Use of native woodlands and traditional olive groves by foraging bats on a Mediterranean island: consequences for conservation

We recorded bat activity on Zakynthos island (Greece) to test the hypotheses that (1) olive ( Olea europea ) groves and native woodlands provide comparable foraging habitat for insectivorous bats, (2) lower foraging activity occurs in olive groves treated with insecticide chemicals. We acoustically sampled bat activity (passes per minute) in four wooded habitats (organic and non-organic olive groves, oak woodland ( Quercus ilex and Quercus coccifera ) and pine ( Pinus halepensis ) woodland from June to August 2005. Habitat type did not affect overall bat activity. A single application of insecticide chemicals annually did not affect activity over traditional olive groves. Habitat use on the island differed in several ways from that reported in studies at mainland sites. Most strikingly, pine woodland supported higher bat activity than expected relative to other habitat types, and we recorded unexpectedly high levels of M. capaccinii activity in woodland habitats. We suggest that traditional olive groves buffer some bat species from the effects of deforestation. Conservation plans for Mediterranean bats should consider the biodiversity value of these groves along with the need to conserve small woodland patches. Finally, understanding island-specific patterns of habitat use is essential to bat conservation on small off-shore islands.     

Habitat selection as a mechanism of resource partitioning in two cryptic bat species Pipistrellus pipistrellus and Pipistrellus pygmaeus

Ecomorphological studies of bat communities often reveal the spatial and temporal coexistence of morphologically similar species, leading to suggestions that these communities are structured by non-deterministic processes. However, the diversification of echolocation call structure in bats allows for considerable morphological similarity while still permitting niche differentiation based on specialisation for prey type and habitat structure. The recent separation of a common Palaearctic bat, the pipistrelle, into Pipistrellus pipistrellus and P. pygmaeus, which are sympatrically distributed throughout their range, raises the question as to whether these two morphologically similar species partition resources in time and space.
To test the hypothesis that the coexistence of these cryptic species is facilitated by differential habitat use, 14 P . pipistrellus , and 12 P. pygmaeus were radio-tracked from adjacent maternity roosts, in northeast Scotland, from May to September 2002/2003. The two species showed distinct habitat partitioning with P. pygmaeus foraging predominantly in riparian woodland and over water, and P. pipistrellus foraging along woodland edges and short isolated tree lines. Inter-specific overlap in habitat use was low and consequently foraging ranges were segregated spatially.
The degree of habitat partitioning revealed in these species, which show considerable overlap in echolocation call parameters and functional morphology, suggests that morphological features, whilst useful in separating chiropteran species into coarse-grained foraging guilds, may not predict fine-grained ecological segregation.     

Factors affecting the distribution pattern of bats in Uppland, central Sweden

Bats were surveyed and insects were trapped in several different habitats to test the hypothesis that insect abundance and types are the most important factors affecting the regional distribution of bats. A bat community of ten species dominated by Eptesicus nilssoni and Pipistrellus pipistrellus was analysed. The results show that the general pattern of bat distribution in different habitats, and the seasonal changes in habitat utilization, could be explained in terms of the abundance of swarming insects, mainly chironomids. Deciduous woodland near water provided essential habitat in early summer for each bat species. Later on, in July, all species spread out to many different habitats. In autumn street-lamps were an important factor affecting bat distribution.     

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.     

Natterer's bats prefer foraging in broad-leaved woodlands and river corridors

We studied habitat selection by radio tracking Natterer's bats Myotis nattereri foraging in a grassland–woodland landscape. We tested the hypothesis that selection of foraging habitat is random at two levels: firstly, the selection of individual foraging ranges and secondly, the choice of foraging habitats made by individuals within these foraging ranges. Habitat selection was random at neither level. When selecting foraging ranges, bats maximized the area of semi-natural broad-leaved woodland and improved grassland and minimized that of dense coniferous plantations. During foraging, semi-natural broad-leaved woodland and river corridors were preferred, while dense coniferous plantations were avoided. Within individual foraging ranges, the intensity of foraging activity over river corridor habitat and semi-natural broad-leaved woodland was 8.2 and 3.8 times higher, respectively, than that over improved grassland. For successful management of Natterer's bat populations, semi-natural broad-leaved woodland should be retained. Clear felling of large blocks of native broad-leaved woodland should be avoided and conifers should not be used for reforestation. Tree cover along river banks should be encouraged and protected.     

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.     

