are related to the spatial-temporal movement of animals. The application of stable isotopes in conservation biology is growing rapidly and shows great promise for the conservation of endangered species. Bats are one of the most taxonomically diverse groups of mammals, however, only a few species are known to migrate long distances. These species have evolved a combination of physiological and morphological traits to allow long-distance migration. The European Nathusius’ pipistrelle (Pipistrellus nathusii) is one of the most well-studied migratory bat species worldwide. The species is known to maintain a long-distance migration with both coastal and offshore pathways. Recent studies have shown that Nathusius’ pipistrelles have been increasing their geographical range, even reaching 60° N in latitude. Their long-distance migratory behaviour and the increase in European wind farms make them highly vulnerable to environmental changes. Considering their broad European range, high vulnerability to environmental changes, and the current need to protect migratory species, Nathusius’ pipistrelles are an interesting model species in which to apply stable isotope analysis to investigate migration patterns. In this thesis, I use stable isotope analysis to identify the northern migratory corridors of Nathusius’ pipistrelles (chapter 1) and differentiate the mortality risk posed by wind turbines on different demographic groups during migration (chapter 2). In chapter 1, I use a dual-isotope approach (δ2H and 87Sr/86Sr) to determine the origin of bats found on three islands in the north of Germany. Although δ2H analysis suggested a possible Fennoscandian origin, 87Sr/86Sr analysis refuted this possibility and proposed they would be originating in Russia and the Baltic states. In chapter 2, I use a comprehensive dataset of fur samples from carcasses collected beneath wind turbines and living individuals across Germany to assess the vulnerability of different demographic groups to wind turbine mortality. Compared to adults, juveniles were more vulnerable at low wind turbine densities; this effect was minimised at high density, with both ages equally affected. In addition, I found more females and regional migrants in both living and carcass populations. Overall, this dissertation demonstrates the importance of stable isotope analysis in wildlife research and provides a practical example of how it can help inform species conservation.

Simple Summary Migratory bats regularly fly over the North Sea, where the number of offshore wind farms will increase rapidly in the next decades. Information is urgently needed on the timing and the conditions bats can be expected offshore, since windfarms can cause fatalities amongst bats. We therefore collected acoustic data on the presence of bats at four nearshore locations at sea between 2012 and 2016. Modelling the occurrence of Nathusius’ pipistrelle for 480 nights in autumn showed that its migration is strongest in early September, with east-northeasterly tailwinds, low wind speeds, and relatively high temperatures. The species’ migration did not show a strong relationship with other factors, i.e., moon phase, cloud cover, atmospheric pressure, rain, and visibility. Our results provide valuable input to policy-makers to prescribe mitigation measures to reduce bat fatalities in offshore wind farms. Abstract Bats regularly migrate over the North Sea, but information on the environmental conditions when this occurs is scarce. Detailed information is urgently needed on the conditions under which bats can be expected offshore, as the number of offshore windfarms that can cause fatalities amongst bats in the North Sea is increasing rapidly. We performed ultrasonic acoustic monitoring at multiple nearshore locations at sea between 2012 and 2016 for, in total, 480 monitoring nights. We modelled the offshore occurrence of Nathusius’ pipistrelle in autumn as a function of weather conditions, seasonality, and the lunar cycle using a generalized additive mixed model (GAMM). We investigated which covariates are important using backward selection based on a likelihood ratio test. Our model showed that important explanatory variables for the offshore occurrence of Nathusius’ pipistrelle are seasonality (night in year), wind speed, wind direction, and temperature. The species’ migration is strongest in early September, with east-northeasterly tailwinds, wind speeds < 5 m/s, and temperatures > 15 °C. Lunar cycle, cloud cover, atmospheric pressure, atmospheric pressure change, rain, and visibility were excluded during the model selection. These results provide valuable input to reduce bat fatalities in offshore wind farms by taking mitigation measures.

The discovery of a significant number of viral pathogens in bat tissue samples and excrement point to a potential prominent role of chiropters in the maintenance and spread of human and animal diseases. It also indicates the potential sensitivity of bat cells to a broad spectrum of viruses. The migratory pipistrelle bat, Pipistrellus nathusii, inhabits northeastern Europe and typically migrates to the southwest. Our study revealed the sensitivity (susceptibility) of a diploid cell line, derived from the kidney of P. nathusii to several transmissible animal disease causative agents such as Epizootic hemorrhagic disease, Akabane disease, Vesicular stomatitis virus and Peste des petit ruminants. High sensitivity of the P. nathusii kidney diploid cell line to viruses from various taxonomic groups allows them to be recommend for extensive use in virological studies.

