Population density of the greater mouse-eared bat (Myotis myotis), local diet composition and availability of foraging habitats

We studied the regional variation in population density of Myotis myotis (Borkhausen 1797) in south-eastern Bavaria, Germany, and its relations to diet composition and the availability of potential foraging habitats. We monitored colony size and juvenile mortality from 1991 to 2003, conducted faecal analyses in 1993 and determined land-use patterns around colonies. The numbers of individuals counted in the nursery colonies showed only small fluctuations over the years. However, data on colony size demonstrated a pronounced regional variation. Epigeic arthropods, mainly Carabidae, were the most important prey. The diet included prey taxa of forest as well as grassland habitats. The percentage of those prey taxa in the diet that originated in grassland managed with different intensity varied according to the availability of these potential foraging habitats around the nursery roosts. The calculated population density of the bats was positively correlated to forest area and especially to the area of mixed forest around the nursery roosts. Our results indicate that the availability of foraging habitats is a limiting factor for local population densities in the greater mouse-eared bat.     

What you need is what you eat? Prey selection by the bat Myotis daubentonii

Optimal foraging theory predicts that predators are selective when faced with abundant prey, but become less picky when prey gets sparse. Insectivorous bats in temperate regions are faced with the challenge of building up fat reserves vital for hibernation during a period of decreasing arthropod abundances. According to optimal foraging theory, prehibernating bats should adopt a less selective feeding behaviour – yet empirical studies have revealed many apparently generalized species to be composed of specialist individuals. Targeting the diet of the bat Myotis daubentonii, we used a combination of molecular techniques to test for seasonal changes in prey selectivity and individual‐level variation in prey preferences. DNA metabarcoding was used to characterize both the prey contents of bat droppings and the insect community available as prey. To test for dietary differences among M. daubentonii individuals, we used ten microsatellite loci to assign droppings to individual bats. The comparison between consumed and available prey revealed a preference for certain prey items regardless of availability. Nonbiting midges (Chironomidae) remained the most highly consumed prey at all times, despite a significant increase in the availability of black flies (Simuliidae) towards the end of the season. The bats sampled showed no evidence of individual specialization in dietary preferences. Overall, our approach offers little support for optimal foraging theory. Thus, it shows how novel combinations of genetic markers can be used to test general theory, targeting patterns at both the level of prey communities and individual predators.     

Reintroduction of beavers <Emphasis Type="Italic">Castor fiber</Emphasis> may improve habitat quality for vespertilionid bats foraging in small river valleys

Beavers strongly modify their environment by not only building dams and creating ponds, which slow the water flow, but also by selective cutting and removing of trees, which change the spatial structure of the forest. We aimed to test the hypothesis that beaver activity promotes new foraging sites for insectivorous bats. The beaver’s influence can be especially significant on aerial hawkers that prefer moderate structural clutter, like the Pipistrellus species (by creating new canopy gaps), and on water-surface foragers, like Myotis daubentonii (by creating ponds with smooth water surface). The study was conducted on small streams in forest areas of northern Poland, which were colonized by the European beaver (Castor fiber). Bat activity was recorded with a Pettersson D-980 ultrasound detector on line transects. The number of bat passes was significantly higher in the stream sections modified by beavers (flooded and subjected to intensive tree cutting) than in the unmodified sections (for Pipistrellus nathusii, Pipistrellus pipistrellus, Pipistrellus pygmaeus, Nyctalus noctula, and all species lumped together). Contrary to our expectations, the activity of the Myotis species was significantly lower on the transect with the largest beaver ponds, possibly due to the blanket of duckweed (Lemnaceae), which is known to produce clutter echoes, thereby reducing prey detection by echolocating M. daubentonii.     

Analysis of the diet of Daubenton’s batMyotis daubentonii in Ireland

The diet of Daubenton’s batMyotis daubentonii (Kuhl, 1817), which takes prey by aerial hawking and from the surface of water, was investigated by analysis of faeces collected in summer at 7 roosts, all close to rivers in pastoral land, in three widely-separated districts in Ireland. Forty-seven categories of arthropod prey were identified; several were insect taxa found in and around water. Most categories were recovered at most roosts, but mainly in small amounts. The main categories were the same throughout, accounting for 82% of the diet by percentage frequency in droppings for pooled data: Chironomidae/Ceratopogonidae 24% (adults 14%, preadult stages 10%), other nematoceran Diptera 21%, other Diptera 10%, and Trichoptera 26% (adults 20%, preadult 6%). A quarter of the prey had evidently been obtained from the water’s surface (eg aquatic insects, their larvae and pupae). Although consumption of several food items varied significantly by month at one or more roosts, little of such variation was consistent between roosts.     

