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.     

A study of species richness and diversity in seed banks and its use for the environmental mitigation of a proposed holiday village development in a coniferized woodland in south east England

A survey was carried out to determine the density and species composition of germinable seed in the surface soil layers of 30 plots within a coniferized lowland woodland in East Kent in order to establish the resources available for habitat creation in the event of some areas being modified during a proposed holiday village development. The selected plots included conifer plantations (up to 69 years old), broad-leaved plantations and the semi-natural broad-leaved edges that remain on parts of the perimeter of the site which were used as the control. A total of 13 682 seedlings emerged from the soil samples during the four-month germination trials. Fifty-two species were identified of which eight were ancient woodland indicator species for south-east England. The most abundant species represented in the seed banks were: Juncus effusus, Rubus fruticosus, Carex sylvatica, Betula pendula and Agrostis tenuis. Between-site comparisons of coniferous plots of different ages revealed a marked reduction in the seed species and seed density in plantations over 65 years old. Results of soil nutrient and texture analyses ruled out the likelihood of edaphic factors being responsible for between-site differences in seed bank composition. Seed species richness and diversity (Shannon-Wiener diversity index) were greatest in the semi-natural broad-leaved edges, but the diversity index used also showed that two replanted conifer sites had high values despite few species being present. The usefulness and limitations of diversity indices in the context of seed bank studies is discussed. From the results of the study, management proposals for the site have been put forward in order to maintain floristic diversity and mitigate the impact of the proposed development.     

Behavioural context of multi-scale species distribution models assessed by radio-tracking

Incorporating ecological processes and animal behaviour into Species Distribution Models (SDMs) is difficult. In species with a central resting or breeding place, there can be conflict between the environmental requirements of the ‘central place’ and foraging habitat. We apply a multi-scale SDM to examine habitat trade-offs between the central place, roost sites, and foraging habitat in Myotis nattereri. We validate these derived associations using habitat selection from behavioural observations of radio-tracked bats. A Generalised Linear Model (GLM) of roost occurrence using land cover variables with mixed spatial scales indicated roost occurrence was positively associated with woodland on a fine scale and pasture on a broad scale. Habitat selection of radio-tracked bats mirrored the SDM with bats selecting for woodland in the immediate vicinity of individual roosts but avoiding this habitat in foraging areas, whilst pasture was significantly positively selected for in foraging areas. Using habitat selection derived from radio-tracking enables a multi-scale SDM to be interpreted in a behavioural context. We suggest that the multi-scale SDM of M. nattereri describes a trade-off between the central place and foraging habitat. Multi-scale methods provide a greater understanding of the ecological processes which determine where species occur and allow integration of behavioural processes into SDMs. The findings have implications when assessing the resource use of a species at a single point in time. Doing so could lead to misinterpretation of habitat requirements as these can change within a short time period depending on specific behaviour, particularly if detectability changes depending on behaviour.     

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

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

Home range and habitat selection of northern bobwhite coveys in an agricultural landscape

Northern bobwhites thrive in fine-grained landscapes with a diversity of early succession woodland, grassland, and agriculture-associated habitat types. Bobwhite conservation has proved challenging in the increasingly coarse-grained Midwestern landscape as simplified agricultural cropping systems are implemented at larger spatial scales. Regardless, managing agricultural landscapes on private lands is the primary opportunity to restore bobwhite populations in the Midwestern United States. Although bobwhite habitat requirements are well understood, habitat selection in contemporary Midwestern landscapes is not well understood, especially on private lands where populations are declining. We used compositional analysis to investigate second- (study area) and third- (home range) order habitat selection by radiomarked bobwhite coveys on 4 private land study areas in southwestern Ohio. Mean covey home range size was 26.1 ± 2.2 ha (n = 48). Although home ranges were established in areas with more grassland cover, bobwhites most strongly selected early succession woody habitat (e.g., fencerows and ditches) at all scales, and selection for grassland diminished between the study area and home range scales. Grassland selection varied among sites and was strongest on sites with more row crop area. Woodlots were avoided at the study area scale, but were selected within home ranges. Grassland cover, like that provided by contemporary conservation programs, is an essential component of bobwhite habitat in the Midwest, but our results suggest more emphasis should be placed on early succession woody cover. Woody cover associated with fencerows, ditches, and woodlots adjacent to food sources and breeding habitat will likely improve non-breeding season survival, which is an influential vital rate in northern populations. © 2012 The Wildlife Society.     

