Habitat selection of endangered and endemic large flying-foxes in Subic Bay, Philippines

Large flying-foxes in insular Southeast Asia are the most threatened of the Old World fruit bats due to high levels of deforestation and hunting and effectively little local conservation commitment. The forest at Subic Bay, Philippines, supports a rare, large colony of vulnerable Philippine giant fruit bats (Pteropus vampyrus lanensis) and endangered and endemic golden-crowned flying-foxes (Acerodon jubatus). These large flying-foxes are optimal for conservation focus, because in addition to being keystone, flagship, and umbrella species, the bats are important to Subic Bay’s economy and its indigenous cultures. Habitat selection information streamlines management’s efforts to protect and conserve these popular but threatened animals. We used radio telemetry to describe the bats’ nighttime use of habitat on two ecological scales: vegetation and microhabitat. The fruit bats used the entire 14,000 ha study area, including all of Subic Bay Watershed Reserve, as well as neighboring forests just outside the protected area boundaries. Their recorded foraging locations ranged between 0.4 and 12 km from the roost. We compared the bats’ use to the availability of vegetative habitat types, riparian areas, and bat trees. The fruit bats’ locations showed a preference for undisturbed forest types and selection against disturbed and agricultural areas. Bat locations also showed selection for particular fruiting/flowering bat trees. The bats showed strong preference for riparian areas; locations were in riparian areas over four times more than expected. From these results we recommend that management focus flying-fox conservation efforts on undisturbed forest and riparian areas.     

Habitat use and conservation of bumblebees (Bombus spp.) under different grassland management regimes

Declines in the natural populations of several bumblebee species across Britain and Europe are an increasing cause for concern. In this study the habitat use of bumblebees was investigated on Salisbury Plain Training Area, the largest remaining area of unimproved chalk grassland in north-west Europe. Habitat characteristics influencing the overall abundance, species richness and foraging activity of bumblebees included the diversity and abundance of flowering plant species (particularly of favoured forage plants such as Trifolium pratense), vegetation structure and height. It is suggested that different Bombus species respond to these habitat characteristics depending on their specific foraging and nesting requirements, the case of Bombus humilis being especially relevant. The effects of several grassland management practices were considered in terms of their suitability for the conservation of bumblebee habitats. Cattle grazing was shown to be preferable to both sheep grazing and the absence of any management, although the timing and intensity of such grazing was important. Small-scale disturbances caused by vehicle activity were also of value in producing locally abundant forage resources in less intensively managed grasslands.     

Species-specific barrier effects of a motorway on the habitat use of two threatened forest-living bat species

It is generally recognized that roads can adversely affect local animal populations but little is known how roads affect bats. In particular, no study compared the response of bats that differ in foraging ecology to motorways that cut through the breeding habitat. As bats are key species in conservation, such data are urgently needed for designing management plans. Using radio-telemetry, mist netting, and mark-recapture data we investigated the effects of a motorway with heavy traffic on the habitat use of two threatened forest-living bats. We compared barbastelle bats (Barbastella barbastellus), which forage in open space, to Bechstein’s bats (Myotis bechsteinii), which glean prey from the vegetation. Five of six radio-tracked barbastelle bats crossed the motorway during foraging and roost switching, flying through underpasses and directly over the motorway. In contrast, only three of 34 radio-tracked Bechstein’s bats crossed the motorway during foraging, all three using an underpass. Bechstein’s bats, unlike barbastelle bats, never crossed the motorway during roost switching. Moreover, only in Bechstein’s bats individuals foraging close to the motorway had smaller foraging areas than individuals foraging further away, whereas other forest edges had no such effect. Our data show that motorways can restrict habitat accessibility for bats but the effect seems to depend on the species’ foraging ecology and wing morphology. We suggest that motorways have stronger barrier effects on bats that forage close to surfaces than on bats that forage in open space, and discuss the implications of our findings for bat conservation during road construction.     