A male Pipistrellus nathusii ringed in Pape Natural Park (S Latvia) in August 2015 was recovered recently dead in Pitillas’ Lagoon Natural Reserve (N Spain) in March 2017. At 2224 km in SSW direction, this is the first documented bat migration between these countries and worldwide the longest migration record of a bat. We also report other observations of this species in autumn in Northern Spain, suggesting that the Iberian Peninsula may be an important wintering area for Nathusius’ pipistrelles. Conservation measures should be agreed on by countries along the migration routes to improve the protection of this species.

Modern climate change and environmental degradation have become planetary. The impact of these factors on complex biological systems must be monitored using bioindicator taxa, including bats. Bioindicative potential of bats is assessed, a prerequisite of which are their widespread distribution in the world, high specialization, availability of study of population trends, and effects of short- and long-term impacts. An effective use of this potential can be fulfilled through the formation of a global monitoring network of bat populations. These populations are declining due to the negative impact of a number of factors, including heavy metals in the world. The content of heavy metals (Pb, Cr, Zn and Cu) is determined and the patterns of their quantitative representation in the wool of five bat species in nature conservation and urban areas of the west of Ukraine are characterized. These species are as follows: Eptesicus serotinus (Schreber, 1774), Myotis daubentonii (Kuhl, 18170), Plecotus auritus (Linnaeus, 1758), Pipistrellus kuhlii (Kuhl, 1819) and Pipistrellus nathusii (Keyserling et Blasius, 1839). The results of study of all bat species observed have shown the highest concentrations for zinc, the lowest – for lead and copper. The concentration of lead in the wool of M. daubentonii and P. nathusii was significantly lower than in other bat bodied indicating the remoteness of the feed and biotopic preferences of these species from anthropogenic influence. The study has revealed that the concentrations of metals in bat wool on the territory of Shatsky National Park and in Lviv show a certain hierarchical sequence – Zn > Cu > Pb or Zn > Pb > Cu, depending on the ecological and biological features of the species. The values of Cr were found to be the most variable for all bats and varied widely. The metal content was determined in the muscles of Ept serotinus, which died in an accident in Lviv and Rivne in 2014. The content of chromium and zinc in muscle was much lower compared to the content of these metals in the bats' wool. The work evaluates the importance of the use of bats as biological indicators of heavy metal contamination and points the expediency of using in bat species environmental monitoring the method of determining the content of heavy metals in wool as non-invasive and highly informative.

Large numbers of bats are killed by wind turbines globally, yet the specific demographic consequences of wind turbine mortality are still unclear. In this study, we compared characteristics of Nathusius' pipistrelles (Pipistrellus nathusii) killed at wind turbines (N = 119) to those observed within the live population (N = 524) during the summer migration period in Germany. We used generalized linear mixed-effects modeling to identify demographic groups most vulnerable to wind turbine mortality, including sex (female or male), age (adult or juvenile), and geographic origin (regional or long-distance migrant; depicted by fur stable hydrogen isotope ratios). Juveniles contributed with a higher proportion of carcasses at wind turbines than expected given their frequency in the live population suggesting that juvenile bats may be particularly vulnerable to wind turbine mortality. This effect varied with wind turbine density. Specifically, at low wind turbine densities, representing mostly inland areas with water bodies and forests where Nathusius' pipistrelles breed, juveniles were found more often dead beneath turbines than expected based on their abundance in the live population. At high wind turbine densities, representing mostly coastal areas where Nathusius' pipistrelles migrate, adults and juveniles were equally vulnerable. We found no evidence of increased vulnerability to wind turbines in either sex, yet we observed a higher proportion of females than males among both carcasses and the live population, which may reflect a female bias in the live population most likely caused by females migrating from their northeastern breeding areas migrating into Germany. A high mortality of females is conservation concern for this migratory bat species because it affects the annual reproduction rate of populations. A distant origin did not influence the likelihood of getting killed at wind turbines. A disproportionately high vulnerability of juveniles to wind turbine mortality may reduce juvenile recruitment, which may limit the resilience of Nathusius' pipistrelles to environmental stressors such as climate change or habitat loss. Schemes to mitigate wind turbine mortality, such as elevated cut-in speeds, should be implemented throughout Europe to prevent population declines of Nathusius' pipistrelles and other migratory bats.