Use of Forest Edges by Bats in a Managed Pine Forest Landscape

ABSTRACT Forest edges often have increased species richness and abundance (edge effect) and affect spatial behaviors of species and dynamics of species interactions. Landscapes of intensively managed pine (Pinus spp.) stands are characterized by a mosaic of patches and linear forest edges. Managed pine forests are a primary landscape feature of the southeastern United States, but the effects of intensive management on bat communities are poorly understood. Insectivorous bats are important top predators in nocturnal forest food webs. We examined bat foraging behavior along forest edges and in 4 structurally distinct stand types (open-canopy pine, prethinned pine, thinned pine, and unmanaged forest) within a managed pine forest in the coastal plain of North Carolina, USA. During May-August, 2006 and 2007, we recorded echolocation calls using Pettersson D240X bat detectors linked to digital recorders at 156 sites. We also sampled nocturnal flying insects at each site using Malaise insect traps. We used negative binomial count regression models to describe bat foraging behavior relative to forest edges, stand types, and prey availability. Although some species showed affinities for certain stand types and prey items, bat activity patterns were most strongly related to forest edges. Edges were used extensively by 6 aerial-hunting bat species, but avoided by Myotis species. Forest edges function similarly to natural forest gaps, by providing foraging opportunities for aerial-hunting bat species. Therefore, the maintenance of forest edges in managed pine landscapes may enhance foraging habitat for aerial-hunting bat species.     

Aggregative response in bats: prey abundance versus habitat

In habitats where prey is either rare or difficult to predict spatiotemporally, such as open habitats, predators must be adapted to react effectively to variations in prey abundance. Open-habitat foraging bats have a wing morphology adapted for covering long distances, possibly use information transfer to locate patches of high prey abundance, and would therefore be expected to show an aggregative response at these patches. Here, we examined the effects of prey abundance on foraging activities of open-habitat foragers in comparison to that of edge-habitat foragers and closed-habitat foragers. Bat activity was estimated by counting foraging calls recorded with bat call recorders (38,371 calls). Prey abundance was estimated concurrently at each site using light and pitfall traps. The habitat was characterized by terrestrial laser scanning. Prey abundance increased with vegetation density. As expected, recordings of open-habitat foragers clearly decreased with increasing vegetation density. The foraging activity of edge- and closed-habitat foragers was not significantly affected by the vegetation density, i.e., these guilds were able to forage from open habitats to habitats with dense vegetation. Only open-habitat foragers displayed a significant and proportional aggregative response to increasing prey abundance. Our results suggest that adaptations for effective and low-cost foraging constrains habitat use and excludes the guild of open-habitat foragers from foraging in habitats with high prey abundance, such as dense forest stands.     

Three‐dimensional stratification pattern in an old‐growth lowland forest: How does height in canopy and season influence temperate bat activity?

The study of animal–habitat interactions is of primary importance for the formulation of conservation recommendations. Flying, gliding, and climbing animals have the ability to exploit their habitat in a three‐dimensional way, and the vertical canopy structure in forests plays an essential role for habitat suitability. Forest bats as flying mammals may seasonally shift their microhabitat use due to differing energy demands or changing prey availability, but the patterns are not well understood. We investigated three‐dimensional and seasonal habitat use by insectivorous bats in a temperate lowland old‐growth forest, the Belovezhskaya Pushcha in Belarus. We acoustically sampled broadleaved and mixed coniferous plots in the forest interior and in gaps in three heights during two reproductive periods (pregnancy/lactation vs. postlactation). In canopy gaps, vertical stratification in bat activity was less pronounced than in the forest interior. Vertical activity patterns differed among species. The upper canopy levels were important foraging habitats for the open‐space forager guild and for some edge‐space foragers like the Barbastelle bat Barbastella barbastellus and the soprano pipistrelle Pipistrellus pygmaeus. Myotis species had highest activity levels near the ground in forest gaps. Moreover, we found species‐dependent seasonal microhabitat shifts. Generally, all species and species groups considered except Myotis species showed higher activity levels during postlactation. Myotis species tended toward higher activity in the forest interior during postlactation. P. pygmaeus switched from high activity levels in the upper canopy during pregnancy and lactation to high activity levels near the ground during postlactation. We conclude that a full comprehension of forest bat habitat use is only possible when height in canopy and seasonal patterns are considered.     