Intra-specific effects of forest loss on the distribution of the forest-dependent northern long-eared bat (Myotis septentrionalis)

Forest fragmentation by the expansion of human activities is acknowledged as an important factor driving declines of forest species worldwide and quantitatively characterizing this trend is essential to understanding the response of populations of forest-dependent species. To examine the impacts of forest cover loss and changes in composition on forest-dependent bats, we examined the effects of these factors on the distribution of the northern long-eared bat (Myotis septentrionalis) in a case study. We surveyed forest fragments in the forest-agricultural landscape of Prince Edward Island, Canada, to assess the occupancy of bats in fragments. Logistic regression and multimodel inference using Akaike’s information criteria were used to identify potentially important predictor variables influencing the distribution of the northern long-eared bat at the fragment and landscape level and quantified their effects. A compositional variable, area of deciduous stands, was found to be the best predictor of the presence of M. septentrionalis. We found the effects of fragmentation were greatest at the fragment level for females and at the landscape level for males. This case study emphasizes the importance of examining intra-specific resource selection in how it affects the response of a forest-dependent species to fragmentation.     

Scale-independent significance of river and riparian zones on three sympatric Myotis species in an agricultural landscape

Conserving biodiversity in agricultural landscapes is an urgent issue. The effective conservation of biodiversity requires plans based on species’ habitat preferences at multiple spatial scales. We examined how the foraging habitat selection of bats varied with grain size (50, 150, 250, and 350 m) and how habitat selection in the home range differed from that in the foraging habitat in an agricultural landscape. Focusing on three sympatric Myotis species (Myotis petax, Myotis gracilis, and Myotis frater), we radio-tracked 10 individuals of each species for 121 nights in Hokkaido, northern Japan. Rivers and broadleaved riparian forests were commonly preferred at multiple scales, although the preferred land-cover type was dependent on both extent and grain size for most species. The best grain sizes for predicting the foraging-habitat use of M. petax and M. gracilis were 50 and 150 m. By contrast, M. frater showed no tendency across the grain sizes. Our results indicate that it is necessary to consider both extent and grain size to understand the habitat selection of bats. Our findings also suggest that focusing primarily on preferred land-cover types at multiple scales is effective for conservation planning, given the limited resources in terms of time, manpower, and finances. Although arable did not negatively affect the selection of foraging habitats and home ranges in these bat species, preventing the loss of rivers and forests should be prioritized over preventing arable land expansion.     

Influence of wetland networks on bat activity in mixed-use landscapes

Parks and other protected lands can provide important source habitat and act as valuable dispersal corridors in urbanizing environments. However, most wetlands within protected areas are managed in isolation without consideration of the broader landscape connections. We studied the importance of wetland habitat connectivity and landscape context to bat activity in five National Parks along a gradient of increasing urbanization within the Mid-Atlantic United States. Ninety-six Anabat stations were set up throughout the parks, from which we derived the characteristic spatial scales at which bat activity was associated with wetlands. This information was used in a graph theoretic framework to construct network models of potential landscape connectivity for those species that had positive associations with wetland land cover. We found that the importance of wetlands as a predictor of bat activity varied on a species-by-species basis and increased when network measures were used that accounted for connected area in a broad spatial context. The results demonstrate that both area and connectivity of wetland foraging habitat may act as orthogonal variables to availability of roosting habitat in explaining the distribution of highly mobile species. We use the results to illustrate the value of network analysis to guide the coordinated management of two of the parks’ most valued natural resources - wetlands and bats.     

Application of central-place foraging theory shows the importance of Mediterranean dehesas for the conservation of the cinereous vulture, Aegypius monachus