Highly mobile species are considered to be the first to respond to climate change by transforming their ranges of distribution. There is evidence suggesting that Pipistrellus nathusii, a species capable of long-distance migration, is expanding both its reproduction and overwintering ranges to the North. We recorded the echolocation calls of bats at 16 sites in South-Western Finland on two consecutive winters, and detected calls of P. nathusii at one of the sites throughout the second winter. To our knowledge, this is the northernmost record of an overwintering P. nathusii, and contributes to evidence that the species is already responding to climate change.

A male Pipistrellus nathusii ringed in Pape Natural Park (S Latvia) in August 2015 was recovered recently dead in Pitillas’ Lagoon Natural Reserve (N Spain) in March 2017. At 2224 km in SSW direction, this is the first documented bat migration between these countries and worldwide the longest migration record of a bat. We also report other observations of this species in autumn in Northern Spain, suggesting that the Iberian Peninsula may be an important wintering area for Nathusius’ pipistrelles. Conservation measures should be agreed on by countries along the migration routes to improve the protection of this species. nonPeerReviewed

The exact anatomical location for an iron particle-based magnetic sense remains enigmatic in vertebrates. For mammals, findings from a cornea anaesthesia experiment in mole rats suggest that it carries the primary sensors for magnetoreception. Yet, this has never been tested in a free-ranging mammal. Here, we investigated whether intact corneal sensation is crucial for navigation in migrating Nathusius’ bats, Pipistrellus nathusii, translocated from their migratory corridor. We found that bats treated with corneal anaesthesia in both eyes flew in random directions after translocation and release, contrasting bats with a single eye treated, and the control group, which both oriented in the seasonally appropriate direction. Using a Y-maze test, we confirmed that light detection remained unaffected by topical anaesthesia. Therefore our results suggest the cornea as a possible site of magnetoreception in bats, although other conceivable effects of the anaesthetic are also explored. Furthermore, we demonstrate that the corneal based sense is of bilateral nature but can function in a single eye if necessary., Lindecke and colleagues experimentally investigate the effect of corneal anesthesia on bat migratory orientation. Their results suggest that the cornea is a promising organ for the search of biological magnetic compass sensors, and also indicate that a single functional cornea is sufficient for migratory navigation.

Although labeled as environmentally friendly, wind power can have negative impacts on the environment, such as habitat destruction or wildlife fatalities. Considering the distribution and migratory characteristics of European bats, the negative effects of wind power should be addressed on an appropriate scale. This review summarizes the current state of knowledge on interactions between wind farms and bats in Europe, and compares it with the situation in the countries of the European boreal biogeographic region. We analyzed data from papers published in international and national scientific journals, focusing on studies conducted in Europe. The issue of the impacts wind power has on bats is clearly overlooked in most of the countries of the European boreal region, with low volumes of research available on the topic. This is probably due to fewer wind farms in the area, making this recent issue a less-prioritized topic. However, the Baltic Sea, and the countries surrounding it, are of extreme importance with regards to bat migration, especially for the Pipistrellus nathusii. Therefore, more research on wind power and bats is needed in this region, as well as more cooperation between all the stakeholders.

he local landscape Feofaniya is located on the outskirts of the Kyiv and represents communities of native oak wood with high density of ancient trees and hornbeam forest, steppe meadow, culture phytocenoses, cascade of ponds, stream. For the period 2012-2018 11 species of fish, 8 amphibians, 7 reptiles, 111 birds, 29 mammals were identified. Of these, 82.0 % of species are protected internationally, 8.9 % at the state level of the Ukraine, 12.6 % are regionally rare. In 81% of animal species, the reproductive site passes through the tract, and in half of them the whole life cycle. 19 % of the species composition of birds (Aves) are sedentary, 58 % arrive for nesting, 12 % for feeding, 9 % for wintering, 2 % for migratory (Regulus ignicapillus, Nucifraga caryocatactes). 82.8 % of mammals (Mammalia) are sedentary, 10.3 % are migratory (Pipistrellus nathusii, Pipistrellus pygmaeu, Nyctalus leisleri), 6.9 % with unexplained status (Neovison vison, Capreolus capreolus). The local landscape has 12 alien species. Carassius gibelio, Ctenopharyngodon idella, Hypophthalmichthys molitrix released into ponds with the purpose of stocking. Trachemys seripta elegans bring vacationers to the ponds. Pseudorasbora parva, Perccottus glenii, Streptopelia decaocto, Dendrocopos syriacus, Phoenicurus ochruros, Serinus serinus, Ondatra zibet, Pipistrellus kuhlii, Neovison vison penetrated the local landscape independently from neighboring biotopes. Thanks to biotechnical measures aimed at improving the living conditions of animals (protection of natural nests and the creation of artificial ones), hanging feeders in the winter, hanging shelters for daytime and wintering bats. The negative impact on the biotopes of Feofaniya from the side of recreants is somewhat leveled at the expense of educational work with vacationers. The most relevant propaganda work is aimed at educating correct behavior in nature, explaining the need to protect all plant and animal species, their biotopes, and explaining the consequences of disturbing the balance of sustainable development of ecosystems.

