Low foraging rates drive large insectivorous bats away from urban areas.

Urbanization has significant impacts on wildlife and ecosystems and acts as an environmental filter excluding certain species from local ecological communities. Specifically, it may be challenging for some animals to find enough food in urban environments to achieve a positive energy balance. Because urban environments favor small-sized bats with low energy requirements, we hypothesized that common noctules (Nyctalus noctula) acquire food at a slower rate and rely less on conspecifics to find prey in urban than in rural environments due to a low food abundance and predictable distribution of insects in urban environments. To address this, we estimated prey sizes and measured prey capture rates, foraging efforts, and the presence of conspecifics during hunting of 22 common noctule bats equipped with sensor loggers in an urban and rural environment. Even though common noctule bats hunted similar-sized prey in both environments, urban bats captured prey at a lower rate (mean: 2.4 vs. 6.3 prey attacks/min), and a lower total amount of prey (mean: 179 vs. 377 prey attacks/foraging bout) than conspecifics from rural environments. Consequently, the energy expended to capture prey was higher for common noctules in urban than in rural environments. In line with our prediction, urban bats relied less on group hunting, likely because group hunting was unnecessary in an environment where the spatial distribution of prey insects is predictable, for example, in parks or around floodlights. While acknowledging the limitations of a small sample size and low number of spatial replicates, our study suggests that scarce food resources may make urban habitats unfavorable for large bat species with higher energy requirements compared to smaller bat species. In conclusion, a lower food intake may displace larger species from urban areas making habitats with high insect biomass production key for protecting large bat species in urban environments.

Coastal onshore wind turbines lead to habitat loss for bats in Northern Germany.

Wind energy production is particularly rewarding along coastlines, yet coastlines are often important as migratory corridors for wildlife. This creates a conflict between energy production from renewable sources and conservation goals, which needs to be considered during environmental planning. To shed light on the spatial interactions of a high collision risk bat species with coastal wind turbines (WT), we analysed 32 tracks of 11 common noctule bats (Nyctalus noctula) in Northern Germany with miniaturized global positioning system units yielding 6266 locations. We used three spatial models to infer on the preferred and avoided landscape features in interaction with WT. We found 3.4% of all locations close to WT, with bats preferring areas with high levels of impervious surface, identified as farmhouses. Common noctule bats were also more present close to WT adjacent to paths and waterbodies. At the local scale, >70% of common noctule bats avoided WT, yet if bats approached WT we counted more positions at large WT, specifically close to known roosts. Our study highlights that coastal WT should not be placed next to feeding grounds and bat roosts. Additionally, avoidance of WT by bats indicates that foraging bats may suffer from habitat loss in coastal landscapes with high turbine densities. To mitigate the conflict between wind energy power production and conservation goals at coastal sites, wind turbines should be placed at distance to habitat features preferred by bats and turbine densities should be limited.

Free-ranging bats combine three different cognitive processes for roost localization.

Animals have evolved different cognitive processes to localize crucial resources that are difficult to find. Relevant cognitive processes such as associative learning and spatial memory have commonly been studied in a foraging related context under controlled laboratory conditions. However, in natural environments, animals can use multiple cognitive processes to localize resources. In this field study, we used a pairwise choice experiment and automatic roost monitoring to assess how individually marked, free-ranging Bechstein's bats belonging to two different colonies use associative learning, spatial memory and social information when localizing suitable day roosts. To our knowledge, this study tests for the first time how associative learning, spatial memory and social information are used in the process of roost localization in bats under the natural conditions. We show that, when searching for new roosts, bats used associative learning to discriminate between suitable and unsuitable roosts. For re-localizing previously occupied roosts, bats used spatial memory rather than associative learning. Moreover, bats significantly improved the localization of suitable unfamiliar roosts and tended to increase their accuracy to re-localize previously occupied day roosts using social information. Our field experiments suggest that Bechstein's bats make hierarchical use of different cognitive processes when localizing day roosts. More generally, our study underlines that evaluating different cues under natural conditions is fundamental to understanding how natural selection has shaped the cognitive processes used for localizing resources.

Effects of tag mass on the physiology and behaviour of common noctule bats.

BACKGROUND: External tags, such as transmitters and loggers, are often used to study bat movements. However, physiological and behavioural effects on bats carrying tags have rarely been investigated, and recommendations on the maximum acceptable tag mass are rather based on rules of thumb than on rigorous scientific assessment. METHODS: We conducted a comprehensive three-step assessment of the potential physiological and behavioural effects of tagging bats, using common noctules Nyctalus noctula as a model. First, we examined seasonal changes in body mass. Second, we predicted and then measured potential changes in flight metabolic rate in a wind tunnel. Third, we conducted a meta-analysis of published data to assess effects of different tag masses on the weight and behaviour of bats. RESULTS: Individual body mass of common noctules varied seasonally by 7.0 ± 2.6 g (range: 0.5-11.5 g). Aerodynamic theory predicted a 26% increase in flight metabolic rate for a common noctule equipped with a 3.8 g tag, equating to 14% of body mass. In a wind tunnel experiment, we could not confirm the predicted increase for tagged bats. Our meta-analysis revealed a weak correlation between tag mass and emergence time and flight duration in wild bats. Interestingly, relative tag mass (3-19% of bat body mass) was not related to body mass loss, but bats lost more body mass the longer tags were attached. Notably, relatively heavy bats lost more mass than conspecifics with a more average body mass index. CONCLUSION: Because heavy tags (> 3 g) were generally used for shorter periods of time than lighter tags (~ 1 g), the long-term effects of heavy tags on bats cannot be assessed at this time. Furthermore, the effects of disturbance and resource distribution in the landscape cannot be separated from those of tagging. We recommend that tags weighing 5-10% of a bat's mass should only be applied for a few days. For longer studies, tags weighing less than 5% of a bat's body mass should be used. To avoid adverse effects on bats, researchers should target individuals with average, rather than peak, body mass indices.