The dehesa (oak woodland) is an extensive agro-pastoral ecosystem characteristic of the Western Mediterranean countries which is suffering a great transformation process since 1950. Although its distribution largely overlaps with several endangered species, there is scarce information on how they use this human-transformed habitat. We studied the foraging habitat selection of one of them, the cinereous vulture Aegypius monachus. We radio-tracked 14 cinereous vultures in one of the largest European colonies from 1998 to 2000. Used and available habitats were compared at two scales using compositional analysis. Moreover, we developed a distance-based GLMM for assessing habitat selection in this central-place forager species, by taking into account the spatial distribution of habitat patches in relation to the location of the colony. Home ranges overlapped over a total surface of 592,527 ha around the colony, and both individual home ranges and travel foraging distances (mean 27.86 km, maximum 86 km) were larger during the breeding season. All cinereous vultures avoided agricultural lands within their home ranges throughout the year. Habitat use in relation to the distance to the colony pointed out that dehesas were positively selected in spite of being on average far away from the colony than other habitats, a result that was consistent among individuals and seasons. The cinereous vulture thus depends for its conservation not only on the protection of breeding areas, as has been so far considered, but also on the maintenance of well-conserved dehesas close to the colonies. Preserving the cinereous vultures could contribute to the economic sustainability of dehesas by attracting PAC funds for their traditional low-intensity exploitation. Although other species may also benefit from this study since cinereous vulture could be a “flagship” for the large-scale conservation of Mediterranean oak woodlands and associated biodiversity, more fine local management guidelines should be performed on the basis of studies on more sensitive species.     

Estimating habitat suitability and potential population size for brown bears in the Eastern Alps

According to the Habitats Directive of the European Union, a favorable conservation status for the brown bear (Ursus arctos) should be targeted at the population level in large contiguous habitats such as the Alps, the largest mountain range in Europe. However, in most of the Alps brown bears are extinct and habitat suitability in these areas is often questionable. For this paper, radio-tracking data from four projects with 42 individual bears was compiled to assess habitat suitability. Discrete-choice models with random bear effects were fitted and compared to results obtained from compositional analysis and logistic regression. Sound definition of the available area in the discrete-choice model turned out to be essential. Brown bears showed a preference for forested and steep habitats and an avoidance of roads.Results from the three approaches were used to predict habitat suitability across the entire range of the Eastern Alps. Minimum potential population size was projected based on observed densities in Trentino and Central Austria, and ranged from 1228 to 1625 individuals, with 518–686 mature bears. This would satisfy a favorable conservation status. The developed methodology also has wide applicability to quantification of habitat suitability and potential population size in other cases where species are at risk.     

Foraging site fidelity shapes the spatial organisation of a population of female western barbastelle bats

Information about the spatial distribution of individual foraging habitats, which determines the space required by a population to be viable, is vitally important for the conservation of bats. Detailed knowledge of this kind is crucial for the design of nature reserves and management plans. Recent field studies that examined habitat use and home range distribution of bats largely ignored factors like traditional range use vs. intra- and interspecific competition, which may be responsible for the spatial organisation of a population home range. We investigated the home range sizes and distribution of a maternity colony of the western barbastelle bat via radio telemetry in four consecutive tracking sessions (2004-2007). Based on 19 data sets with a total of 2737 fixes obtained from 12 females, we examined (i) how colony members partition the population home range (home range overlap analysis), and (ii) if individuals tracked over several years exhibit site fidelity. Home range sizes ranged from 125 to 2551 ha (median: 403 ha), with a median number of 2 core areas (range: 1-5 core areas per individual per year). The core area sizes ranged from 5 to 285 ha (median: 67 ha). A home range and core area overlap analysis showed that site fidelity across years seems to be more important for home range distribution than competition among colony members. This allows researchers to combine information from several years to get a deeper insight into the population’s spatial requirements.     

Differential habitat selection by Pipistrellus pipistrellus and Pipistrellus pygmaeus identifies distinct conservation needs for cryptic species of echolocating bats

Cryptic species are similar in morphology, and make interesting subjects for relating morphological differentiation to ecological resource partitioning. Can species that are morphologically almost identical occupy different ecological niches, and hence potentially need distinct conservation planning? The discovery that the most widespread bat in Europe - the pipistrelle - comprised two cryptic species (Pipistrellus pipistrellus and Pipistrellus pygmaeus) that emit echolocation calls at different frequencies provides a remarkable model system for investigating links between morphology, echolocation call design and resource partitioning. We investigated resource partitioning between the two cryptic species of sympatric pipistrelle bats by radio tracking breeding females. Habitat selection was investigated by using compositional analysis. P. pygmaeus selected riparian habitats over all other habitat types in its core foraging areas, whereas P. pipistrellus, although preferring deciduous woodland overall, was more of a generalist, spreading its foraging time in a wider range of habitats. Although morphologically very similar, the cryptic species show quite different patterns of habitat use. Our findings suggest that large-scale differences in habitat preferences can occur between sympatric bat species that are virtually identical in flight morphology; hence morphological differences may be a weak indication of ecological differences between taxa. Conservation planning needs to take account of these differences to meet policy and legal obligations associated with these protected cryptic species.     

