Foraging habitats of Myotis emarginatus in Central Europe

We radio-tracked Myotis emarginatus in Upper Bavaria, Germany to identify the key-foraging habitats and to enable an adequate habitat management for this endangered species. The studied females foraged at a distance of up to 8.1 km around their colony roost. The average distance of the foraging area was 3.7 km, where 70% of foraging areas were located within a distance of 5 km and 90% within 6 km of the nurseries. Moreover, these bats spent about 75% of their foraging time within 5 km and 85% within a 6-km radius. To reach the foraging areas, the bats usually used riparian woodlands, hedges and tree lines as flight paths. Specifically, 46.9% of the foraging areas were located in forests, 24.5% in cow sheds, 18.4% in riparian woodlands along streams and 10.2% in fields, villages, orchards, hedges and groves on open land. On average, the bats foraged in forests for 56.2% of the time, during which habitat allocation was possible. In cow sheds the percentage was 29.2%, in riparian deciduous woodland 11.5% and in the other habitats 3.1%. Within forests M. emarginatus avoided foraging in spruce monocultures. Pure stands of spruce covered 45% of the total forest area, but only 10% of the foraging areas were located in this forest type. Deciduous forests on the other hand were much more common as foraging sites (40% versus 11%). Therefore, the availability of native deciduous forest and of fly-infested stables within a radius of 6 km around the colony roosts should be the focus of conservation concepts for M. emarginatus.     

The spatial ecology of the whiskered bat (Myotis mystacinus) at the western extreme of its range provides evidence of regional adaptation

Although widespread, the ecology of the whiskered bat, Myotis mystacinus in Europe remains poorly understood. Ireland is positioned at the most western extreme of this species’ range. To ascertain the ecology of M. mystacinus at its geographic range extreme, the roosting behaviour, home range and habitat use of females in a maternity roost in Ireland was investigated by radio-tracking. M. mystacinus were active in a diversity of habitats: namely, mixed woodland, riparian vegetation, arable land and rough grassland. However, only mixed woodland and riparian habitats were selected as core foraging areas. This is in contrast to a previous study from Britain where only pasture was utilised but is in agreement with data from Slovakia, where woodland was also selected, whilst riparian areas were also utilised by this species in Germany. A high degree of overlap in the foraging areas of individuals was observed. A total of seven roosts were utilised by tracked bats and roost switching behaviour was observed. We discuss our contrasting results in respect to range limitations, regional variability in landscape structure and the composition of bat communities. The present results have implications for the conservation of M. mystacinus within Ireland and other parts of its range, highlighting the need for range wide ecological studies. Regional variability in the ecology of bats related to landscape factors is an important consideration for bat conservation and therefore must be incorporated into future management plans.     

Use of foraging habitats by bats in a Mediterranean area determined by acoustic surveys: conservation implications

We determined habitat use by foraging bats by broad-band acoustic surveys in 10 habitat types from a Mediterranean area (southern Italy). We applied discriminant functions to identify time-expanded echolocation calls from free-flying bats.
Moon phase and cloud cover had no effect on bat activity. Only Hypsugo savii was influenced by temperature, and activity of Myotis daubentonii and Myotis capaccinii was reduced at higher wind speeds. Both total numbers of bat passes and feeding buzzes were highest over rivers and lakes. Pipistrellus kuhlii and H. savii were most frequently recorded. Pipistrellus kuhlii , Pipistrellus pipistrellus and Tadarida teniotis proved generalists in using foraging habitats.
Water sites and conifer plantations were respectively the most and the least used habitats by H. savii . Rivers were especially important to Myotis bats, Miniopterus schreibersii and Pipistrellus pygmaeus . Unlike P. kuhlii , P. pipistrellus was frequent in beech woodlands; P. pygmaeus made a considerable use of chestnut woodlands and Myotis spp. were moderately active in both these woodland types.
A large number of endangered or vulnerable species featured in riparian habitats, broadleaved woodlands and olive groves. Riparian and woodland habitats constitute an important target for conservation. Typical land use forms such as woodlands used for chestnut production and traditionally managed olive groves should be encouraged in conservation plans. The negative impact of urbanisation on bats might be counteracted by fostering trees, gardens and small cultivated patches. Farmland practices should encourage landscape complexity and limit the use of pesticides.     