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.     

Multi-scale habitat selection and foraging ecology of the eurasian hoopoe (<Emphasis Type="Italic">Upupa epops</Emphasis>) in pine plantations

Bird conservation can be challenging in landscapes with high habitat turnover such as planted forests, especially for species that require large home ranges and juxtaposition of different habitats to complete their life cycle. The eurasian hoopoe (Upupa epops) has declined severely in western Europe but is still abundant in south-western France. We studied habitat selection of hoopoes in pine plantation forests using a multi-scale survey, including point-counts at the landscape level and radio-tracking at the home-range scale. We quantified habitat use by systematically observing bird behaviour and characterized foraging sites according to micro-habitat variables and abundance of the main prey in the study area, the pine processionary moth (Thaumetopoea pityocampa). At the landscape scale, hoopoes selected habitat mosaics of high diversity, including deciduous woods and hedgerows as main nesting sites. At the home-range scale, hoopoes showed strong selection for short grassland vegetation along sand tracks as main foraging habitats. Vegetation was significantly shorter and sparser at foraging sites than random, and foraging intensity appeared to be significantly correlated with moth winter nest abundance. Hoopoe nesting success decreased during the three study years in line with processionary moth abundance. Thus, we suggest that hoopoes need complementation between foraging and breeding habitats to establish successfully in pine plantations. Hoopoe conservation requires the maintenance of adjacent breeding (deciduous woods) and foraging habitats (short swards adjacent to plantation edges), and consequently depends on the maintenance of habitat diversity at the landscape scale.     

Heard but not seen: Comparing bat assemblages and study methods in a mosaic landscape in the Western Ghats of India

We used capture (mist‐netting) and acoustic methods to compare the species richness, abundance, and composition of a bat assemblage in different habitats in the Western Ghats of India. In the tropics, catching bats has been more commonly used as a survey method than acoustic recordings. In our study, acoustic methods based on recording echolocation calls detected greater bat activity and more species than mist‐netting. However, some species were detected more frequently or exclusively by capture. Ideally, the two methods should be used together to compensate for the biases in each. Using combined capture and acoustic data, we found that protected forests, forest fragments, and shade coffee plantations hosted similar and diverse species assemblages, although some species were recorded more frequently in protected forests. Tea plantations contained very few species from the overall bat assemblage. In riparian habitats, a strip of forested habitat on the river bank improved the habitat for bats compared to rivers with tea planted up to each bank. Our results show that shade coffee plantations are better bat habitat than tea plantations in biodiversity hotspots. However, if tea is to be the dominant land use, forest fragments and riparian corridors can improve the landscape considerably for bats. We encourage coffee growers to retain traditional plantations with mature native trees, rather than reverting to sun grown coffee or coffee shaded by a few species of timber trees.     

Foraging by Nightjars Caprimulgus europaeus away from their nesting areas

Radio-tagged Nightjars Caprimulgus europaeus nesting in conifer plantation clearings and on lowland heath in Dorset travelled an average of 3.1 km away from their nesting areas at night to atypical habitats, presumably to feed. Comparison of habitat availability with use showed that Nightjars preferred deciduous or mixed woodland habitats, while they avoided conifer plantations and arable or improved grassland. Absence from the nesting area was recorded on 72 % of bird-nights and birds were least likely to be away during the middle third of the night. The use of atypical habitats may have implications for Nightjar conservation.     