The amino acid spectrum of blood plasma in three bat species (Myotis dasycneme, Pipistrellus nathusii, Vespertilio murinus) inhabiting the Urals has been studied for the first time. The bats were trapped in the zone of their high abundance in Chelyabinsk oblast (2013–2014). Free amino acids were determined by liquid ion exchange chromatography (a total of 384 determinations). It has been shown that the plasma amino acid spectrum consists of 22 amino acids in subadult bats of all three species, but there are species-specific differences in their concentrations. The total amino-acid pool concentration in migratory P. nathusii and V. murinus exceeds that in resident M. dasycneme by factors of 2.9 and 1.8, respectively. Migratory species are characterized by a high concentration of plasma arginine: it is six times higher in V. murinus than in M. dasycneme, and in P. nathusii arginine accounts for 25.4% of the amino acid pool. The group of glucogenic amino acids is prevalent in the blood plasma of migratory species (75% in V. murinus and 79% in P. nathusii), while in M. dasycneme the total proportion of lysine, glycine, and glutamic acid is 2.3 times lower than in P. nathusii and 1.7 lower than in V. murinus (p < 0.05). These results provide evidence for significant differences in the contents of free blood plasma amino acids between migratory and resident bat species.

The occurrence of bats in mountains at high altitudes above 2300 m has not been investigated in the Alps until recently. In other European mountains, only a few studies on bats in high regions e.g., in the Pyrenees or in Switzerland at the Col de Bretolet had been conducted. Here we report a study carried out between 2012 and 2016 at several high-alpine sites at altitudes ranging from 2250 m up to 2761 m in the mountains of the Hohe Tauern, Salzburg, Austria. In spite of extreme climatic conditions, a surprisingly high bat activity and diversity of bat species was recorded. Eleven bat species were detected by different methods. Species identified by echolocation calls were: Barbastella barbastellus, Eptesicus nilssonii, Nyctalus noctula, N. leisleri, Pipistrellus pipistrellus, P. pygmaeus, Myotis daubentonii, and Vespertilio murinus. Additional findings were the species pairs Myotis mystacinus/M. brandtii and Pipistrellus nathusii/P. kuhlii as well as the genus Plecotus. The presence of most of these species was confirmed by capturing individuals with mist nets. Permanent monitoring of ultrasound calls was undertaken throughout the active period of bats at the site located at 2315 m above sea level. A total of over 30,000 series of bat calls were recorded and the activity period lasted from the end of March to the beginning of November. Long-distance migratory bat species were recorded in spring under wintry conditions and much more prominently from August to October. Presumably these individuals were crossing the Alps on their way between summer and winter roosts. Sedentary bats used the high alpine zone for foraging during summer. At these altitudes bats were active at high wind speeds of up to 13.9 m/s as well as at low temperatures around freezing point and down to -5.8°C.