Resource partitioning in two species of vespertilionid bats (Chiroptera) occupying the same roost

Colonies of Plecotus auritus (L.) and Myotis daubentoni (Kuhl) occupying the same roost showed no interspecific interaction. They utilized separate areas of the roost, departed from and returned to it at different times and foraged in different habitats on different groups of insects. P. auritus emerged from the roost before dusk, foraged singly in woodland areas with slow, hovering flight and fed on insects commonly found flying in woodland, on insects and spiders gleaned from foliage and on insects caught inside the roost. M. daubentoni emerged from the roost after dusk, flew fast and low over water and around riparian vegetation and foraged on long, straight beats. They frequently foraged in groups, and fed opportunistically on insects which fly in swarms in riparian habitats. The diets of the two species occupying the same roost did not differ from those established in areas where each species was found alone.     

Roosting and foraging behaviour of Natterer's bats (Myotis nattereri) close to the northern border of their distribution

Seven nursery roosts and four roosts of male Myotis nattereri , Kuhl 1818 were found in central Scotland at latitude 56–57 N. Most were in crevices in the stonework of man-made structures other than occupied houses. Emergence occurred late in the evening, at an average light intensity of 3.5 lux and emerging bats circled in dark, sheltered areas outside roosts before departing along flyways towards foraging areas. Individuals departed from, and returned to, roosts in groups of 2-6, and circling behaviour was repeated on returning to the roost. During pregnancy, bats from anursery roost made one flight each per night. This increased to an average maximum of 1.84 early in lactation and then decreased again to one around weaning. Night roosts were situated in foraging areas and were used by M. nattereri for resting and grooming, for suckling volant but incompletely weaned young and also, possibly, for information transfer. Important foraging habitats were woodland edges, parkland, roadside vegetation and sheltered areas of water. Arthropod prey was captured both on the wing and by gleaning from foliage, and the bats were able to vary their diet according to arthropod availability. Overall, important prey included Diptera (both Nematocera and higher flies), Trichoptera, Coleoptera and non-flying groups such as Hemiptera, Dermaptera, Arachnida and Opiliones.     

Pteropus lylei primarily forages in residential areas in Kandal,Cambodia

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Roosting and Foraging Social Structure of the Endangered Indiana Bat (Myotis sodalis)

Social dynamics are an important but poorly understood aspect of bat ecology. Herein we use a combination of graph theoretic and spatial approaches to describe the roost and social network characteristics and foraging associations of an Indiana bat (Myotis sodalis) maternity colony in an agricultural landscape in Ohio, USA. We tracked 46 bats to 50 roosts (423 total relocations) and collected 2,306 foraging locations for 40 bats during the summers of 2009 and 2010. We found the colony roosting network was highly centralized in both years and that roost and social networks differed significantly from random networks. Roost and social network structure also differed substantially between years. Social network structure appeared to be unrelated to segregation of roosts between age classes. For bats whose individual foraging ranges were calculated, many shared foraging space with at least one other bat. Compared across all possible bat dyads, 47% and 43% of the dyads showed more than expected overlap of foraging areas in 2009 and 2010 respectively. Colony roosting area differed between years, but the roosting area centroid shifted only 332 m. In contrast, whole colony foraging area use was similar between years. Random roost removal simulations suggest that Indiana bat colonies may be robust to loss of a limited number of roosts but may respond differently from year to year. Our study emphasizes the utility of graphic theoretic and spatial approaches for examining the sociality and roosting behavior of bats. Detailed knowledge of the relationships between social and spatial aspects of bat ecology could greatly increase conservation effectiveness by allowing more structured approaches to roost and habitat retention for tree-roosting, socially-aggregating bat species.     