The influence of wave exposure on the foraging activity of marine otter, <Emphasis Type="Italic">Lontra felina</Emphasis> (Molina, 1782) (Carnivora: Mustelidae) in northern Chile

The marine otter Lontra felina has been said to prefer wave-exposed habitats over more protected sites in response to a greater prey abundance in exposed habitat. We examined how the foraging activity of L. felina is affected by the regime of wave exposure and prey availability at Isla Choros, northern Chile. Through focal sampling we recorded time spent by otters in foraging, the duration of dives, and the hunting success on a wave-exposed and a wave-protected site on the island. In addition, we quantified the abundance of prey in both habitats. Marine otters spent more time foraging in the wave-protected site compared with the wave-exposed habitat. Successful dives reached 26.9% in the wave-exposed habitats, and 38.2% in the wave-protected habitat. Foraging dives were 18% shorter in wave-exposed as compared with wave-protected habitat. Numerically, available prey did not differ significantly with habitat. Our results are more consistent with the hypothesis that wave-exposed habitats represent a sub-optimal habitat to foraging marine otters. Marine otters’ use of wave-exposed patches through northern and central Chile coastal areas probably reflects a low availability of suitable protected areas and greater human disturbance of more protected habitat.     

Habitat Use,Roost Selection and Conservation of Bats in Tsingy De Bemaraha National Park,Madagascar

Although the land mammals of Madagascar have been the subject of many studies, the island’s bats have yet to feature prominently on the research or conservation agenda. In this study we used mist nets, acoustic sampling and cave surveys to assess habitat use, seasonality and roost selection. Four microchiropteran species (Triaenops rufus, T. furculus, Miniopterus manavi and Myotis goudoti) appeared to be strongly associated with the forest interior based on trapping, but analysis of time-expanded echolocation recordings revealed that T. rufus and M.␣manavi were frequently recorded in forest edges and clearings. Bat activity was significantly lower inside the forest than at the interface between agricultural land and forest. The caves visited most often by tourists were low in bat abundance and species richness. Anjohikinakina Cave, which was visited infrequently by people, was used by five species and contained between 54% (winter) and 99% (summer) of bats counted in 16 caves and is a site of national importance for bat conservation. Hipposideros commersoni was only netted in our study area during October and may be a migrant to the site or present but inactive during the austral winter. The forest surrounding the caves is therefore important because it provides cover for emerging bats and a potential source of invertebrate prey whilst the forest edge is important to foraging bats.     

Constructed wetlands support bats in agricultural landscapes

Bats are known to use aquatic habitats as foraging habitats. Agricultural intensification is perceived to be a main reason for the loss of wetlands. However, artificial wetland creation (i.e. the construction of retention-ponds) in the agricultural landscape aiming at water or nutrient retention has recently gained importance. We evaluated to what extent bats use these artificial wetlands as foraging habitats in an agricultural landscape.Bat activity and prey density were compared in matched pairs at retention-ponds and neighbouring vineyard sites using stationary bat-detectors and sticky-traps, respectively. To examine if bat activity is related to the number of bat individuals, a thermal infrared imaging camera was used. Pipistrellus pipistrellus, the dominant species, served as an example to assess habitat selection between retention-ponds and vineyards. This was performed by relating foraging activity to the available area available within the potential home-range.Total bat activity and nocturnal prey density were significantly higher above the retention-ponds than above vineyards. High differences of activity levels between the ponds and the respective vineyard sites were found for Pipistrellus spp. (P. pipistrellus and P. nathusii) and Myotis spp. (M. daubentonii and M. mystacinus), being about 180 times and 50 times higher above the retention-ponds, respectively. A significant correlation was found between recorded bat activity and the maximum number of bat individuals observed with a thermal infrared imaging camera. When relating foraging activity to habitat availability within the assumed home-range of P. pipistrellus, retention-ponds had on average a higher importance as a foraging habitat than the complete vineyard area although they covered less than 0.1% of its area.This study indicates that artificial wetlands such as retention-ponds provide foraging habitats for bats. Therefore, creation of wetlands in intensively used agricultural landscapes benefits bats.     