Introduction Bats are threatened by agricultural intensification, and although bat ecology in agricultural landscapes is in the focus of current research, the effects of interacting spatiotemporal factors on species-specific bat activity above farmland remain understudied. Our aim was to identify spatiotemporal factors and their interactions relevant for the activity of bat species above conventionally managed arable fields. Methods We repeatedly monitored relative bat activity above open arable fields in Germany using acoustic monitoring. We used site-related biotic and abiotic factors and landscape characteristics across five spatial scales, their combinations, and interactions to identify those factors which best explain variation in bat activity. Results Numerous interactions between landscape characteristics and the insect abundance affected bat activity above fields. For instance, Pipistrellus pipistrellus became more active with increasing insect abundance, but only above fields with a low proportion of woody vegetation cover in the surroundings. Additionally, the level of bat activity in summer depended on landscape characteristics. For example, the activity of Pipistrellus nathusii was relatively low in summer above fields that were surrounded by vegetation patches with a high degree of edge complexity (e.g., hedgerow). However, the activity remained at a relatively high level and did not differ between seasons above fields that were surrounded by vegetation patches with a low degree of edge complexity (e.g., roundly shaped forest patch). Conclusions Our results revealed that landscape characteristics and their interactions with insect abundance affected bat activity above conventionally managed fields and highlighted the opportunistic foraging behavior of bats. To improve the conditions for bats in agricultural landscapes, we recommend re-establishing landscape heterogeneity to protect aquatic habitats and to increase arthropod availability.