Wind energy production in forests conflicts with tree-roosting bats.

Many countries are investing heavily in wind power generation,1 triggering a high demand for suitable land. As a result, wind energy facilities are increasingly being installed in forests,2,3 despite the fact that forests are crucial for the protection of terrestrial biodiversity.4 This green-green dilemma is particularly evident for bats, as most species at risk of colliding with wind turbines roost in trees.2 With some of these species reported to be declining,5,6,7,8 we see an urgent need to understand how bats respond to wind turbines in forested areas, especially in Europe where all bat species are legally protected. We used miniaturized global positioning system (GPS) units to study how European common noctule bats (Nyctalus noctula), a species that is highly vulnerable at turbines,9 respond to wind turbines in forests. Data from 60 tagged common noctules yielded a total of 8,129 positions, of which 2.3% were recorded at distances <100 m from the nearest turbine. Bats were particularly active at turbines <500 m near roosts, which may require such turbines to be shut down more frequently at times of high bat activity to reduce collision risk. Beyond roosts, bats avoided turbines over several kilometers, supporting earlier findings on habitat loss for forest-associated bats.10 This habitat loss should be compensated by developing parts of the forest as refugia for bats. Our study highlights that it can be particularly challenging to generate wind energy in forested areas in an ecologically sustainable manner with minimal impact on forests and the wildlife that inhabit them.

Long-term individualized monitoring of sympatric bat species reveals distinct species- and demographic differences in hibernation phenology.

BACKGROUND: Hibernation allows species to conserve energy and thereby bridge unfavorable environmental conditions. At the same time, hibernation imposes substantial ecological and physiological costs. Understanding how hibernation timing differs within and between species can provide insights into the underlying drivers of this trade-off. However, this requires individualized long-term data that are often unavailable. Here, we used automatic monitoring techniques and a reproducible analysis pipeline to assess the individualized hibernation phenology of two sympatric bat species. Our study is based on data of more than 1100 RFID-tagged Daubenton's bats (Myotis daubentonii) and Natterer's bats (Myotis nattereri) collected over seven years at a hibernaculum in Germany. We used linear mixed models to analyze species-, sex- and age-specific differences in entrance, emergence and duration of the longest continuous period spent in the hibernaculum. RESULTS: Overall, Daubenton's bats entered the hibernaculum earlier and emerged later than Natterer's bats, resulting in a nearly twice as long hibernation duration. In both species, adult females entered earlier and emerged from hibernation later than adult males. Hibernation duration was shorter for juveniles than adults with the exception of adult male Natterer's bats whose hibernation duration was shortest of all classes. Finally, hibernation timing differed among years, but yearly variations in entrance and emergence timing were not equally shifted in both species. CONCLUSIONS: Our results suggest that even in sympatric species, and across sex and age classes, hibernation timing may be differentially affected by environmental conditions. This highlights the necessity of using individualized information when studying the impact of changing environments on hibernation phenology.

Differences in seasonal survival suggest species-specific reactions to climate change in two sympatric bat species.

Long-lived animals with a low annual reproductive output need a long time to recover from population crashes and are, thus, likely to face high extinction risk, if the current global environmental change will increase mortality rates. To aid conservation of those species, knowledge on the variability of mortality rates is essential. Unfortunately, however, individual-based multiyear data sets that are required for that have only rarely been collected for free-ranging long-lived mammals. Here, we used a five-year data set comprising activity data of 1,445 RFID-tagged individuals of two long-lived temperate zone bat species, Natterer's bats (Myotis nattereri) and Daubenton's bats (Myotis daubentonii), at their joint hibernaculum. Both species are listed as being of high conservation interest by the European Habitats Directive. Applying mixed-effects logistic regression, we explored seasonal survival differences in these two species which differ in foraging strategy and phenology. In both species, survival over the first winter of an individual's life was much lower than survival over subsequent winters. Focussing on adults only, seasonal survival patterns were largely consistent with higher winter and lower summer survival but varied in its level across years in both species. Our analyses, furthermore, highlight the importance of species-specific time periods for survival. Daubenton's bats showed a much stronger difference in survival between the two seasons than Natterer's bats. In one exceptional winter, the population of Natterer's bats crashed, while the survival of Daubenton's bats declined only moderately. While our results confirm the general seasonal survival pattern typical for hibernating mammals with higher winter than summer survival, they also show that this pattern can be reversed under particular conditions. Overall, our study points toward a high importance of specific time periods for population dynamics and suggests species-, population-, and age class-specific responses to global climate change.