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The Earth's magnetic field is used as a navigational cue by many animals. For mammals, however, there are few data to show that navigation ability relies on sensing the natural magnetic field. In night-time migrating bats, experiments demonstrating a role for the solar azimuth at sunset in the calibration of the orientation system suggest that the magnetic field is a candidate for their compass. Here, we investigated how an altered magnetic field at sunset changes the nocturnal orientation of the bat Pipistrellus pygmaeus . We exposed bats to either the natural magnetic field, a horizontally shifted field (120°), or the same shifted field combined with a reversal of the natural value of inclination (70° to -70°). We later released the bats and found that the take-off orientation differed among all treatments. Bats that were exposed to the 120° shift were unimodally oriented northwards in contrast to controls which exhibited a bimodal north-south distribution. Surprisingly, the orientation of bats exposed to both a 120° shift and reverse inclination was indistinguishable from a uniform distribution. These results suggest that these migratory bats calibrate the magnetic field at sunset, and for the first time, they show that bats are sensitive to the angle of magnetic inclination.

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., (© 2021 The Authors. Ecological Applications published by Wiley Periodicals LLC on behalf of The Ecological Society of America.)

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.

Integrating information on species-specific sensory perception with spatial activity provides a high-resolution understanding of how animals explore environments, yet frequently used exploration assays commonly ignore sensory acquisition as a measure for exploration. Echolocation is an active sensing system used by hundreds of mammal species, primarily bats. As echolocation call activity can be reliably quantified, bats present an excellent model system to investigate intraspecific variation in environmental cue sampling. Here, we developed an in situ roost-like novel environment assay for tree-roosting bats. We repeatedly tested 52 individuals of the migratory bat species, Pipistrellus nathusii, across 24 h, to examine the role of echolocation when crawling through a maze-type arena and test for consistent intraspecific variation in sensory-based exploration. We reveal a strong correlation between echolocation call activity and spatial activity. Moreover, we show that during the exploration of the maze, individuals consistently differed in spatial activity as well as echolocation call activity, given their spatial activity, a behavioral response we term 'acoustic exploration'. Acoustic exploration was correlated with other exploratory behaviors, but not with emergence latency. We here present a relevant new measure for exploration behavior and provide evidence for consistent (short-term) intra-specific variation in the level at which wild bats collect information from a novel environment.

Rationale: Identifying migratory corridors of animals is essential for their effective protection, yet the exact location of such corridors is often unknown, particularly for elusive animals such as bats. While migrating along the German coastline, Nathusius' pipistrelles (Pipistrellus nathusii) are regularly killed at wind turbines. Therefore, we explored the paths taken on their annual journey., Methods: We used isotope ratio mass spectrometry to measure stable hydrogen and strontium isotope ratios in fur keratin of 59 Nathusius' pipistrelles captured on three offshore islands. Samples were pre-treated before analysis to report exclusively stable isotope ratios of non-exchangeable hydrogen. We generated maps to predict summer origins of bats using isoscape models., Results: Bats were classified as long-distance migrants, mostly originating from Eastern Europe. Hydrogen analysis suggested for some bats a possible Fennoscandian origin, yet additional information from strontium analysis excluded this possibility. Instead, our data suggest that most Nathusius' pipistrelles migrating along the German coastline were of continental European summer origin, but also highlight the possibility that Nathusius' pipistrelles of Baltorussian origin may travel offshore from Fennoscandia to Germany., Conclusions: Our findings demonstrate the benefit of using complementary isotopic tracers for analysing the migratory pathways of bats and also potentially other terrestrial vertebrate species. Furthermore, data from our study suggest an offset of fur strontium isotope ratios in relation to local bedrock., (© 2020 The Authors. Rapid Communications in Mass Spectrometry published by John Wiley & Sons Ltd.)

Maintaining a competent immune system is energetically costly and thus immunity may be traded against other costly traits such as seasonal migration. Here, we tested in long-distance migratory Nathusius' pipistrelles (Pipistrellus nathusii), if selected branches of immunity are expressed differently in response to the energy demands and oxidative stress of aerial migration. During the migration period, we observed higher baseline lymphocyte and lower neutrophil levels than during the pre-migration period, but no stronger response of cellular effectors to an antigen challenge. Baseline plasma haptoglobin, as a component of the humoral innate immunity, remained similar during both seasons, yet baseline plasma haptoglobin levels increased by a factor of 7.8 in migratory bats during an immune challenge, whereas they did not change during the pre-migration period. Oxidative stress was higher during migration than during pre-migration, yet there was no association between blood oxidative status and immune parameters, and immune challenge did not trigger any changes in oxidative stress, irrespective of season. Our findings suggest that humoral effectors of the acute phase response may play a stronger role in the first-line defense against infections for migrating bats compared to non-migrating bats. We conclude that Nathusius' pipistrelles allocate resources differently into the branches of their immune system, most likely following current demands resulting from tight energy budgets during migration.

