Preservation of wide-ranging top predators by site-protection: Black and red kites in Doñana National Park

We used data from a long-term study on two medium-sized generalist predators, the black kite (Milvus migrans) and the red kite (Milvus milvus), to illustrate the complexity of managing wide-ranging top predators by site-protection. The study was conducted between 1989 and 2000 in the Reserva Biológica de Doñana, located at the core of Doñana National Park. Both species occurred at high density and showed high productivity. Black kites were slightly increasing, and red kites showed a non-significant trend. The main cause of breeding failure was nest predation, mainly by mammalian predators, most likely Iberian lynx (Lynx pardinus) or common genet (Genetta genetta). The main cause of mortality was illegal poisoning, which sharply increased in the late 1990s. Nest-site selection models showed that: (1) black kites selected areas near the seasonal marshes, rich in their main prey species and (2) red kites selected areas rich in marshland and far from the park border, possibly in response to illegal poisoning episodes. Productivity was positively related to the availability of marshland and of open terrestrial habitats for black and red kites, respectively, and negatively related for both species to the density of black kites within 200 m of the nest. Our results showed that: (1) intraguild predation and competition among predators make outcomes of interspecific interactions extremely complex, leading to unpredictable side-effects of priority actions targeting one species at a time and (2) though the park was fenced and patrolled by wardens, the wide ranging behaviour of the two species made park borders permeable to negative human pressures, such as poisoning, leading to extreme edge effects and absence of a true, safe core of the park. Park management needs to be better integrated with its regional socio-economic and cultural context. Education programmes against illegal poisoning should be urgently promoted.     

A test of the hierarchical model of habitat selection using eastern massasauga rattlesnakes (Sistrurus c. catenatus)

Understanding the process animals follow to select habitat, rather than just documenting the habitat they use, will improve our ability to predict how the animals use habitat in other locations and how they will respond to changes in habitat. Animals are usually assumed to select habitats hierarchically, preferentially using specific macrohabitats at a landscape scale and specific microhabitats within the preferred macrohabitats. We used four years of telemetry data from 34 individuals to test this hierarchical model of habitat selection with eastern massasauga rattlesnakes (Sistrurus c. catenatus) in Ontario. Snakes were selective at the microhabitat scale, preferentially using locations with closer retreat sites and shrubs than random. Gravid females were most selective, using sites with more rock cover and less canopy closure than sites used by males and nongravid females. Snakes preferred forested habitats for hibernation and steadily increased their use of open, wetland, and edge habitats to a peak in mid-summer. Landscape-scale habitat preferences were generally mild and could be explained by the relative availability of suitable microhabitat within habitats, suggesting habitat selection was primarily driven by microhabitat preferences. The lack of selectivity at the landscape scale may be a consequence of fine-grained differences between habitats that allow massasauga rattlesnakes to find suitable microhabitats in all available macrohabitats. For species that select habitat primarily at the microhabitat scale (e.g., the rattlesnakes we studied), landscape-scale modeling of habitat use will only be effective to the extent habitats reflect the availability of suitable microhabitat within.     

Satellite tracking of breeding black storks Ciconia nigra: new incomes for spatial conservation issues

Satellite tracking of black storks was used to estimate home range sizes and to study habitat selection during the breeding season. Breeding and non-breeding adults foraged over very large areas (ca. 54 000 ha for 12 territories), preferentially in woodlands with high number of river sources, mirroring the species needs for high quality water resource. Rearing and post-fledging ranges of breeding partners largely overlapped. Home ranges of non-breeding adults largely overlapped ranges of breeding birds, so that assessing home range size of breeding pairs from observed densities is not reliable. Protection and management of breeding and feeding habitats appear to be the most important conservation measures to be considered. This study allowed to evaluate how large these protected areas should be, and which habitat types they should encompass. Conservation measures for the species in western Europe should include protection of very large forest areas and also focus on managing river networks to ensure a high water quality as far as 20 km away from nests.     