Native vegetation structure and persistence of endangered Tehuantepec jackrabbits in a neotropical savanna in Oaxaca, México

Because of its crucial importance for the persistence of the endangered Tehuantepec jackrabbit (Lepus flavigularis), we identified structural characteristics of native vegetation (1) selected by Tehuantepec jackrabbits to establish home ranges, (2) used within home ranges, and (3) in relation to activity and inactivity periods. A neotropical savanna in the Isthmus of Tehuantepec, Oaxaca, México, offered diversity in native flora and heterogeneity in vegetation structure to the studied population of Tehuantepec jackrabbit. Adults and juveniles differed in habitat use patterns. No effect on habitat selection was detected for sex or season. Jackrabbits established home ranges on grassy habitat with discontinuous overstory of nanche (Byrsonima crassifolia) shrubbery and morro (Crescentia) trees. Adults underused dense vegetation where predators may ambush them; juveniles avoided dense vegetation and grassland without overstory. Within adults’ home ranges, habitat selection favoring grassy habitats with nanche and morro was relatively stronger during the activity period (nocturnal and crepuscular hours) and during one of 2 years which had more rainfall. Scattered trees and open shrubbery likely allowed foraging jackrabbits with both visibility and escape routes used to detect and outrun predators. Moreover, during the inactivity period (diurnal hours) adults and juveniles favored nanche shrubbery that provided resting jackrabbits with shelter to hide from predators. Clearly, structure and diversity of native vegetation in the savanna needs to be preserved for conservation of Tehuantepec jackrabbits because deteriorated habitat may pose higher predation rates for a jackrabbit population in risk of extirpation.     

The value of plantation forests for plant,invertebrate and bird diversity and the potential for cross-taxon surrogacy

As the area of plantation forest expands worldwide and natural, unmanaged forests decline there is much interest in the potential for planted forests to provide habitat for biodiversity. In regions where little semi-natural woodland remains, the biodiversity supported by forest plantations, typically non-native conifers, may be particularly important. Few studies provide detailed comparisons between the species diversity of native woodlands which are being depleted and non-native plantation forests, which are now expanding, based on data collected from multiple taxa in the same study sites. Here we compare the species diversity and community composition of plants, invertebrates and birds in Sitka spruce- (Picea sitchensis-) dominated and Norway spruce- (Picea abies-) dominated plantations, which have expanded significantly in recent decades in the study area in Ireland, with that of oak- and ash-dominated semi-natural woodlands in the same area. The results show that species richness in spruce plantations can be as high as semi-natural woodlands, but that the two forest types support different assemblages of species. In areas where non-native conifer plantations are the principle forest type, their role in the provision of habitat for biodiversity conservation should not be overlooked. Appropriate management should target the introduction of semi-natural woodland characteristics, and on the extension of existing semi-natural woodlands to maintain and enhance forest species diversity. Our data show that although some relatively easily surveyed groups, such as vascular plants and birds, were congruent with many of the other taxa when looking across all study sites, the similarities in response were not strong enough to warrant use of these taxa as surrogates of the others. In order to capture a wide range of biotic variation, assessments of forest biodiversity should either encompass several taxonomic groups, or rely on the use of indicators of diversity that are not species based.     

Habitat selection of three cryptic Plecotus bat species in the European Alps reveals contrasting implications for conservation

