Implications for the protection of nocturnal migrants

The replacement of conventional lighting with energy‐saving light emitting diodes (LED) is a worldwide trend, yet its consequences for animals and ecosystems are poorly understood. Strictly nocturnal animals such as bats are particularly sensitive to artificial light at night (ALAN). Past studies have shown that bats, in general, respond to ALAN according to the emitted light color and that migratory bats, in particular, exhibit phototaxis in response to green light. As red and white light is frequently used in outdoor lighting, we asked how migratory bats respond to these wavelength spectra. At a major migration corridor, we recorded the presence of migrating bats based on ultrasonic recorders during 10‐min light‐on/light‐off intervals to red or warm‐white LED, interspersed with dark controls. When the red LED was switched on, we observed an increase in flight activity for Pipistrellus pygmaeus and a trend for a higher activity for Pipistrellus nathusii. As the higher flight activity of bats was not associated with increased feeding, we rule out the possibility that bats foraged at the red LED light. Instead, bats may have flown toward the red LED light source. When exposed to warm‐white LED, general flight activity at the light source did not increase, yet we observed an increased foraging activity directly at the light source compared to the dark control. Our findings highlight a response of migratory bats toward LED light that was dependent on light color. The most parsimonious explanation for the response to red LED is phototaxis and for the response to warm‐white LED foraging. Our findings call for caution in the application of red aviation lighting, particularly at wind turbines, as this light color might attract bats, leading eventually to an increased collision risk of migratory bats at wind turbines

Nathusius’ bats, Pipistrellus nathusii, bypass mating opportunities of their own species, but respond to foraging heterospecifics on migratory transit flights

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

In situ novel environment assay reveals acoustic exploration as a repeatable behavioral response in migratory bats

Integrating information on species-specific sensory perception together 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 animal model to investigate intra-specific variation in environmental cue sampling. Here, we developed an in situ roost-like novel environment assay for tree-cave roosting bats. We repeatedly tested 52 individuals of the migratory bat species, Pipistrellus nathusii, across 24 hours, to examine the role of echolocation when crawling through a maze-type arena and test for consistent intra-specific 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

First record of a Nathusius' pipistrelle (Pipistrellus nathusii) overwintering at a latitude above 60°N

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.</p

Polarized skylight does not calibrate the compass system of a migratory bat

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

Nathusius' bats optimize long-distance migration by flying at maximum range speed

Metabolic rate of 12 Pipistrellus nathusii in relation to varying airspeed. We measured the metabolic rate of flying bats in a wind tunnel using the 13C labeled Na-bicarbonate method. The relationship between metabolic rate and airspeed was U-shaped in the majority of individuals. We could not find a U-shaped curve in a few individuals that engaged in flight manoeuvers, including landing. We used the shape of the U-shaped power curve to estimate minimum flight speed and maximum range speed for the study species. Further we present data on migration speed (n=37) and foraging flight speed (n=40) in wild Pipistrellus nathusii at a major miratory corridor in Latvia

Migratory bats respond to artificial green light with positive phototaxis

Artificial light at night is spreading worldwide at unprecedented rates, exposing strictly nocturnal animals such as bats to a novel anthropogenic stressor. Previous studies about the effect of artificial light on bats focused almost exclusively on non-migratory species, yet migratory animals such as birds are known to be largely affected by light pollution. Thus, we conducted a field experiment to evaluate if bat migration is affected by artificial light at night. In late summer, we presented artificial green light of 520 nm wavelength to bats that were migrating south along the shoreline of the Baltic Sea. Using a light on-off treatment, we observed that the activity of Pipistrellus nathusii and P. pygmaeus, the two most abundant migratory species at our site, increased by more than 50% in the light-on compared to the light-off treatment. We observed an increased number of feeding buzzes during the light-on compared to the light-off treatment for P. nathusii. However, feeding activity was low in general and did not increase disproportionately during the light-on treatment in relation to the overall echolocation call activity of bats. Further, P. nathusii were attracted towards the green light at a distance of about 23 m, which is way beyond the echolocation detection range for insects of Nathusius’ bats. We therefore infer that migratory bats were not attracted to artificial green light because of high insect densities, but instead by positive phototaxis. We conclude that artificial light at night may potentially impact bat migration in a yet unrecognized way

Data from: Diet of the insectivorous bat Pipistrellus nathusii during autumn migration and summer residence

Migration is widespread among vertebrates. Yet bat migration has received little attention and only in the recent decades knowledge of it has been gained. Migration can cause significant changes in behaviour and physiology, due to increasing energy demands and aerodynamic constraints. Dietary shifts, for examples, have been shown to occur in birds before onset of migration. For bats it is not known if a change in diet occurs during migration, although especially breeding season related dietary preference has been documented. It is known that fat-rich diets, and subsequent accumulation of high fat deposits, do increase the flight range of migratory bats. Some bat species can be regarded as long-distance migrants, covering up to 2,000 km on their way between summer and winter roosting areas. Pipistrellus nathusii (Vespertilionidae), a European long-distant migrant, travels each year along the Baltic Sea from north-eastern Europe to hibernate in central and southern Europe. This study presents data on the dietary habits of migrating Pipistrellus nathusii compared with dietary habits during the breeding season. We analysed faecal samples from bats on fall migration caught at the Ornithological Field Station in Pape, Latvia and from samples collected in North-Latvian summer roosts. We applied both morphological identification and molecular methods, as morphological methods also recognize life stages of prey and can contribute frequency data. The diets of bats on migration and breeding bats were similar, with Diptera and Lepidoptera comprising the major prey categories. However certain prey groups could be explained by the different hunting habitats used during migration vs. summer residence