Reconciling the dichotomy between single species and ecosystem conservation: black kites (Milvus migrans) and eutrophication in pre-Alpine lakes

In recent times there has been a growing dichotomy between preservation of single species and broader, ecosystem-based approaches to conservation. Freshwater habitats are among the most highly human-impacted ecosystems. We used a long-term data set on black kites (Milvus migrans), a threatened raptor dependent on aquatic habitats, to explore ways to reconcile single species and biodiversity-driven approaches to conservation. In the pre-Alpine lakes of northern Italy, black kite populations showed medium-low density, extremely low breeding success and widespread declines. Spatio-temporal variations showed density and breeding success to be positively related to ecosystem productivity (as estimated by phosphorus concentrations in lakes), availability of aquatic habitats and grassland, and to be negatively related to extent of farmland and fish harvest by professional fishermen. Fish species richness, used as a surrogate of biodiversity, was highest in one oligotrophic lake, but on average increased with increasing ecosystem productivity (i.e. lake eutrophication). Given expected future declines in ecosystem productivity, kite conservation will be helped by enhancing populations of alternative prey in terrestrial habitats (e.g. through incentives for grassland), and higher regulation of fish harvest. On the other hand, ecosystem management may profit from the use of black kites or other aerial piscivores as indicators of biodiversity and of diffuse ecosystem stress, such as sustainability of fishing practices.     

Effects of an exotic prey species on a native specialist: Example of the snail kite

Despite acknowledging that exotic species can exhibit tremendous influence over native populations, few case studies have clearly demonstrated the effects of exotic prey species on native predators. We examined the effects of the recently introduced island apple snail (Pomacea insularum) on the foraging behavior and energetics of the endangered snail kite (Rostrhamus sociabilis plumbeus) in Florida. We conducted time-activity budgets: (i) on kites foraging for native Florida apple snails (Pomacea paludosa) in major wetland units within the kites’ range that had not been invaded by the exotic island apple snail and (ii) on kites foraging for exotic apple snails in Lake Tohopekaliga, the only major wetland utilized by the snail kite that had suffered a serious invasion of P. insularum. When foraging for P. insularum, snail kites dropped a greater proportion of snails, and they experienced increased handling times and decreased consumption rates; however, kites foraging for P. insularum also spent a smaller proportion of the day in flight. Estimates of net daily energy balances between kites feeding on P. insularum versus P. paludosa were comparable for adults, but juveniles experienced energetic deficiencies when feeding on the exotic snail. Due to this discrepancy, we hypothesize that wetlands invaded by P. insularum, such as Lake Tohopekaliga, may function as ecological traps for the snail kite in Florida by attracting breeding adults but simultaneously depressing juvenile survival. This study highlights the conservation implications and importance of elucidating the effects that exotic species have on native specialists, especially those that are endangered, because subtle influences on behavior may have significant population consequences.     

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.     

Does land irrigation actually reduce foraging habitat for breeding lesser kestrels? The role of crop types

The lesser kestrel is a Globally Threatened Species which large decline has been related to recent agricultural changes in European pseudo-steppes. Irrigation is considered as one of the major threats for this and other steppe birds, but the actual effects of irrigation on foraging habitat selection have been scarcely examined. We studied the selection of traditional dry cereal farming and irrigated habitats by foraging lesser kestrels during the breeding cycle, paying especial attention to possible differences among crop types. Field margins were the scarcest but the most positively selected habitat, and different stages of cereals cultivated following traditional practices were selected depending on the breeding and agriculture cycles. Effects of irrigation were dual. While irrigated maize and other crop types were avoided, alfalfa was used in proportion to its availability and later highly selected after harvesting. Moreover, field margins in irrigated land were selected in a similar way than in traditional dry farmland. Therefore, although maintaining low-intensity farming is still the main recommendation for this species, new management options arise when social pressure makes irrigation unavoidable. Further agri-environmental schemes in these circumstances should thus promote cultivation of alfalfa with a low input of biocides while avoiding maize, together with increasing field margins, to make compatible irrigation with lesser kestrel conservation.     

