How Bats Escape the Competitive Exclusion Principle—Seasonal Shift From Intraspecific to Interspecific Competition Drives Space Use in a Bat Ensemble

Predators that depend on patchily distributed prey face the problem of finding food patches where they can successfully compete for prey. While the competitive exclusion principle suggests that species can only coexist if their ecological niches show considerable differences, newer theory proposes that local coexistence can be facilitated by so-called stabilizing and equalizing mechanisms. A prerequisite to identify such mechanisms is the understanding of the strength and the nature of competition (i.e., interference or exploitation). We studied the interaction between two open-space foraging bats by testing if common noctule bats Nyctalus noctula shift their space use in response to simulated aggregations of conspecifics or heterospecific Pipistrellus nathusii. When confronted with playbacks of heterospecifics, N. noctula increased their activity in early summer, but decreased activity in late summer. This pattern was accompanied by a decrease in the proportion of large insects in late summer, suggesting a more intense competition for food in late compared to early summer. When confronted with playbacks of conspecifics, N. noctula did not change their activity, irrespective of season. Our results indicate that in early summer, intraspecific competition is more severe than interspecific competition for insectivorous bats. Likely, conspecifics engage in interference competition for flight space, and may suffer from reduced prey detectability as echolocation calls of conspecifics interfere with each other. During insect rich times, interspecific competition on the other hand may be mediated by fine scale vertical partitioning and the use non-interfering echolocation frequencies. In contrast, when food is scarce in late summer, bats may engage in exploitation competition. Our data suggests that N. noctula avoid aggregations of more agile bats like P. nathusii, probably due to impeded hunting success. Yet, as fast and efficient fliers, N. noctula may be able to escape this disadvantage by exploiting more distant foraging patches

Aerial-hawking bats adjust their use of space to the lunar cycle

BACKGROUND: Animals change their habitat use in response to spatio-temporal fluctuation of resources. Some resources may vary periodically according to the moonphase. Yet it is poorly documented how animals, particularly nocturnal mammals, adjust their use of space in response to the moonphase. Here, we asked if an obligate nocturnal mammal, the aerial-hawking common noctule bat (Nyctalus noctula), adjusts its 3-dimensional flight behaviour and habitat use to the lunar period. Using miniaturized GPS loggers, we recorded 3-dimensional flight tracks of N. noctula and related these to a canopy height model derived from aerial laser scans to investigate whether bats adjust forest strata use to moonlight intensities. RESULTS: Noctules frequently foraged above the canopy of coniferous forest at low moonlight intensities, but switched to using open grasslands and arable fields in nights with high moonlight intensities. During the few occasions when noctules used the forest during moonlit nights, they mostly restricted their use of space to flying below the canopy level. The median overall flight altitude of N. noctula equalled 13 ± 16 m but reached up to 71 m above ground (97.5% quantile). CONCLUSIONS: Our findings argue against general lunar phobic behaviour of aerial-hawking bats. We suggest that the preferred use of open fields around full moon may be a strategy of noctules to increase the success of hunting airborne insects at night. Specifically, the adjustment in use of space may allow bats to hunt for insects that emerge and disperse over open fields during bright moonlight

Data_Sheet_3_How Bats Escape the Competitive Exclusion Principle—Seasonal Shift From Intraspecific to Interspecific Competition Drives Space Use in a Bat Ensemble.docx

<p>Predators that depend on patchily distributed prey face the problem of finding food patches where they can successfully compete for prey. While the competitive exclusion principle suggests that species can only coexist if their ecological niches show considerable differences, newer theory proposes that local coexistence can be facilitated by so-called stabilizing and equalizing mechanisms. A prerequisite to identify such mechanisms is the understanding of the strength and the nature of competition (i.e., interference or exploitation). We studied the interaction between two open-space foraging bats by testing if common noctule bats Nyctalus noctula shift their space use in response to simulated aggregations of conspecifics or heterospecific Pipistrellus nathusii. When confronted with playbacks of heterospecifics, N. noctula increased their activity in early summer, but decreased activity in late summer. This pattern was accompanied by a decrease in the proportion of large insects in late summer, suggesting a more intense competition for food in late compared to early summer. When confronted with playbacks of conspecifics, N. noctula did not change their activity, irrespective of season. Our results indicate that in early summer, intraspecific competition is more severe than interspecific competition for insectivorous bats. Likely, conspecifics engage in interference competition for flight space, and may suffer from reduced prey detectability as echolocation calls of conspecifics interfere with each other. During insect rich times, interspecific competition on the other hand may be mediated by fine scale vertical partitioning and the use non-interfering echolocation frequencies. In contrast, when food is scarce in late summer, bats may engage in exploitation competition. Our data suggests that N. noctula avoid aggregations of more agile bats like P. nathusii, probably due to impeded hunting success. Yet, as fast and efficient fliers, N. noctula may be able to escape this disadvantage by exploiting more distant foraging patches.</p

Data_Sheet_1_How Bats Escape the Competitive Exclusion Principle—Seasonal Shift From Intraspecific to Interspecific Competition Drives Space Use in a Bat Ensemble.DOCX