Assessing the ecological requirements of species coexisting within a community is an essential requisite for developing sound conservation action. A particularly interesting question is what mechanisms govern the stable coexistence of cryptic species within a community, i.e. species that are almost impossible to distinguish. Resource partitioning theory predicts that cryptic species, like other sympatric taxa, will occupy distinct ecological niches. This prediction is widely inferred from eco-morphological studies. A new cryptic long-eared bat species, Plecotus macrobullaris, has been recently discovered in the complex of two other species present in the European Alps, with even evidence for a few mixed colonies. This discovery poses challenges to bat ecologists concerned with planning conservation measures beyond roost protection. We therefore tested whether foraging habitat segregation occurred among the three cryptic Plecotus bat species in Switzerland by radiotracking 24 breeding female bats (8 of each species). We compared habitat features at locations visited by a bat versus random locations within individual home ranges, applying mixed effects logistic regression. Distinct, species-specific habitat preferences were revealed. P. auritus foraged mostly within traditional orchards in roost vicinity, with a marked preference for habitat heterogeneity. P. austriacus foraged up to 4.7 km from the roost, selecting mostly fruit tree plantations, hedges and tree lines. P. macrobullaris preferred patchy deciduous and mixed forests with high vertical heterogeneity in a grassland dominated-matrix. These species-specific habitat preferences should inform future conservation programmes. They highlight the possible need of distinct conservation measures for species that look very much alike.     

The importance of pre-thicket conifer plantations for nesting Hen Harriers Circus cyaneus in Ireland

Hen Harriers Circus cyaneus are threatened across much of their range and their conservation requires appropriate habitat management. The locations of 148 Hen Harrier nests found in the Republic of Ireland during national breeding surveys in 2000 and 2005 were used to assess nest-site selection. The distribution of these nests was compared to distributions of randomly located points to investigate selection at the scale of the nest-site and landscape. The main nesting habitats selected were pre-thicket stage of first and, particularly, second rotation plantations, mostly of exotic conifers. There was no evidence that the area of post-closure plantations negatively affected Hen Harrier nest distribution. There was a positive correlation across study areas between changes in numbers of Hen Harrier nests between 2000 and 2005 and changes in the area of pre-thicket second rotation plantations over the same period. The overall effect of plantation forests on breeding Hen Harriers in Ireland therefore appears to be positive. However, this study did not consider the effects of plantation habitats on breeding success. Improved grassland was strongly avoided as a nesting habitat. Furthermore, after controlling for the influence of nesting habitat on nest location, landscapes with a high percentage cover of improved grassland were also avoided. Further agricultural intensification of grassland in areas where Hen Harriers breed is likely to have a negative impact on this species. These results are required for the development of management strategies for the conservation of this species.     

Geographic Variation in Roost-Site Selection of Long-Legged Myotis in the Pacific Northwest

Abstract: Several species of bats in the Pacific Northwest of the United States, including long-legged myotis (Myotis volans), are dependent on snags in coniferous forests during summer for roosting and rearing young. Thus, data on roosting preferences of this species are needed to integrate their habitat requirements into shifting plans for management of forests in this region. Therefore, from 2001 to 2006, we radiotracked adult female long-legged myotis (n = 153) to day roosts (n = 395) across 6 watersheds in Washington, Oregon, and Idaho, USA, and compared characteristics associated with roosting sites to those of random snags (n = 260) sampled in the same watersheds using use-availability logistic regression and an information-theoretic approach. Model rankings varied among geographic locations, with quantity of stem surface for roosting the best model for explaining roost-site selection of long-legged myotis in both Washington and Oregon. Model rankings for populations of bats in Idaho found stand- and landscape-scale features to be important in roost-site selection, with a habitat fragmentation model and a foraging habitat quality model both demonstrating strong support as best model. Choice of day roosts by long-legged myotis was associated with snags that were taller, intact at the top of the stem, possessing a greater amount of exfoliating bark, in stands with a larger basal area of dead stems, and in landscapes that were unfragmented (i.e., supporting lesser amounts of edge). Results indicate that roost-site selection of bats in western coniferous forests, particularly long-legged myotis, is likely to be region-specific. We encourage land managers to consider importance of geographic variation in intraspecific habitat use in forest-dwelling bats when implementing silvicultural systems to promote biological diversity in actively managed forests of the Pacific Northwest region.     

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.     

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.     