Correction factors used for estimating prey biomass in the diet of American minkMustela vison

Feeding experiment on nine feral minkMustela vison Schreber, 1777 was carried out to find differences in digestibility of particular prey types. The values of correction factors (CF), which show the ratio between consumed prey biomass and undigested remains, were calculated for seven food types. The experiment revealed that results obtained by commonly used percentage of occurrence methods differed from the real intake of food. Food types which were intensively digested were underestimated by percentage of occurrence methods and prey which had hardly digestive elements of their body were overestimated. The calculated values of CF are as follows: crayfish — 14.8, fish — 30.8, frog — 61.3, small passerine bird — 17.2, chicken — 41.3, rodent — 17.3, egg — 687.5. The variability in CF values did not depend on mink individuals but could be explained by the mean weight of prey items eaten by mink. The positive correlation between the prey size and its digestibility was recorded for crayfish, fish and chicken. The biomass of frog, passerine and rodent remains was not related to the body mass of eaten prey. The use of evaluated correction factors enables more accurate estimates of the food composition of wild living mustelids.     

Flexibility in Diet and Activity Patterns of <Emphasis Type="Italic">Macaca tonkeana</Emphasis> in Response to Anthropogenic Habitat Alteration

I examined how Sulawesi Tonkean macaques (Macaca tonkeana) respond in terms of their diet and activity patterns to anthropogenic habitat alteration in Lore Lindu National Park, Sulawesi, Indonesia. Anthropogenic habitat alteration comprised clearing of forest for agriculture and small-scale forest product collection. I quantified the diet and activity of 2 groups (CH and Anca), occupying habitats with different levels of alteration, via scan sampling. Tree abundance, key food specific density, and fruit production were greater in the minimally altered habitat (CH), substantiating the characterization of the group’s habitat as higher quality. For the group in the heavily altered habitat (Anca), alternative foods, e.g., insects, fungus, young and mature leaves, shoots, and stems, accounted for a significantly greater proportion of the diet. Dietary diversity is significantly lower in the Anca group, with 52% of their diet being palm fruits from Arenga pinnata. The activity patterns of the Anca group—more time foraging, less time moving, and more time resting than the CH group—reflect the lower resource availability in their habitat and their reliance on more alternative food items, coupled with their extremely small group size (6–9 individuals). The group may be at the optimal size in which foraging efficiency is maximized for the habitat, a response, in conjunction with dietary and behavioral flexibility, to alteration of their habitat. The results are contextualized with respect to the conservation value of human-modified landscapes.     

Analysis of the diet of Natterer's bat Myotis nattereri and the common long-eared bat Plecotus auritus in the West of Ireland

The diets of Natterer's bat Myotis nattereri and the common long-eared bat Plecotus auritus were investigated at a nursery colony of each species by analysis of droppings collected monthly from May to September. Respectively, 68% and 42% of the diets comprised items presumed to have been gleaned from foliage or other surfaces: diurnal insects, insects which rarely fly, and non-flying arthropods. Such surfaces included the ground: centipedes were eaten by both bats, and the muscid, Scatophaga stercoraria , usually associated with cattle dung, was a common prey. Indeed, the prevalence of cattle-farming around both of the bat roosts almost certainly contributed indirectly, in the form of S. stercoraria , to a significant part of the Diptera consumed. The chief food of M. nattereri was the larger Diptera, but Trichoptera, Hymenoptera and Arachnida were also important. Lepidoptera, Coleoptera and Hemiptera were minor prey, with Dermaptera and Chilopoda taken occasionally. Diptera, closely followed by Lepidoptera, together accounted for nearly two-thirds of the diet of P. auritus. Also taken, in descending order of importance, were Trichoptera, Arachnida, Chilopoda, Coleoptera, Dermaptera, Hymenoptera and Hemiptera. Plecotus auritus , but not M. nattereri , was evidently able to take a few small insects such as aphids and lesser nematoceran Diptera.     