<p>Predators that depend on patchily distributed prey face the problem of finding food patches where they can successfully compete for prey. While the competitive exclusion principle suggests that species can only coexist if their ecological niches show considerable differences, newer theory proposes that local coexistence can be facilitated by so-called stabilizing and equalizing mechanisms. A prerequisite to identify such mechanisms is the understanding of the strength and the nature of competition (i.e., interference or exploitation). We studied the interaction between two open-space foraging bats by testing if common noctule bats Nyctalus noctula shift their space use in response to simulated aggregations of conspecifics or heterospecific Pipistrellus nathusii. When confronted with playbacks of heterospecifics, N. noctula increased their activity in early summer, but decreased activity in late summer. This pattern was accompanied by a decrease in the proportion of large insects in late summer, suggesting a more intense competition for food in late compared to early summer. When confronted with playbacks of conspecifics, N. noctula did not change their activity, irrespective of season. Our results indicate that in early summer, intraspecific competition is more severe than interspecific competition for insectivorous bats. Likely, conspecifics engage in interference competition for flight space, and may suffer from reduced prey detectability as echolocation calls of conspecifics interfere with each other. During insect rich times, interspecific competition on the other hand may be mediated by fine scale vertical partitioning and the use non-interfering echolocation frequencies. In contrast, when food is scarce in late summer, bats may engage in exploitation competition. Our data suggests that N. noctula avoid aggregations of more agile bats like P. nathusii, probably due to impeded hunting success. Yet, as fast and efficient fliers, N. noctula may be able to escape this disadvantage by exploiting more distant foraging patches.</p

Data_Sheet_2_How Bats Escape the Competitive Exclusion Principle—Seasonal Shift From Intraspecific to Interspecific Competition Drives Space Use in a Bat Ensemble.DOCX

<p>Predators that depend on patchily distributed prey face the problem of finding food patches where they can successfully compete for prey. While the competitive exclusion principle suggests that species can only coexist if their ecological niches show considerable differences, newer theory proposes that local coexistence can be facilitated by so-called stabilizing and equalizing mechanisms. A prerequisite to identify such mechanisms is the understanding of the strength and the nature of competition (i.e., interference or exploitation). We studied the interaction between two open-space foraging bats by testing if common noctule bats Nyctalus noctula shift their space use in response to simulated aggregations of conspecifics or heterospecific Pipistrellus nathusii. When confronted with playbacks of heterospecifics, N. noctula increased their activity in early summer, but decreased activity in late summer. This pattern was accompanied by a decrease in the proportion of large insects in late summer, suggesting a more intense competition for food in late compared to early summer. When confronted with playbacks of conspecifics, N. noctula did not change their activity, irrespective of season. Our results indicate that in early summer, intraspecific competition is more severe than interspecific competition for insectivorous bats. Likely, conspecifics engage in interference competition for flight space, and may suffer from reduced prey detectability as echolocation calls of conspecifics interfere with each other. During insect rich times, interspecific competition on the other hand may be mediated by fine scale vertical partitioning and the use non-interfering echolocation frequencies. In contrast, when food is scarce in late summer, bats may engage in exploitation competition. Our data suggests that N. noctula avoid aggregations of more agile bats like P. nathusii, probably due to impeded hunting success. Yet, as fast and efficient fliers, N. noctula may be able to escape this disadvantage by exploiting more distant foraging patches.</p

Habitat use of bats in relation to wind turbines revealed by GPS tracking

Worldwide, many countries aim at countering global climate change by promoting renewable energy. Yet, recent studies highlight that so-called green energy, such as wind energy, may come at environmental costs, for example when wind turbines kill birds and bats. Using miniaturized GPS loggers, we studied how an open-space foraging bat with high collision risk with wind turbines, the common noctule Nyctalus noctula (Schreber, 1774), interacts with wind turbines. We compared actual flight trajectories to correlated random walks to identify habitat variables explaining the movements of bats. Both sexes preferred wetlands but used conventionally managed cropland less than expected based on availability. During midsummer, females traversed the land on relatively long flight paths and repeatedly came close to wind turbines. Their flight heights above ground suggested a high risk of colliding with wind turbines. In contrast, males recorded in early summer commuted straight between roosts and foraging areas and overall flew lower than the operating range of most turbine blades, suggesting a lower collision risk. Flight heights of bats suggest that during summer the risk of collision with wind turbines was high for most studied bats at the majority of currently installed wind turbines. For siting of wind parks, preferred bat habitats and commuting routes should be identified and avoided

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

Landscape structure influences the use of social information in an insectivorous bat

In anthropogenic landscapes, aerial insectivores are often confronted with variable habitat complexity, which may influence the distribution of prey. Yet, high mobility may allow aerial insectivores to adjust their foraging strategy to different prey distributions. We investigated whether aerial-hunting common noctules Nyctalus noctula adjust their foraging strategy to landscapes with different habitat complexity and assumingly different prey distribution. We hypothesized that the movement behaviour of hunting common noctules and changes of movement behaviour in reaction towards conspecifics would depend on whether they hunt in a structurally poor cropland dominated landscape or a structurally rich forest dominated landscape. We tracked flight paths of common noctules in northeastern Germany using GPS loggers equipped with an ultrasonic microphone that recorded foraging events and presence of conspecifics. Above cropland, common noctules hunted mainly during bouts of highly tortuous and area restricted movements (ARM). Bats switched from straight flight to ARM after encountering conspecifics. In the forested landscape, common noctules hunted both during ARM and during straight flights. The onset of ARM did not correlate with the presence of conspecifics. Common noctules showed a lower feeding rate and encountered more conspecifics above the forested than above the cropland dominated landscape. We conjecture that prey distribution above cropland was patchy and unpredictable, thus making eavesdropping on hunting conspecifics crucial for bats during search for prey patches. In contrast, small scale structural diversity of the forested landscape possibly led to a more homogeneous prey distribution at the landscape scale, thus enabling bats to find sufficient food independent of conspecific presence. This suggests that predators depending on ephemeral prey can increase their foraging success in structurally poor landscapes by using social information provided by conspecifics. Hence, a minimum population density might be obligatory to enable successful foraging in simplified landscapes

Additional file 1: of Aerial-hawking bats adjust their use of space to the lunar cycle

Habitat types within the study area. The location of the artificial roosts is indicated by the white star. (PNG 3728 kb