Scattered trees in rural landscapes: foraging habitat for insectivorous bats in south-eastern Australia

Scattered trees and small patches of vegetation among farmland are typical of rural landscapes throughout the world, often comprising a significant proportion of remaining habitats for native fauna. Insectivorous bats can use such isolated resources owing to the high mobility of most species, but little is known of the relationship between bats and tree density, or of the impact of incremental loss of trees in the landscape. Bats were surveyed at 30 sites in south-eastern Australia, in five habitat categories representing a range of tree densities from remnant woodland blocks (>35 trees/ha) to sparsely scattered trees (<1 tree/ha), and open paddocks devoid of trees. Sampling was undertaken by using harp traps and ultrasonic bat detectors. The abundance of potential arthropod prey was assessed by using light traps. Eleven species of insectivorous bats were recorded and bats were widespread in all habitat categories; all species were recorded around scattered trees. Overall activity, as revealed by detectors, did not differ significantly between the four treed categories, but in open paddocks there was a lower level of activity and a different community composition. However, a regression model revealed a significant quadratic relationship between activity of bats and tree density, with highest activity at 20-30 trees/ha. All species were recorded in open paddocks, but for eight of the ten species this represented <7% of their total activity recorded across all habitat categories. For six species, significant positive correlations were found between activity levels and the abundance of potential arthropod prey. In contrast to bat detectors, trapping results showed a significant difference in overall abundance of bats between wooded habitats, with decreased abundance as tree density declined. This study highlights the value of scattered trees as foraging habitat for bats, and emphasizes that, in rural land mosaics, such small and isolated habitat components have value for the conservation of biodiversity, and require greater recognition and protection.     

Effects of tropical forest fragmentation on aerial insectivorous bats in a land-bridge island system

Habitat fragmentation causes drastic changes in the biota and it is crucial to understand these modifications to mitigate its consequences. While studies on Neotropical bats have mainly targeted phyllostomid bats, impacts of fragmentation on the equally important aerial insectivores remain largely unexplored. We studied species richness, composition, count abundance and feeding activity of aerial insectivorous bats in a system of land-bridge islands in Panama with acoustic sampling. We predicted negative effects of fragmentation on forest species while bats foraging in open space should remain essentially unaffected. Rarefaction analyses indicated higher species richness for islands than mainland sites. For forest species, multivariate analyses suggested compositional differences between sites due to effects of isolation, area and vegetation structure. Contrary to our expectations, count abundance of forest species was similar across site categories. Feeding activity, however, was curtailed on far islands compared to near islands. As expected, bats hunting in open space did not reveal negative responses to fragmentation. Interestingly, they even displayed higher abundance counts on far and small islands. On the species level, two forest bats responded negatively to size reduction or site isolation, respectively, while a forest bat and a bat hunting in open space were more abundant on islands, irrespectively of island isolation or size. Our findings suggest that small forest remnants are of considerable conservation value as many aerial insectivores intensively use them. Hence high conservation priority should be given to retain or re-establish a high degree of forest integrity and low levels of isolation.     

Public concern about bats (Chiroptera) in Britain: An analysis of enquiries in 1982–1983

The Wildlife and Countryside Act 1981 protected all species of bat and required that, in certain circumstances, advice be sought about them. Analysis of these enquiries produced data about their geographic distribution, the types of problem that arose, the acceptability of the advice provided, the species of bat involved and the organisations receiving the enquiries. The great majority of enquiries related to dwelling-houses and, of these, most resulted from the discovery of bats roosting in the roof.The conservation organisations, mainly the Nature Conservancy Council, proved very successful in persuading householders to leave bats undisturbed; almost all colonies were permitted to remain temporarily and many were permitted to return in succeeding years. Repairs or remedial timber treatment to roofs used by bats were also important sources of enquiries and detailed advice was provided. Most enquiries came from southern England, with a sustained decline towards the north. Pipistrelles were the most common species, followed by long-eared bats. The other 13 species appeared to be involved in relatively few cases.     