Habitat usage of Daubenton's bat (Myotis daubentonii), common pipistrelle (Pipistrellus pipistrellus), and soprano pipistrelle (Pipistrellus pygmaeus) in a North Wales upland river catchment

Distributions of Daubenton's bat (Myotis daubentonii), common pipistrelle, (Pipistrellus pipistrellus), and soprano pipistrelle (Pipistrellus pygmaeus) were investigated along and altitudinal gradient of the Lledr River, Conwy, North Wales, and presence assessed in relation to the water surface condition, presence/absence of bank‐side trees, and elevation. Ultrasound recordings of bats made on timed transects in summer 1999 were used to quantify habitat usage. All species significantly preferred smooth water sections of the river with trees on either one or both banks; P. pygmaeus also preferred smooth water with no trees. Bats avoided rough and cluttered water areas, as rapids may generate high‐frequency echolocation‐interfering noise and cluttered areas present obstacles to flight. In lower river regions, detections of bats reflected the proportion of suitable habitat available. At higher elevations, sufficient habitat was available; however, bats were likely restricted due to other factors such as a less predictable food source. This study emphasizes the importance of riparian habitat, bank‐side trees, and smooth water as foraging habitat for bats in marginal upland areas until a certain elevation, beyond which bats in these areas likely cease to forage. These small‐scale altitudinal differences in habitat selection should be factored in when designing future bat distribution studies and taken into consideration by conservation planners when reviewing habitat requirements of these species in Welsh river valleys, and elsewhere within the United Kingdom.     

Hematological data and hemoglobin components in bats (Vespertilionidae)

1. Statistically significant variations were observed in the RBC counts and Hb concentration in pregnant females of Pipistrellus pipistrellus. 2. Basic hematological values in 59 animals of five species of insectivorous bats were estimated. 3. Electrophoretic separation of the hemoglobins of Plecotus austriacus, Myotis nattereri and Myotis myotis showed two components, whereas in Miniopterus schreibersi and Pipistrellus pipistrellus appeared three and four components, respectively.     

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.     

Local patterns of distribution and resource utilization of four bat species (Myotis brandti, Eptesicus nilssoni, Plecotus auritus and Pipistrellus pipistrellus) in patchy and continuous environments

The local patterns of distribution and resource utilization of four bat species ( Myotis brandti, Eptesicus nilssoni, Plecotus auritus and Pipistrellus pipistrellus ) were examined in patchy and continuous environments, using bat detectors. The effects of two different kinds of open matrix habitats (crop-fields and water) on species occurrence were compared in the patchy areas. A crop-field matrix seemed to have a greater negative influence on species occurrence than a water matrix. Presence and absence of species in the patchy areas were analysed against island area, area of some habitats, and isolation. All species were positively affected by one or more forest habitat parameters. Two species ( M. brandti and P. auritus ) were negatively affected by isolation, which suggests that they may be particularly vulnerable to increased forest patchiness. These species occurred mainly on large islands. Two hypotheses that might explain the habit of open area avoidance in M. brandti and P. auritus were tested: 1. Insect abundance hypothesis; 2. Foraging behaviour hypothesis. Both failed to explain why these two species avoid open habitats and as a consequence are negatively affected by isolation.     

Phylogeography of pipistrelle-like bats within the Canary Islands,based on mtDNA sequences

Evolution of three Canary Island Vespertilionid bat species, Pipistrellus kuhlii, Pipistrellus maderensis, and Hypsugo savii was studied by comparison of approximately 1 kbp of mtDNA (from cytochrome b and 16S rRNA genes) between islands. mtDNA reveals that both P. kuhlii and P. maderensis exist in sympatry on Tenerife (and possibly other islands). Their morphological similarity explains why their co-occurrence had not been detected previously. Levels of sequence divergence are quite low within P. maderensis. Haplotypes were either identical or separated by /=12 mutational steps) indicating colonization of the latter from the former sometime during the last approximately 1.2 Ma, with low subsequent gene flow. Unlike P. maderensis the El Hierro population alone appears to represent an ESU. The H. savii haplotypes detected in Gran Canaria and Tenerife are identical or separated by 1 mutational step.     

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.     

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.     

Studies on the prevalence of haematozoa of British bats

Bloodfilms were collected from 491 bats representing 12 of the 15 species of British bats, from a number of sites in England and Scotland, and examined for haematozoa. Trypanosoma (Schizotrypanum ) sp. were found in Pipistrelluspipistrellus, Nyctalus noctula, N. leisleri, Eptesicus serotinus and Myotis brandtii , and Trypanosoma (Megatrypanum) incertum was found in P. pipistrellus. Both subgenera of trypanosomes appear to be transmitted by the bat bug Cimex pipistrelli. Gametocytes of the haemoproteid Poly-chromophilus murinus were found in one-third of Myotis daubentonii examined, and stages of the parasite were found in the vector, the wingless blood-sucking fly, Nycteribia kolenatii , collected from these bats. The piroplasm Babesia vesperuginis was found in the blood of P. pipistrellus and Myotis mystacinus , and was the only parasite to show evidence of pathogenicity to bats.     