Diet and prey selection in the trawling long-fingered bat

The diet of the long-fingered bat Myotis capaccinii is poorly known, and there is no previously recorded information on this species' prey preferences. To investigate these subjects, we captured 51 individuals at a nursery cave in the Iberian Peninsula, from pre-breeding to post-lactation seasons. Each bat's diet composition was assessed by faecal content analysis and its foraging places (rivers, pools and channels) identified by radio-telemetry. To estimate prey availability, we sampled arthropods in the individual bats' identified foraging places and also emulated the bats' hunting technique. The bats' diet comprised of arthropods, dominated by small insects with aquatic larvae and flying adult phases. The most consumed taxon was Nematocera (mainly Chironomidae), including adults and pupae, which were also found to be the most abundant prey over water. Other frequently consumed prey were brachycerans, lepidopterans, arachnids, trichopterans and neuropterans. Diet proportions were compared with prey availability to infer a rank of preferences. The preferred prey were lepidopterans and arachnids, both having a terrestrial life cycle and a bigger size than any other taxa consumed. Without discarding the possible underestimation of prey's aerial availability, the observed preference pattern seems to be a consequence of selection for size more than for specific taxa. Apparently M. capaccinii efficiently exploit water-related prey according to availability when the bats hunt low over the water's surface, and are also able to take more profitable prey found higher in the air.     

Distribution and population densities of seven species of bat in northern England

Distribution and minimum population densities for seven UK bat species known to be resident in northern England were calculated in an area covering 2500 km². The species present were pipistrelle ( Pipistrellus pipistrellus ), brown long-eared ( Plecotus auritus ), Daubenton's ( Myotis daubentonii ), whiskered ( Myotis mystacinus ), Natterer's (Myotis nattereri) , noctule ( Nyctalus noctula ) and Brandt's (Myotis brandtii). Data were collected primarily from counts at summer roosts over the period 1983 to 1990. A total of 310 bat roosts were discovered within the study area. Of the 256 roosts at which the species present was identified, the majority, 127 (49.6%) were P. pipistrellus , with a mean maternity roost size of 69.6 bats. A minimum population density of 12.6 batskm⁻² was estimated for P. pipistrellus , based on summer (maternity) roosts. The minimum population density estimate was higher than previous studies in northern England but substantially lower than those reported in Scotland (18.2 bats km-⁻²). The combined density of M. mystacinus, M. brandtii, and P. pipistrellus , which have similar foraging styles (15.8 bats km⁻²), is comparable to Scottish P. pipistrellus densities. The density of M. duubentonii was also lower than in Scotland, although the density of P. auritus was comparable. The majority of summer roosts for all species were found in buildings, except N. noctula and M. duubentonii which used bridges/tunnels or trees.     

The mitochondrial and nuclear genetic structure of Myotis capaccinii (Chiroptera: Vespertilionidae) in the Eurasian transition, and its taxonomic implications

Allopatric isolation in glacial refugia has caused differentiation and speciation in many taxa globally. In this study, we investigated the nuclear and mitochondrial genetic differentiation of the long fingered bat, Myotis capaccinii during the ice ages in south-eastern Europe and Anatolia. The mitochondrial DNA (mtDNA) analyses indicated a suture zone similar to those recorded in other animal species, including bats, suggesting the association of more than one refugium with the region. Contrary to most of the other species where a suture zone was seen in Anatolia, for M. capaccinii the geographical location of the genetic break was in south-eastern Europe. This mitochondrial differentiation was not reflected in the nuclear microsatellites, however, suggesting that the lack of contact during the ice ages did not result in reproductive isolation. Hence taxonomically, the two mitochondrial clades cannot be treated as separate species.     

Reproductive and age classes do not change spatial dynamics of foraging long-fingered bats (Myotis capaccinii)