Regional conservation priorities for a large predator: golden eagles (Aquila chrysaetos) in the Alpine range

We compared the density of golden eagles (Aquila chrysaetos) and their nest dispersion, productivity and diet in Alpine and pre-Alpine areas. The comparison was made at two spatial scales: (1) at the territory level within two contiguous populations, and (2) at the population level, using published data for 22 populations scattered across four countries (France, Switzerland, Germany and Italy). Availability of golden eagle's main prey species was higher in Alpine than in pre-Alpine regions. The potential foraging ranges of pairs of eagles in the pre-Alps had larger amounts of woodland and lower amounts of suitable foraging habitat than those of Alpine pairs. At both spatial scales, density and productivity were lower in the pre-Alps. Also, pre-Alpine populations were characterised by a lower percentage of main prey species in the diet, which was significantly correlated with productivity, and by a higher breadth of diet, which was negatively correlated with density. Overall, higher prey availability and habitat suitability in the Alps resulted in higher density and breeding success, in a manner compatible with theoretical metapopulation models. We suggest that priority be given to conservation of eagle populations in high quality Alpine habitat, and that conservation action be carried out by a mixture of site-protection measures and conservation of the wider environment.     

Sex-specific roost movements and population dynamics of the vulnerable long-fingered bat, Myotis capaccinii

The roosts of many IUCN-listed cave-roosting bat species are under threat from tourist development in SE Europe and other regions of the world. Much-needed conservation strategies require, among other information, an understanding of their roost movements and population dynamics, which can now be obtained relatively quickly using advanced models. We have studied the long-fingered bat, Myotis capaccinii, an obligate cave-dweller, in Dadia National Park, Greece. The species formed colonies of up to a few thousand individuals and was highly mobile, frequently switching summer roosts up to 39 km apart, even during late pregnancy. The bats migrated to distant hibernacula including a cave in Bulgaria 140 km NW of the Park. Adult recapture probabilities varied with season and sex: low female recapture rates in autumn, relative to spring and summer, indicated non-random temporary emigration following nursery colony dispersal. The opposite pattern was seen in males: increasing recapture rates in the autumn suggest that males gather in these roosts to mate with females in transit. Adult survival (0.86-0.94) was similar in females and males, similar in winter and summer, and comparable to recent estimates for other bats based on similar modelling techniques. Sex-based differences in juvenile recapture suggest female philopatry and male-biased dispersal. Our work shows that protection of M. capaccinii roosts must extend beyond the Park’s and indeed the country’s boundaries: its conservation requires large-scale, trans-national integrated conservation plans. Our results will apply to many other warm-temperate species with similar life history cycles.     

GIS-models work well, but are not enough: Habitat preferences of Lanius collurio at multiple levels and conservation implications

Species conservation largely depends on knowledge of habitat needs of target species. GIS-models are increasingly used to assess habitat preferences and distribution of target species, but their accuracy is constrained by availability of digital data layers. We developed a two-steps approach aiming at showing pros and cons of landscape (GIS)- and site-level habitat models, identifying key habitat factors for conservation of a threatened bird species, the red-backed shrike Lanius collurio. A spatially explicit GIS-model was generated using landscape variables, and a second model at site level was developed using fine-scale variables measured on the ground. The GIS-based model was then extrapolated to the entire region to obtain a map of distribution of suitable habitats. Positive associations between shrike occurrence and both hedgerow length and partial shrub cover were detected at both scales. Shrikes were also positively associated with grassland cover at landscape level and with partial cover of untilled herbaceous vegetation at the finer scale, and negatively affected by lucerne cover. The GIS-model led to an affordable map of predicted habitat suitability which should help conservationists to focus on different local priorities, but was unable to identify effects of untilled and lucerne cover. Site-level model gave fine details for habitat management, but its application elsewhere requires ground-measurements of factors. Combining the multiscale models could indicate more urgent actions at large scales (e.g. maintaining suitable habitats, or improving connectivity among isolated patches) and draw a detailed figure of the most suitable habitat for the species. Shrike occurrence was associated with a higher number of shrub and tree species: the indicator value of the species should ensure general benefits for biodiversity from dedicated management.     