Bats in a changing landscape: Linking occupancy and traits of a diverse montane bat community to fire regime

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Long-term changes in abundance of bats as revealed by their frequency in tawny owls’ diet

Percentage of bats in tawny owls’ diet was compared in three periods: I — before 1976, II — 1976–1992, III — 1993–2009, by using the published and unpublished material from Poland (only samples over 100 vertebrate prey items). This species of owl showed an opportunistic predation on bats and took them more frequently in periods of higher abundance. Before the mass use of toxic pesticides in Poland, in the period I bats constituted more than 2% of vertebrates in four out of five diet samples (median 2.4%). The lowest bat abundance occurred in Poland in the 1980s and resulted in the lowest percentage of bats taken by owls in the period II (n = 11, median 0.2%). Due to the recovery of bat populations in the period III, the percentage of bats in tawny owls’ diet increased (n = 23, median 0.7%). In large samples (over 200 vertebrate items, n = 21) collected in central and north-eastern Poland the percentage of bats increased from 1980 to 2009 (the estimated average value at the end of that period slightly exceeded 1%). Samples collected at the same five sites in 1975–1992 and again in 2000–2009, confirmed the increasing trend in percentage of bats captured by tawny owls noted in last years.     

Bat activity and species richness in different land‐use types in and around Chome Nature Forest Reserve,Tanzania

Bats have important ecological roles in ecosystems, but many species are threatened because of anthropogenic impacts. Tanzania has limited information on how bats respond to habitat modification. This makes it difficult to anticipate which bat species are at risk. Bat activity and species richness were assessed in five land‐use types: forest and banana–coffee (upland habitats), rice paddy, riverine and sisal estate (lowland habitats). Mist nets, harp traps and bat detectors were used to sample bats. Species richness differed between habitats. Bat activity levels were higher in lowland habitats than upland habitats. Riverine and rice paddy habitats were shown to have an important role as foraging sites for many insectivorous bats as bat species richness and activity were generally higher than other habitats. Fruit‐eating bats preferred riverine and banana–coffee habitats. We recommend using organic manure as alternatives to chemical fertilisers, and pesticide use should be avoided in rice paddies. Riparian vegetation along rivers and water bodies should be maintained as important faunal nesting, roosting and/or foraging grounds. The requirement that farming practices be at least 60 m from the river should be strictly enforced. These recommendations will help in the conservation of bats and their habitats in modified agricultural landscapes.     

Assessing niche partitioning of co‐occurring sibling bat species by DNA metabarcoding

Niche partitioning through foraging is a mechanism likely involved in facilitating the coexistence of ecologically similar and co‐occurring animal species by separating their use of resources. Yet, this mechanism is not well understood in flying insectivorous animals. This is particularly true of bats, where many ecologically similar or cryptic species coexist. The detailed analysis of the foraging niche in sympatric, cryptic sibling species provides an excellent framework to disentangle the role of specific niche factors likely involved in facilitating coexistence. We used DNA metabarcoding to determine the prey species consumed by a population of sympatric sibling Rhinolophus euryale and Rhinolophus mehelyi whose use of habitat in both sympatric and allopatric ranges has been well established through radio tracking. Although some subtle dietary differences exist in prey species composition, the diet of both bats greatly overlapped (O_jk = 0.83) due to the consumption of the same common and widespread moths. Those dietary differences we did detect might be related to divergences in prey availabilities among foraging habitats, which prior radio tracking on the same population showed are differentially used and selected when both species co‐occur. This minor dietary segregation in sympatry may be the result of foraging on the same prey‐types and could contribute to reduce potential competitive interactions (e.g., for prey, acoustic space). Our results highlight the need to evaluate the spatial niche dimension in mediating the co‐occurrence of similar insectivorous bat species, a niche factor likely involved in processes of bat species coexistence.     