Spatial dynamics of foraging long-fingered bats (Myotis capaccinii) were studied in the Eastern Iberian Peninsula. We analysed the locations of 45 radio-tracked individuals during three discrete periods through the breeding season and measured the spatial parameters related to their foraging behaviour in order to test whether variations in spatial use occur. Colony range, measured as the minimum convex polygon through all the radiolocations, was 345 km², but the area used during each period was smaller. During pre-breeding, foraging bats gathered at two stretches of different tributary rivers; during lactation, they scattered throughout the river system; and during weaning, they aggregated at a stretch of the main river. Individuals on average flew 5.7 km from roosts to foraging areas, with a maximum absolute distance of 22.7 km. Individual foraging ranges were measured linearly, because the bats foraged mostly along rivers; their values averaged 1.3 km/night and overlapped extensively between neighbouring bats (>65% on average). The sampling period, rather than the bats’ reproductive status, age, or sex, explained the observed variability in spatial distribution and size of hunting sites. We did not find differences in spatial parameters between lactating females and non-lactating bats, nor between juveniles and adults. This is the first study to split the independent effects of season and population class in order to enable unconfounded interpretations of the spatial dynamics of foraging reproductive females and juveniles. We speculate that the relationship between colony size and prey availability ruled the observed changes in foraging area through seasons. The considerable overlap in individual foraging ranges may be a necessary adaption to large colonies forced by the specific roost requirements of the long-fingered bat and the narrow foraging niche they appear to occupy.     

Influences of Landscape Features on Bat Activity in North Dakota

Land conversion and modification threatens many wildlife and plant species in the northern Great Plains, including bats. Our objective was to assess the association of bat species with landscape features in the northern Great Plains of North Dakota, USA, taking the first step towards understanding the habitat needs of bats in this region. We examined patterns of bat activity across different landscapes, identified those landscape features associated with high levels of bat activity, and determined which specific land features (i.e., vegetation and water types) were most commonly associated with each bat species. We used passive acoustic monitoring to measure bat activity at sites across North Dakota, and assessed detailed land characteristics at each site. We used nonmetric multidimensional scaling and multivariate regression tree analysis to examine relationships between bat activity and landscape variables. Bat foraging activity was influenced by structural landscape characteristics and the availability of specific water resources. High levels of bat activity were associated with riparian forested areas of varying structural complexity, ponds, and, to a lesser extent, open riparian lands. Individual bat species were influenced by land type and water resources differently. We identified big brown bats (Eptesicus fuscus) and little brown bats (Myotis lucifugus) as indicators of open riparian and pond landscapes, respectively. These results highlight the importance of prairie riparian landscapes and maintaining heterogeneity across the landscape for conservation and management of bat communities. Further, we identified ponds as an important landscape feature for little brown bats, a species of conservation concern, indicating that this specific feature should be a focus of conservation efforts on prairie wetlands. © 2019 The Wildlife Society.     

Patterns of island occupancy in bats: influences of area and isolation on insular incidence of volant mammals

Aim  The influence of landscape structure on the distribution patterns of bats remains poorly understood for many species. This study investigates the relationship between area and isolation of islands and the probability of occurrence of six bat species to determine whether persistence and immigration abilities vary among bat species and foraging guilds.
Location  Thirty-two islands in the Gulf of California near the Baja California peninsula in north-west Mexico.
Methods  Using logistic regression and Akaike information criterion (AIC) model selection, we compared five a priori models for each of six bat species to explain patterns of island occupancy, including random patterns, minimum area effects, maximum isolation effects, additive area and isolation effects and compensatory area and isolation effects.
Results  Five species of insectivorous bats ( Pipistrellus hesperus , Myotis californicus , Macrotus californicus , Antrozous pallidus and Mormoops megalophylla ) displayed minimum area thresholds on incidence. The probability of occurrence tended to decrease at moderate distances of isolation ( c . 10–15 km) for these species (excepting A. pallidus ). The distributions of two non-insectivorous species ( Leptonycteris curasoae and Myotis vivesi ) were not influenced by island size and isolation.
Main conclusions  Minimum area thresholds on incidence suggest that island occupancy by insectivorous bats may be limited by resource requirements. Islands smaller than 100 ha typically did not support occupancy or use by insectivorous bats, except at minimal isolation distances. Insectivorous bat species may also be more sensitive to moderate levels of habitat isolation in some landscape contexts than previously expected. Our results suggest that differences in foraging habits may have important implications for understanding the distribution patterns of bats.     