Associations of bats with local structure and landscape features of forested stands in western Oregon and Washington

Understanding the processes that underlie bat distribution and activity patterns requires examination of habitat associations at multiple scales. We examined the association of both local structure and landscape context with bat activity in forested stands using ultrasonic detectors. Forty-eight stands in western Oregon and Washington were monitored for bat activity on at least six occasions for each of two field seasons. At the stand level, bat activity was negatively associated with tree density. The standard deviation of tree density and the density of newly created snags were positively associated with bat activity. In combination, these three variables explained 46% of the total variance in bat activity among stands. Landscape-level variables did not explain any significant variation among a subset of stands (n=22). Our study suggests that management of forest-dwelling bats should focus primarily on structural attributes at the stand level and the effects of these features on feeding and roosting opportunities.     

Habitat associations of bats in Northern Ireland: implications for conservation

Agricultural intensification in Northern Ireland has brought about large-scale changes to the landscape with a detrimental effect on biodiversity. Between 1996 and 1998, we surveyed a stratified random sample of 1 km squares for bats using a spot-sample technique and time expansion bat detector to establish linear and area habitat associations. Bats strongly selected water bodies with bankside vegetation, treelines, and deciduous and mixed woodland edge, avoiding open areas such as upland/unimproved grassland and improved grassland. Of three sympatric pipistrelle species (Pipistrellus pipistrellus, P. pygmaeus and P. nathusii), two were shown to forage in different habitats; P. pipistrellus being more of a ‘generalist’, foraging in a wider variety of habitats than P. pygmaeus, a ‘specialist’ which foraged in a smaller range of habitats. Nyctalus leisleri selected parkland/amenity grassland, deciduous woodland edge and rivers/canals and avoided improved grassland, hedgerows, whilst Myotis spp. selected deciduous woodland centre, lake/reservoir margins and rivers/canals. Reduction in area and quality of inland water, deciduous woodlands and field boundaries in Northern Ireland may impact on bat populations. Habitat management should focus on improving those habitats which are selected by bats, including the maintenance and enhancement of connecting linear habitats.     

When the species is also a habitat: Comparing the predictively modelled distributions of Lophelia pertusa and the reef habitat it forms

Internationally there is political momentum to establish networks of marine protected areas for the conservation of threatened species and habitats. Practical implementation of such networks requires an understanding of the distribution of these species and habitats. Predictive modelling provides a method by which continuous distribution maps can be produced from limited sample data. This method is particularly useful in the deep sea where a number of biological communities have been identified as vulnerable ‘habitats’, including Lophelia pertusa reefs. Recent modelling efforts have focused on predicting the distribution of this species. However the species is widely distributed where as reef habitat is not. This study uses Maxent predictive modelling to investigate whether the distribution of the species acts as a suitable proxy for the reef habitat. Models of both species and habitat distribution across Hatton Bank and George Bligh Bank are constructed using multibeam bathymetry, interpreted substrate and geomorphology layers, and derived layers of bathymetric position index (BPI), rugosity, slope and aspect. Species and reef presence records were obtained from video observations. For both models performance is fair to excellent assessed using AUC and additional threshold dependant metrics. 7.17% of the study area is predicted as highly suitable for the species presence while only 0.56% is suitable for reef presence, using the sensitivity–specificity sum maximisation approach to determine the appropriate threshold. Substrate is the most important variable in the both models followed by geomorphology in the RD model and fine scale BPI in the SD model. The difference in the distributions of reef and species suggest that mapping efforts should focus on the habitat rather than the species at fine (100 m) scales.