Terrestrial locomotion imposes high metabolic requirements on bats

The evolution of powered flight involved major morphological changes in Chiroptera. Nevertheless, all bats are also capable of crawling on the ground and some are even skilled sprinters. We asked if a highly derived morphology adapted for flapping flight imposes high metabolic requirements on bats when moving on the ground. We measured the metabolic rate during terrestrial locomotion in mastiff bats, Molossus currentium, a species that is both a fast-flying aerial-hawking bat and an agile crawler on the ground. Metabolic rates of bats averaged 8.0±4.0 ml CO2 min–1 during a 1-min period of sprinting at 1.3±0.6 km h–1. With rising average speed, mean metabolic rates increased, reaching peak values that were similar to those of flying conspecifics. Metabolic rates of M. currentium were higher than those of similar-sized rodents that sprinted at similar velocities under steady-state conditions. When M. currentium sprinted at peak velocities, its aerobic metabolic rate was 3–5 times higher than those of rodent species running continuously in steady-state conditions. Costs of transport (J kg–1 m–1) were more than 10 times higher for running than for flying bats. We conclude that at the same speed bats experience higher metabolic rates during short sprints than quadruped mammals during steady-state terrestrial locomotion, yet running bats achieve higher maximal mass-specific aerobic metabolic rates than non-volant mammals such as rodents

Rain increases the energy cost of bat flight

Similar to insects, birds and pterosaurs, bats have evolved powered flight. But in contrast to other flying taxa, only bats are furry. Here, we asked whether flight is impaired when bat pelage and wing membranes get wet. We studied the metabolism of short flights in Carollia sowelli, a bat that is exposed to heavy and frequent rainfall in neotropical rainforests. We expected bats to encounter higher thermoregulatory costs, or to suffer from lowered aerodynamic properties when pelage and wing membranes catch moisture. Therefore, we predicted that wet bats face higher flight costs than dry ones. We quantified the flight metabolism in three treatments: dry bats, wet bats and no rain, wet bats and rain. Dry bats showed metabolic rates predicted by allometry. However, flight metabolism increased twofold when bats were wet, or when they were additionally exposed to rain. We conclude that bats may not avoid rain only because of sensory constraints imposed by raindrops on echolocation, but also because of energetic constraints

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

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

Equivariant cyclic homology

In der vorliegenden Arbeit wird äquivariante zyklische Homologie definiert und untersucht. Von zentraler Bedeutung ist die Tatsache, dass die zugrundeliegenden Objekte der Theorie keine Kettenkomplexe im Sinne der homologischen Algebra sind. Eine Konsequenz hiervon ist, dass im äquivarianten Kontext im wesentlichen nur die periodische zyklische Homologie sinnvoll definiert werden kann. Wir zeigen, dass die äquivariante bivariante periodische zyklische Theorie homotopieinvariant und stabil ist und Ausschneidung in beiden Variablen erfüllt. Weiter beweisen wir ein Analogon des Satzes von Green-Julg für endliche Gruppen und einen dualen Satz von Green-Julg für beliebige diskrete Gruppen. Schließlich untersuchen wir Wirkungen von diskreten Gruppen auf simplizialen Komplexen. Wir zeigen, dass die äquivariante zyklische Homologie dieser Algebren in enger Beziehung zu einer von Baum und Schneider entwickelten äquivarianten Kohomologietheorie steht

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

Increasing species richness along elevational gradients is associated with niche packing in bat assemblages

1. The change in species richness along elevational gradients is a well-known pattern in nature. Niche theory predicts that increasing species richness in assemblages can either lead to denser packing of niche space (‘niche packing’) or an expansion into its novel regions (‘niche expansion’). Traditionally, these scenarios have been studied using functional traits but stable isotopes provide advantages such as identifying the degree of resource specialisation, or niche partitioning among functionally similar species. 2. In this study, we evaluate the relevance of niche packing versus niche expansion by investigating stable carbon and nitrogen isotopic niche width and overlap among 23 bat species from six functional groups across a 1500 m elevational gradient in the Himalaya. 3. Our results suggest that an increase in species richness in the low elevation is accompanied by small niche width with high overlap, whereas the high elevation assemblage shows large niche width with low overlap among functional group members. At the functional group level, edge-space foraging, trawling, and active gleaning bats have the highest niche width while passive gleaning bats that are only found in high elevations are isotopic specialists showing low overlap with other groups. Edge and open-space foraging bats showed idiosyncratic changes in niche width across elevations. We also find that the niches of rhinolophid bats overlap with edge-space and open-space foraging bats despite their unique functional traits. 4. These results support the idea that at low elevations high species richness is associated with niche packing while at high elevations strong niche partitioning prevails in dynamic and resource-poor environments. We conclude that although high elevation animal assemblages are often ‘functionally underdispersed’, that is show homogenous functional traits, our approach based on stable isotopes demonstrates niche partitioning among such functionally similar species

Migratory flight imposes oxidative stress in bats

Many animal species migrate over long distances, but the physiological challenges of migration are poorly understood. It has recently been suggested that increased molecular oxidative damage might be one important challenge for migratory animals. We tested the hypothesis that autumn migration imposes an oxidative challenge to bats by comparing values of 4 blood-based markers of oxidative status (oxidative damage and both enzymatic and nonenzymatic antioxidants) between Nathusius’ bats Pipistrellus nathusii that were caught during migration flights with those measured in conspecifics after resting for 18 or 24 h. Experiments were carried out at Pape Ornithological Station in Pape (Latvia) in 2016 and 2017. Our results show that flying bats have a blood oxidative status different from that of resting bats due to higher oxidative damage and different expression of both nonenzymatic and enzymatic antioxidants (glutathione peroxidase). The differences in oxidative status markers varied between sampling years and were independent from individual body condition or sex. Our work provides evidence that migratory flight might impose acute oxidative stress to bats and that resting helps animals to recover from oxidative damage accrued en route. Our data suggest that migrating bats and birds might share similar strategies of mitigating and recovering from oxidative stress

Immune Profile Predicts Survival and Reflects Senescence in a Small, Long- Lived Mammal, the Greater Sac-Winged Bat (Saccopteryx bilineata)

The immune system imposes costs that may have to be traded against investment of resources in other costly life-history traits. Yet, it is unknown if a trade-off between immunity and longevity occurs in free-ranging mammals. Here, we tested if age and survival, two aspects associated with longevity, are linked to immune parameters in an 8 g bat species. Using a combination of cross-sectional and longitudinal data, we assessed whether total white blood cell (WBC) counts, bacterial killing ability of the plasma (BKA) and immunoglobulin G (IgG) concentration change with age. Furthermore, we asked if these immune parameters impose costs resulting in decreased survival probabilities. We found that WBC counts decreased with age both within and among individuals. IgG concentrations were higher in older individuals, but did not change with age within individuals. Furthermore, individuals with above average WBC counts or IgG concentration had lower probabilities to survive the next six months. High WBC counts and IgG concentrations may reflect infections with parasites and pathogens, however, individuals that were infected with trypanosomes or nematodes showed neither higher WBC counts or IgG concentrations, nor was infection connected with survival rates. BKA was higher in infected compared with uninfected bats, but not related to age or survival. In conclusion, cellular (WBC) and humoral (IgG) parts of the immune system were both connected to age and survival, but not to parasite infections, which supports the hypothesis that energetically costly immunological defences are traded against other costly life-history traits, leading to a reduced lifespan in this free-ranging mammal