Foraging activity of Rhinolophus hipposideros on the Island of Herrenchiemsee, Upper Bavaria

We studied the foraging behaviour of Rhinolophus hipposideros on the island “Herrenchiemsee” in Lake Chiemsee (Upper Bavaria) during summer 2001. The island offers extensively managed woodlands, highly structured open landscapes and a broad reed belt around the shore. On average the flight activity of the 6 radio tracked females outside the roost lasted 229 min per night. The home range size varied between 6.8 and 62.7 ha (mean 25.2 ha). The size of the activity centres varied between 2.8 and 8.2 ha (mean 5.3) and all except one were located almost exclusively in woodland. Within woodlands the bats did not select for specific spatial structures (different age classes of the stands or canopy densities). Only two bats regularly foraged in additional habitats outside woodlands. One of these bats used orchards and tree rows; the other foraged over artificial ponds and gardens adjoining to its woodland foraging area. We never found the bats foraging over the lake or the reed belt. Longer linear landscape elements as tree lines were used during commuting flights but there was no indication of a continuous foraging activity along these elements. Two females left the island to forage on the mainland in August after the fledging of juveniles. To reach the mainland shore, the bats had to fly at least 1.2 km across the lake.

Assuming that most foraging flights on the island occur in woodlands, a bat density in this habitat type of 0.7 bats/ha can be calculated.     

Eating local: influences of habitat on the diet of little brown bats (Myotis lucifugus)

We employ molecular methods to profile the diet of the little brown bat, Myotis lucifugus, and describe spatial and temporal changes in diet over their maternity season. We identified 61 prey species of insects and 5 species of arachnid. The largest proportion of prey (～32%) were identified as species of the mass-emerging Ephemeroptera (mayfly) genus Caenis. Bats roosting in agricultural settings had lower dietary richness than those occupying a roost located on a forest fragment in a conservation area. We detected temporal fluctuations in diet over the maternity season. Dipteran (fly) species dominated the diet early in the season, replaced later by species of mayfly. Because our methodology provides species-level identification of prey, we were able to isolate environmental indicator species in the diet and draw conclusions about the location and type of their foraging habitat and the health of these aquatic systems. The species detected suggested that the bats use variable habitats; members of one agricultural roost foraged on insects originating in rivers or streams while those in another agricultural roost and the forest roost fed on insects from pond or lake environments. All source water for prey was of fair to good quality, though no species detected are intolerant of pollution thus the habitat cannot be classified as pristine. Our study outlines a model system to investigate the abiotic and biotic interactions between habitat factors through this simple food chain to the top predator.     

Social foraging in vampire bats is predicted by long-term cooperative relationships

Stable social bonds in group-living animals can provide greater access to food. A striking example is that female vampire bats often regurgitate blood to socially bonded kin and nonkin that failed in their nightly hunt. Food-sharing relationships form via preferred associations and social grooming within roosts. However, it remains unclear whether these cooperative relationships extend beyond the roost. To evaluate if long-term cooperative relationships in vampire bats play a role in foraging, we tested if foraging encounters measured by proximity sensors could be explained by wild roosting proximity, kinship, or rates of co-feeding, social grooming, and food sharing during 21 months in captivity. We assessed evidence for 6 hypothetical scenarios of social foraging, ranging from individual to collective hunting. We found that closely bonded female vampire bats departed their roost separately, but often reunited far outside the roost. Repeating foraging encounters were predicted by within-roost association and histories of cooperation in captivity, even when accounting for kinship. Foraging bats demonstrated both affiliative and competitive interactions with different social calls linked to each interaction type. We suggest that social foraging could have implications for social evolution if “local” within-roost cooperation and “global” outside-roost competition enhances fitness interdependence between frequent roostmates.

A combination of captive experiments and proximity sensing in the wild show that social bonds in vampire bats that are typically defined by cooperative interactions within the roost also extend beyond the roost and may provide benefits during foraging.     