Can pinewoods provide habitat for a deciduous forest specialist? A two-scale approach to the habitat selection of Bechstein's bat

Populations of Myotis bechsteinii in Sierra de Cazorla, Segura y Las Villas Natural Park (southwestern Spain) survive in a large, old-growth coniferous woodland, which is in contrast with known ecological preferences of the species. We tracked ten lactating females and studied patterns of habitat selection by Bechstein's bat (Myotis bechsteinii) at two spatial scales: macrohabitat and microhabitat, regarding forest essence (coniferous vs. deciduous). The tracked individuals always foraged within forested areas and did not use areas devoid of trees. At the macrohabitat level, no positive selection of deciduous stands was apparent, suggesting selection studies of coarse resolution may not be able to capture subtle selection patterns. At the microhabitat level Myotis bechsteinii selected deciduous patches within the coniferous matrix, therefore, our results corroborate the perception of this species as dependent of services provided by deciduous woodlands. Larger foraging home ranges and commuting distances as compared with other Mediterranean localities suggest that our studied population inhabits a marginal or suboptimal environment in terms of habitat quality. We argue that this population of Bechstein's bat has survived as a relict one probably as a consequence of fragmentation and transformation of deciduous forest ecosystems in the Mediterranean range.     

Foraging in space and time structure an African small mammal community

We used live-trapping and foraging to test for the effect of habitat selection and diet on structuring a community of six small mammals and one bird within the Soutpansberg, South Africa. We established grids that straddled adjacent habitats: woodland, rocky hillside, and grassland. Trapping and foraging were used to estimate abundance, habitat use, and species-specific foraging costs. The species with the highest abundance and foraging activity in a habitat, activity time, or food was considered the most efficient and presumed to have a competitive advantage. All species exhibited distinct patterns of spatial and temporal habitat preference which provided the main mechanism of coexistence, followed by diet selection. The study species were organized into three assemblages (α diversity): grassland, Rhabdomys pumilio, Dendromus melanotis, and Mus minutoides.; woodland, Aethomys ineptus and Micaelamys namaquensis; and rock-dwelling, M. namaquensis and Elephantulus myurus. Francolinus natalensis foraged in open rocky areas and under wooded islands within the grassland. Species organization across the habitats suggested that feeding opportunities are available within all habitats; however, distinct habitat preferences resulted from differing foraging aptitudes and efficiencies of the competing species. At Lajuma, species distribution and coexistence are promoted through distinct habitat preferences that were shaped by competition and species-specific foraging costs. The combination of trapping and foraging provided a mechanistic approach that integrates behavior into community ecology by ‘asking’ the animal to reveal its perspective of the environment. Using spatial and temporal foraging decisions—as behavioral indicators—enables us to guide our understanding for across-taxa species coexistence.     

Combining information from range use and habitat selection: sex-specific spatial responses to habitat fragmentation in tawny owls Strix aluco

How individuals respond to habitat heterogeneity is usually measured as variation in range size and by ranking the relative importance of habitat types (habitat selection). The combined effect of how individuals incorporate different habitat types in their home ranges and allocate their time budget between them is rarely derived. Additionally, when home range size varies between individuals, habitat selection analyses might be flawed if foraging decisions are based on variation in absolute rather than proportional availability. We investigated the suitability of standard analytical approaches by measuring the spatial responses of tawny owls to habitat fragmentation. These owls inhabited woodland of various sizes, representing a fragmentation gradient from open farmland with small, isolated woodland patches, to continuous woodland within their home ranges. In 17 territories within open farmland, the available area covered by woodland increased with the square root of the area of open land embraced in the home range. The owls did not display functional response in habitat selection, but females selected woodland more strongly than males. Females utilised woodland 10 times more intensively in farmland than in continuous woods, whereas males utilised farmland woods 3.2 times more intensively. Moreover, females in farmland exploited woodland 3.2 times as intensively as males, apparently because of higher travel costs in open areas. Since the extensive variation in intensity of use as a function of total availability was not indicated from the analysis of habitat selection, we suggest that information about intensity of use be more widely used as a supplementary measure of habitat use patterns than appears to be the practice at present.