Echolocation calls produced by Kuhl's pipistrelles in different flight situations

Flexibility in the echolocation call structure of bats can improve their performances, because, in some situations, some signal designs are better than others. Hence, at least some bats should adjust their echolocation calls according to the setting in which they are operating but also to the specific task at hand, that is their behavioral intention. We studied variation in the echolocation calls of Pipistrellus kuhlii emitted during four flight situations that were similar in setting but differed in behavioral context: emergence from a roost, commuting to and from foraging sites, foraging and returning to a roost. Echolocation calls produced by P. kuhlii differed significantly according to the flight situation. Call types differed most distinctly between foraging and commuting. We also found a high variance in the emergence calls we recorded, perhaps reflecting pre- and post-takeoff calls. Discriminant function analysis on calls emitted while foraging, commuting or returning to the roost classified the calls to the correct group 73.3% of the time. The differences between bats' echolocation calls in different flight situations might indicate an intrinsic change in the bat's behavior. Recognizing these differences could be crucial when using call variables to identify bat species.     

Gastrointestinal helminths of pipistrelle bats (Pipistrellus pipistrellus/Pipistrellus pygmaeus) (Chiroptera: Vespertilionidae) of England

Although bats are one of the most successful and diverse of mammalian orders, studies that focus upon bat endoparasites are limited. To further knowledge of bat parasitology, pipistrelle bats (Pipistrellus pipistrellus and P. pygmaeus) were acquired from across the Greater Manchester and Lancashire region of England and examined for gastrointestinal helminths using morphological and molecular analyses. Sixty-eight of 90 adult/juvenile bats (76% prevalence) were infected with at least 1 species of helminth and mean helminth abundance was 48·2 (+/-7·0). All helminths were digenean trematodes and the following species were identified in 51 P. pipistrellus specimens (prevalence in parentheses): Lecithodendrium linstowi (80·4%), L. spathulatum (19·6%), Prosthodendrium sp. (35·3%), Plagiorchis koreanus (29·4%) and Pycnoporus heteroporus (9·8%). Statistical analyses, incorporating multifactorial models, showed that male bats exhibited a significantly more aggregated helminth distribution and lower abundance than females. Positive associations were observed between L. linstowi and L. spathulatum, Prosthodendrium sp. and P. heteroporus and between L. spathulatum and P. koreanus. A revised phylogeny of bat-associated Lecithodendriidae, incorporating novel L. spathulatum and Prosthodendrium sp. 28S rRNA sequences, separated the controversial clade formed by L. linstowi and P. hurkovaae. Further studies are likely to assist the understanding of bat-parasite/pathogen relationships, helminth infracommunity structures and phylogenetics.     

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.     

Use of lamplit roads by foraging bats in southern England

Roads illuminated by white streetlamps attracted three times more foraging bats (mostly Pipistrellus pipistrellus ) than did roads lit by orange streetlamps or unlit roads (3.2, 1.2 and 0.7 bat passes/km, respectively). More insects flew around white lamps than around orange lamps (mean 0.67 and 0.083 insects per lamp, respectively). The mean number of bat passes recorded in any 1-km section of road was positively correlated to the number of white streetlamps along the section, and also, independently, to the amount of trees and hedgerows. Bat activity was not related to the number of houses along the road, ambient temperature or cloud cover. The attractive effect of the lamps on the bats was diminished in windy weather.     

Seasonal Shifts in Nocturnal Habitat Use by Coastal Bat Species

Sensitivity of bats to land use change depends on their foraging ecology, which varies among species based on ecomorphological traits. Additionally, because prey availability, vegetative clutter, and temperature change throughout the year, some species may display seasonal shifts in their nocturnal habitat use. In the Coastal Plain of South Carolina, USA, the northern long-eared bat (Myotis septentrionalis), southeastern myotis (Myotis austroriparius), tri-colored bat (Perimyotis subflavus), and northern yellow bat (Lasiurus intermedius) are species of conservation concern that are threatened by habitat loss. Our objective was to identify characteristics of habitat used by these species during their nightly active period and compare use between summer and winter. We conducted acoustic surveys at 125 sites during May–August and at 121 of the same 125 sites December–March 2018 and 2019 in upland forests, bottomland forests, fields, ponds, and salt marsh and used occupancy models to assess habitat use. The northern long-eared bat and southeastern myotis (i.e., myotis bats) used sites that were closer to hardwood stands, pine stands, and fresh water year-round. We did not identify any strong predictors of tri-colored bat habitat use in summer, but during winter they used bottomland forests, fields, and ponds more than salt marsh and upland forests. During summer and winter, northern yellow bats used sites close to fresh water and salt marsh. Additionally, during summer they used fields, ponds, and salt marsh more than upland and bottomland forests, but in winter they used bottomland forests, fields, and ponds more than upland forest and salt marsh. Our results highlight important land cover types for bats in this area (e.g., bottomland forests, ponds, and salt marsh), and that habitat use changes between seasons. Accounting for and understanding how habitat use changes throughout the year will inform managers about how critical habitat features may vary in their importance to bats throughout the year. © 2021 The Wildlife Society.