Day roost selection in female Bechstein's bats (Myotis bechsteinii): a field experiment to determine the influence of roost temperature

The decision where to live has far-reaching fitness consequences for animals. In contrast to most other mammals or birds that use sheltered nest sites, female Bechstein's bats frequently switch day roosts during one breeding season, and therefore must often decide where to spend the day. Selecting the right roost is important, because roost quality, e.g. microclimatic condition, influences survival and reproduction in bats. Although thermal factors are very important for the quality of roosts occupied by bats, whether bats base their day roost selection directly on roost temperature has not been tested in the field. Over one summer, we examined and tested the roost choice of 21 individually marked female Myotis bechsteinii living in one maternity colony. In a field experiment, we allowed the bats to choose between relatively warm versus cold bat boxes, while controlling for site preferences. We expected females to exhibit a preference for warm roosts during pregnancy and lactation to accelerate gestation and shorten the period of growth of their young. Roost occupancy over 160 census days reflected significant temperature differences among 89 surveyed roosts (14 tree holes and 75 bat boxes), and preferences changed with the season. Females significantly preferred cold roosts before parturition, whereas post-partum, they significantly favoured warm roosts. Temperature preferences were independent of the roost site, and thus roost selection was based directly on temperature. Boxes with significantly different daytime temperatures did not differ significantly at night. Consequently, bats would have to spend at least 1 day in a new roost to test it. Information transfer among colony members might facilitate knowledge of roost availability. Access to many roosts providing different microclimates is likely to be important for successful reproduction in the endangered Bechstein's bat.     

Flight performance, foraging tactics and echolocation in the trawling insectivorous bat Myotis adversus (Chiroptera: Vespertilionidae)

The bat Myotis adversus hunts for prey by aerial hawking and by taking prey from the water surface with its feet (trawling). The flight performance and echolocation of this species were studied in Queensland, Australia, and comparisons were made with Myotis daubentoni , a bat filling a similar ecological niche in the Palaearctic Region. The bats foraged in very similar ways, using the same foraging tactics and feeding in similar habitats, yet they were not geometrically similar in shape. The slightly larger Myotis adversus had relatively larger wings than M. daubentoni , conferring a slightly lower wing-loading. Nevertheless, M. adversus flew faster than M. daubentoni during the searching phase of foraging. Myotis daubentoni turned in tighter circles than M. adversus . Both species used short frequency-modulated (FM) echolocation calls of a characteristic sigmoidal structure, and nulls typically observed in the calls were an observational artefact. Myotis adversus also adopted an unusual 'long'FM call while foraging. The relations between echolocation frequencies and body size were explored in male M. adversus . Specialized morphological and acoustic adaptations for prey capture by trawling in insectivorous bats are discussed.     

Impact of landscape fragmentation on a specialised woodland bat,Rhinolophus hipposideros

For habitat specialists, fragmentation has major consequences as it means less suitable habitat for the species to live in. In a fragmented landscape, we would expect larger, but spatially more clustered, foraging ranges. We studied the impact of landscape fragmentation on the foraging range and habitat exploitation of a specialised forest bat by radiotracking 16 female lesser horseshoe bats Rhinolophus hipposideros in a landscape with connected woodland structures and in a highly fragmented landscape in Carinthia, Austria.Contrary to our expectations, spatial foraging behaviour was not influenced by fragmentation. No differences in the behaviour of the bats between the sites were evident for the foraging ranges (minimum convex polygon, MCP), the core foraging areas (50% kernel), nor the mean or the maximum distances from the roost. However, in the highly fragmented landscape, the foraging activity of individuals was spatially more clustered and the overall MCP of all bats of a colony was greater compared to the less fragmented landscape.Woodland was the most important foraging habitat for the lesser horseshoe bats at both study sites. Habitat selection at the individual MCPs was evident only at the site with low fragmentation. However, in the core foraging areas, woodland was significantly selected over all other habitat types at both study sites.We conclude that (1) conservation measures for colonies of lesser horseshoe bats should be undertaken within 2.5 km of the nursery roost, (2) woodland is the key foraging habitat particularly in the vicinity of the roost, and (3) any loss of woodland near the colonial roosts are likely to negatively influence the colony, since these bats do not seem to be able to adapt their spatial foraging behaviour in a degraded landscape. The inflexible spatial behaviour of this specialised bat highlights the need to compensate for any habitat loss within the foraging range of a bat colony.