22. Nathusius’s Pipistrelle Pipistrellus nathusii French: Pipistrelle de Nathusius / German: Rauhautfledermaus / Spanish: Pipistrela de Nathusius Taxonomy. Vesperugo nathusii Keyserling & Blasius, 1839, Berlin, Germany. Pipastrellus nathusii appears to be close to the P. kuhlii complex, which includes paraphyletic P. kuhliiand P. maderensis. Monotypic. Distribution. Most of Europe (from Ireland and Iberian Peninsula E to Russia and W Kazakhstan), Turkey, and Caucasus; also on Mediterranean Is such as Crete or Mallorca. Descriptive notes. Head-body 44-54 mm, tail 30-40 mm, ear 10-5-16 mm, forearm 32-2-37-1 mm; weight 6-15-5 g. Fur color of Nathusius’s Pipistrelle changes seasonally, being dark brown or gray brown in winter and reddish in summer. Fur is long and uniformly colored, but venteris slightly paler or more yellowish than dorsum. Proximal one-half of uropatagium is covered by fur. Bare skin is dark brown, snout is short and rounded, and ears are triangular and short, with slender and rounded tragus. Compared with other pipistrelle species, wings of Nathusius’s Pipistrelle are quite long (wingspan 230-250 mm), perfectly adapted for long flights and its migratory behavior. Although not always reliable, a particular wing venation pattern is used as a diagnostic character (cell from elbow to fifth finger clear ly divided). I* is bicuspid, I’ is longer than second cusp of I, canines are considerably large, P? is small but visible above gum, and lower molars are nyctalodont. Chromosomal complement has 2n = 44, FNa = 50, and FN = 54. Habitat. [Lowland to alpine habitats from sea level up to elevations of ¢. 2200 m in the Alps (probably single vagrant individuals). Habitats selected by Nathusius’s Pipistrelle include woodlands, forest edges, wetlands, open areas, and grasslands. In terms offorests, it occurs in deciduous, lowland, riparian, and coniferous forests and urban gardens with isolated trees. It tends to select those habitats surrounding ponds, lakes, or water bodies; thus,it is common in wetlands, swamps, and floodplains, especially during migration. Food and Feeding. Nathusius’s Pipistrelle reportedly eats Diptera and Lepidoptera, especially non-biting midges, mosquitoes, and black flies. Despite being slower than its congeners, it hunts insects (aerial hawker) at 3-20 m aboveground, commonly on forest edges, linear structures, and forest paths and over water bodies (from small ponds to large lakes), and occasionally around streetlamps. It has also been recorded foraging over sea up to 1-3 km from the coast. Dietary composition varies between maternity, swarming, and migration periods. In Latvia, during migration, Nathusius’s Pipistrelles seem to use a “fly-and-forage” feeding strategy, which consists in stopping several times during migration to feed and then continue the flight. Breeding. Mating of Nathusius’s Pipistrelles occurs in June-November, including when they roost in nursery colonies (June-August), during migration (August-November), and at their destination at swarming and hibernation sites (November). Males can be resident in swarming areas (which they defend from other males), visitors to specific mating locations, or sporadic visitors that try to mate opportunistically. Maternity colonies are only found in northern and Eastern Europe (Britain, eastern Germany, Baltic States, Belarus, Ukraine, and Russia). During breeding season, Nathusius’s Pipistrelles congregate in small colonies of some tens to a few hundred individuals, roosting in tree holes, bat boxes, and buildings, usually in lowland forests and wetlands. They tend to give birth to twins in June and leave colonies by the end ofJuly. Young females become sexually active during their first year of life. Nathusius’s Pipistrelles can live up to 13 years. Activity patterns. Nathusius’s Pipistrelle emerges at dusk to hunt flying insects. Unlike other pipistrelles, its flight is not that erratic, being straighter and more predictable. It roosts in hollow trees, bird and bat boxes, rock crevices, expansion joints in bridges, houses, and a wide range of different buildings, timber claddings, or wooden churches. In winter,it tends to roost in rock crevices, cliffs, caves, mines, and buildings in urban areas (most likely benefiting from urban heat islands). It hibernates from mid-winter to early spring. Its echolocation is very similar to that of Kuhl’s Pipistrelle and has high FM sweeps with CF cue that resembles a hockey stick-shaped call when represented on sonograms. Peak frequencies are ¢.38—43 kHz. Echolocation calls of Nathusius’s Pipistrelle can be confused with those of Kuhl’s Pipistrelle (PF. kuhliz) and Savi’s Pipistrelle (Hypsugo savii). Calls are variable; in open spaces, FM component can disappear or be lower in frequency, making identification difficult; and in cluttered environments, pulses become highly modulated resembling Myotis echolocation. Pulses last up to 12 milliseconds. Socials calls are very distinct from those of Kuhl’s Pipistrelle and are typically used to reliably identify recorded free-flying bat calls. Movements, Home range and Social organization. Nathusius’s Pipistrelle is one of the few long-distance migratory bat species in Europe, for which migration patterns of more than 1900 km have been reported. In Europe, it tends to breed in north-eastern countries in summer and migrate toward south-western countries to hibernate. In these regions,it positively selects natural wetlands for foraging and avoids agricultural lands and pastures. During migration, it presumably follows coastlines (it has been captured or recorded in offshore platforms and fishing boats), but it has also been captured over high mountains or following waterways. It has also been recorded migrating from continental Europe to Great Britain. Females start migration before males. Migration extends from August to October or November. Nathusius’s Pipistrelles can fly up to 29-48 km/night and only occasionally up to 80 km /night. During the maternity period, foraging areas are relatively close to maternity colonies (less than 6-5 km). Maternity colonies usually share space with the Common Pipistrelle, Brandt's Myotis (Myotis brandtii), and the Pond Myotis (M. dasycneme). Status and Conservation. Classified as Least Concern on The IUCN Red List. Although not very common, Nathusius’s Pipistrelle is quite widespread. Although it has been more common in northern countries,it is now more frequently reported in the south, and during the last few decades, it has been reported or established in the Netherlands, UK, Switzerland, Italy, and Spain. Because it is one of the few long-distance migratory bat species in Europe, modification or fragmentation of habitat and commuting routes might compromise its movements. Roost disturbance and destruction, loss of mature trees in wetlands, and direct persecution are recognized as direct threats. Bibliography. Arslan & Zima (2014), Benda, Georgiakakis et al. (2008), Bogdanowicz (1999a), Boshamer & Bekker (2008), Brosset (1990), Dietz & Kiefer (2016), Flaguer, Puig-Montserrat et al. (2009), Flaquer, Ruiz-Jarillo et al. (2005), Harris &Yalden (2008), Jahelkova et al. (2008), Kriiger et al. (2014), Limpens & Schulte (2000), Lundy et al. (2010), Pacifici et al. (2013), Paunovi¢ & Juste (2016), Pétersons (2004), Rodriguez-Munoz et al. (1993), Russ & Racey (2007), Russ, Hutson et al. (2001), Russ, O'Neill & Montgomery (1998), Sachanowicz et al. (2019), Schober & Grimmberger (1998), Speakman et al. (1991), Suba et al. (2012), Trujillo & Garcia (2009), Vierhaus (2004), Voigt et al. (2012)., Published as part of Don E. Wilson & Russell A. Mittermeier, 2019, Vespertilionidae, pp. 716-981 in Handbook of the Mammals of the World – Volume 9 Bats, Barcelona :Lynx Edicions on page 773, DOI: 10.5281/zenodo.6397752