From bats to whales, millions of mammals migrate every year. However, their navigation capacity for accomplishing long-distance movements remains remarkably understudied and lags behind by five decades compared to other animals [1, 2]-partly because, unlike for other taxa, such as birds and sea turtles, no small-scale orientation assay has so far been developed. Yet recently, bats became a model to investigate which cues mammals use for long-range navigation, and, surprisingly for nocturnal animals, sunset cues, and particularly polarized-light cues, appear to be crucial for calibration of the magnetic-compass system in non-migratory bats [3-5]. This does not appear to hold for a species of migratory bat, however [6], and thus the nature of the information used by migratory bats for navigation remains unclear. Here, we asked whether the position of the solar disk per se is relevant for compass orientation in a migratory bat, Pipistrellus pygmaeus. Using a new experimental assay that measures takeoff orientation, we tested the orientation of bats exposed to a shifted sunset azimuth using a mirror at dusk. Bats exposed to a 180°-rotated azimuth of the setting sun and released after translocation during the same night shifted their heading direction by ∼180° compared to control bats. However, first-year migrants had no clear orientation either as controls or after the same treatment. This suggests that learning the migratory direction is a key component in the navigational system of naive bats in this species. Our study provides rare evidence for the specific cues and mechanisms that migratory mammals use for navigation., (Copyright © 2019. Published by Elsevier Ltd.)

The killing of large numbers of migratory bats at wind turbines is a pressing conservation problem. Even though avoidance and mitigation measures could benefit from a better knowledge of the species' migratory habits, we lack basic information about what habitats and corridors bats use during migration. We studied the isotopic niche dimensions of three bat species that are frequently killed at wind turbines in Germany: non-migratory Pipistrellus pipistrellus, mid-distance migratory Nyctalus noctula, and long- distance migratory Pipistrellus nathusii. We measured stable carbon and nitrogen isotope ratios (δ¹³C, δ¹⁵N) in five tissues that differed in isotopic retention time (fur, wing membrane tissue, muscle, liver, blood) to shed light on the species-specific habitat use during the autumn migration period using standard ellipse areas (SEAc). Further, we used stable isotope ratios of non-exchangeable hydrogen (δ²H(K)) in fur keratin to assess the breeding origin of bats. We inferred from isotopic composition (δ¹³C, δ¹⁵N) of fur keratin that isotopic niche dimensions of P. nathusii was distinct from that of N. noctula and P. pipistrellus, probably because P. nathusii was using more aquatic habitats than the other two species. Isoscape origin models supported that traveled distances before dying at wind turbines was largest for P. nathusii, intermediate for N. noctula, and shortest for P. pipistrellus. Isotopic niche dimensions calculated for each sample type separately reflected the species' migratory behavior. Pipistrellus pipistrellus and N. noctula showed similar isotopic niche breadth across all tissue types, whereas SEAc values of P. nathusii increased in tissues with slow turnaround time. Isotopic data suggested that P. nathusii consistently used aquatic habitats throughout the autumn period, whereas N. noctula showed a stronger association with terrestrial habitats during autumn compared to the pre-migration period.

In a recent study, Greif et al. (Greif et al. Nat Commun 5, 4488. (doi:10.1038/ncomms5488)) demonstrated a functional role of polarized light for a bat species confronted with a homing task. These non-migratory bats appeared to calibrate their magnetic compass by using polarized skylight at dusk, yet it is unknown if migratory bats also use these cues for calibration. During autumn migration, we equipped Nathusius' bats, Pipistrellus nathusii, with radio transmitters and tested if experimental animals exposed during dusk to a 90° rotated band of polarized light would head in a different direction compared with control animals. After release, bats of both groups continued their journey in the same direction. This observation argues against the use of a polarization-calibrated magnetic compass by this migratory bat and questions that the ability of using polarized light for navigation is a consistent feature in bats. This finding matches with observations in some passerine birds that used polarized light for calibration of their magnetic compass before but not during migration., (© 2015 The Author(s).)

Over recent years, it became widely accepted that alternative, renewable energy may come at some risk for wildlife, for example, when wind turbines cause large numbers of bat fatalities. To better assess likely populations effects of wind turbine related wildlife fatalities, we studied the geographical origin of the most common bat species found dead below German wind turbines, the noctule bat (Nyctalus noctula). We measured stable isotope ratios of non-exchangeable hydrogen in fur keratin to separate migrants from local individuals, used a linear mixed-effects model to identify temporal, spatial and biological factors explaining the variance in measured stable isotope ratios and determined the geographical breeding provenance of killed migrants using isoscape origin models. We found that 72% of noctule bat casualties (n = 136) were of local origin, while 28% were long-distance migrants. These findings highlight that bat fatalities at German wind turbines may affect both local and distant populations. Our results indicated a sex and age-specific vulnerability of bats towards lethal accidents at turbines, i.e. a relatively high proportion of killed females were recorded among migratory individuals, whereas more juveniles than adults were recorded among killed bats of local origin. Migratory noctule bats were found to originate from distant populations in the Northeastern parts of Europe. The large catchment areas of German wind turbines and high vulnerability of female and juvenile noctule bats call for immediate action to reduce the negative cross-boundary effects of bat fatalities at wind turbines on local and distant populations. Further, our study highlights the importance of implementing effective mitigation measures and developing species and scale-specific conservation approaches on both national and international levels to protect source populations of bats. The efficacy of local compensatory measures appears doubtful, at least for migrant noctule bats, considering the large geographical catchment areas of German wind turbines for this species.