Shade cacao plantations (<Emphasis Type="Italic">Theobroma cacao</Emphasis>) and bat conservation in southern Bahia,Brazil

Nearly 40% of the remaining Atlantic Forest in southern Bahia state, Brazil, is a rustic agroforest of cacao (Theobroma cacao). These traditional shade plantations, locally known as ‘cabrucas’, are habitats for forest dwelling species. Here we investigated the potential role of this traditional crop as a support for bat assemblages in southern Bahia, establishing the extent to which the bat community structure found in this agricultural system is influenced by the proximity of forest tracts. We compared the bat community attributes (richness, diversity and dominance) and species-level response (capture frequency) between native forest tracts and shade cacao plantations located in two distinct landscapes, which varied in total amount of forest (patch sizes and total forested area) and in their proximity to forest tracts. The cabrucas contiguous to forest tracts showed a rich and abundant bat community, with samples showing capture rates, species richness, diversity and evenness significantly higher than those reported for forest tracts. This situation changes, however, when shade plantations are located at some distance from forest tracts ( > 1000 m). Bat communities in these isolated cabrucas are less diverse than those found in forests and nearby cabrucas, but in both cases, species usually associated with pristine habitats were found. These findings suggest that cabrucas per se are not forest surrogates, and their potential to harbor forest dwelling bat species is closely linked with the existence of nearby forest tracts that may act as a source for species populations. Therefore, the entire landscape should be considered for management, taking into account that maintenance of cabrucas together with the preservation and restoration of forest patches is probably direly needed if one wishes to conserve the bat diversity in southern Bahia for the long term.     

A mosaic of opportunities? Spatio-temporal patterns of bat diversity and activity in a strongly humanized Mediterranean wetland

The Baixo Vouga Lagunar (BVL) landscape, in the Portuguese central-west coast, harbours a mosaic of wetland habitat types, interspersed by intensive and extensive agricultural fields, pastures, production forests and urban areas. In this study, we aimed to determine the species composition and the structure of the bat assemblages of the different habitats that constitute this heterogeneous landscape and to investigate seasonal changes in the patterns of bat diversity and activity across habitats. We acoustically sampled bats across 24 sampling sites representative of the eight main habitat types that shape the landscape—Bocage, forests, maize fields, marshlands, reed beds, rice fields, sea rushes and urban settlements. We compared bat richness, diversity and evenness across habitat types and seasons. We analysed habitat-specific and season-specific overall bat activity, and because habitat selection by bats is known to reflect morphological characters, foraging strategies and echolocation call structures, we also analysed the activity of individual species and of eco-morphological guilds. From 1,544 bat-passes recorded, we identified 12 species. Pipistrellus pygmaeus, Pipistrellus pipistrellus and Eptesicus serotinus/Eptesicus isabellinus were the most frequently recorded. Species composition and activity were similar across habitats, whilst exhibiting strong seasonal dynamics within habitats. Our results suggest that the mosaicism of the landscape provides several opportunities for bats, enabling them to explore different resources in distinct habitat patches. However, it may also reflect a forced exploitation of less optimal habitats and resources by bats, due to the scarcity of opportunities provided by fragmented landscapes.     

Numerical and Functional Responses of Forest Bats to a Major Insect Pest in Pine Plantations

Global change is expected to modify the frequency and magnitude of defoliating insect outbreaks in forest ecosystems. Bats are increasingly acknowledged as effective biocontrol agents for pest insect populations. However, a better understanding is required of whether and how bat communities contribute to the resilience of forests to man- and climate-driven biotic disturbances. We studied the responses of forest insectivorous bats to a major pine defoliator, the pine processionary moth pityocampa, which is currently expanding its range in response to global warming. We used pheromone traps and ultrasound bat recorders to estimate the abundance and activity of moths and predatory bats along the edge of infested pine stands. We used synthetic pheromone to evaluate the effects of experimentally increased moth availability on bat foraging activity. We also evaluated the top-down regulation of moth population by estimating T. pityocampa larval colonies abundance on the same edges the following winter. We observed a close spatio-temporal matching between emergent moths and foraging bats, with bat activity significantly increasing with moth abundance. The foraging activity of some bat species was significantly higher near pheromone lures, i.e. in areas of expected increased prey availability. Furthermore moth reproductive success significantly decreased with increasing bat activity during the flight period of adult moths. These findings suggest that bats, at least in condition of low prey density, exhibit numerical and functional responses to a specific and abundant prey, which may ultimately result in an effective top-down regulation of the population of the prey. These observations are consistent with bats being useful agents for the biocontrol of insect pest populations in plantation forests.