This atlas is based on information in museum collections, literature, databases and unpublished data. In the last 150 years, the number of bat species in Finland increased from six to thirteen. Of these, five are common and regularly breeding (Eptesicus nilssonii, Myotis brandtii, Myotis daubentonii, Myotis mystacinus, Plecotus auritus), and eight rare (Eptesicus serotinus, Myotis dasycneme, Myotis nattereri, Nyctalus noctula, Pipistrellus nathusii, Pipistrellus pipistrellus, Pipistrellus pygmaeus, Vespertilio murinus), of which breeding of two (M. nattereri, P. nathusii) have been confirmed. The total number of records in the study is 11 234, of which 9717 are identified to species. The records are from 940 (25%) 10-km2 squares of Finland’s land area. Of the records, 89% are new (1993–2014). Of the recorded bat species, only Eptesicus nilssonii occurs in each of the 21 biogeographical provinces. A decreasing south–north gradient in species richness and abundance exists which may be related to research efforts that are clearly higher in the south. This atlas is based on information in museum collections, literature, databases and unpublished data. In the last 150 years, the number of bat species in Finland increased from six to thirteen. Of these, five are common and regularly breeding (Eptesicus nilssonii, Myotis brandtii, Myotis daubentonii, Myotis mystacinus, Plecotus auritus), and eight rare (Eptesicus serotinus, Myotis dasycneme, Myotis nattereri, Nyctalus noctula, Pipistrellus nathusii, Pipistrellus pipistrellus, Pipistrellus pygmaeus, Vespertilio murinus), of which breeding of two (M. nattereri, P. nathusii) have been confirmed. The total number of records in the study is 11 234, of which 9717 are identified to species. The records are from 940 (25%) 10-km2 squares of Finland’s land area. Of the records, 89% are new (1993–2014). Of the recorded bat species, only Eptesicus nilssonii occurs in each of the 21 biogeographical provinces. A decreasing south–north gradient in species richness and abundance exists which may be related to research efforts that are clearly higher in the south. peerReviewed

Aerial migration is the fastest, yet most energetically demanding way of seasonal movement between habitats. However, for many taxa, and bats in particular, we lack a clear understanding of the energy requirements for migration. Here, we examined the energetic cost and flight speed of the long-distance migratory Nathusius’ bat (Pipistrellus nathusii). We measured flight metabolism in relation to airspeed in a wind tunnel, inferred the optimal traveling speed over long distances, i.e. maximum range speed, and compared this value with flight speed measured in wild conspecifics. Body mass and wing morphologies were similar in captive and wild bats, indicating that the body condition of captive bats was similar to that of migratory bats. Nine out of the 12 captive bats exhibited a U-shaped relationship between flight metabolic power and airspeed when flying in the wind tunnel. The flight metabolic rate across all airspeeds averaged 0.98±0.28 W, which corresponds well to established allometric relationships between flight metabolic rate and body mass for bats. During summer migration, P. nathusii traveled at an average speed of 6.9±0.7 m s−1, which was significantly higher than the minimum power speed (5.8±1.0 m s−1), yet within the range of expected maximum range speed inferred from wind tunnel experiments. This suggests that P. nathusii may migrate at an energetically optimal speed and that aerial refueling does not substantially lower migratory speed in P. nathusii.

Similar to birds, bats also perform long-distance migration between their breeding and wintering sites. In Northern Europe, migratory bat species are often detected along the coastline of the Baltic Sea particularly during migration seasons in the spring and autumn. In spite of regular monitoring of bat migration at coastal sites, the overall distribution of migratory bats in Northern Europe and variability between sites and seasons are still very poorly known. In this study we used automated bat detectors to compare the activity of migratory bat species between coastal and inland monitoring sites along the west coast of Finland (61.5–61.9°N, 21.3−22.3°E). Our main goal was to test whether the activity of migratory bat species is associated with the coastline or whether these species also occur inland. Of migratory bat species observations, 98.6% were covered by Pipistrellus nathusii, which was detected at all our monitoring sites. The activity of the species decreased rapidly, with increasing distance from the coastline towards inland, indicating a sharp activity gradient along the coastline of the Baltic Sea. Because the activity of P. nathusii occurred in migration season and no similar spatial pattern was detected among sedentary species, our results suggest that the aggregation of P. nathusii at the coastline is related to migration as such rather than regular foraging behavior of this species. Our study has direct implications to the wind power planning in Northern Europe. Based on our study we conclude that the impact of wind power on both migratory (namely P. nathusii) and sedentary bat species (namely Eptesicus nilssonii) should be taken into account in wind power planning and impact mitigation in Northern Europe, especially if new wind farms are located along the coastline of the Baltic Sea. When the turbines are located further inland, more attention in the planning process should however be given to the sedentary bat E. nilssonii.