Habitat selection and conservation of Sharpe's longclaw (Macronyx sharpei), a threatened Kenyan grassland endemic

Montane grassland is a severely threatened habitat in Kenya. Despite a high level of faunal endemism, it has received very little conservation attention. We investigated habitat selection in a threatened grassland endemic bird species, Sharpe's longclaw (Macronyx sharpei) (family Motacillidae), in order to understand its likely response to land-use changes. Between November 1995 and May 1996, we studied 41 territories of this species on the Kinangop Plateau in central Kenya. With an overall density of 0.4 birds ha⁻¹, longclaws lived in permanent groups of two to seven individuals. They were sedentary and territorial, with a mean home range size of 0.5 ha. The species avoided non-grassland areas entirely, and within grassland showed a strong preference for short grass with tussocks. Territory sizes and foraging ranges were smaller, and rates of pecking for food higher, in this grassland type than in open short grass or long grass. Land use changes, in particular conversion to cultivation or woodlots and ploughing up of grassland to remove tussocks, pose a serious and immediate threat to this species. Dairy farming is potentially compatible with grassland conservation, and conservation-friendly livestock rearing needs to be promoted through economic incentives, awareness-raising and technical advice.     

Landscape composition affects habitat use and foraging flight distances in breeding European starlings

The recent decline of the European starling in northern Europe has been associated with changes in agricultural land-use. We investigated how agricultural land-use affected starling habitat use and home-range size in a heterogeneous landscape in southern Sweden. Breeding starlings primarily foraged in pastures, a habitat with a high availability of potential food for nestlings. When the availability of pasture close to the nest was low, starlings increased their use of other permanently grass-covered habitats such as field margins and ditches. They also spent a larger proportion of their daytime flying because they visited more distant foraging areas. The further from the nest parents foraged, the higher was the probability that they foraged on a pasture. Feeding frequency of nestlings was negatively related to the distance from the nest parents foraged, but no consequences on nestling survival or growth were found. A lower breeding density at low local availability of pasture may compensate for lack of good foraging habitat. It is suggested that recent changes in the agricultural landscape may have increased the mismatch between the availability of good foraging areas and nest-sites. This might have contributed to the recent population decline of the starling in northern Europe by affecting both foraging habitat quality and flight costs paid by parents during breeding.     

Impact of grassland farming intensification on the breeding ecology of an indicator insectivorous passerine, the Whinchat Saxicola rubetra: Lessons for overall Alpine meadowland management

The decline of insectivorous farmland passerines has been attributed mostly to global decrease in arthropod availability, as a result of intensification of agricultural practices. The diminution of the Alpine Whinchat, once a widespread insectivorous passerine, has been ascribed to nest losses due to earlier and more frequent mowings. However, potential conjugated effects of deteriorated arthropod food availability had yet to be investigated. We compared food supply and nestling diet in intensively vs. traditionally managed grassland. Abundance and diversity of arthropods were much lower in intensive areas, where small-sized invertebrates, which do not enter nestling diet, were also predominant. Parents breeding in intensive habitats fed less biomass to nestlings than adults from traditional habitats. Nestling diet was less diverse and dominated by less profitable prey items in intensive than in traditional habitats. Feeding rate did not differ between the two habitats, but foraging distances from nest tended to be greater in intensive farmland. There were no significant differences in clutch sizes and hatching success with respect to management intensity, but fledging success was higher in traditional habitats. The recent intensification of farming practices has led to a decrease in the availability of grassland invertebrates, and of important Whinchat nestling food in particular, affecting parents’ foraging efficiency and reproductive success. Conservation actions must not only reduce nest losses by postponing mowing, but should also promote grassland farming that is less detrimental to invertebrates. Dominant at the study site, organic grassland farming does seemingly not provide sufficient conditions for Alpine Whinchats.     

Bighorn sheep habitat use and selection near an urban environment

To further understand the impact of urban development on wildlife populations, we examined habitat use and selection by female bighorn sheep Ovis canadensis in two endangered subpopulations near a metropolitan area in southern California. One subpopulation, which had previously been found to have low reproductive success, selected urban environments while the other did not use urban areas. In the subpopulation that used urban areas, females had smaller core activity areas and selected lower elevations and gentler slopes. These females used urban sources of water but a clear relationship between levels of urban use and periods of increased water need was not evident. Diet quality was higher among females that selected urban areas, and this increase was correlated with the level of urban use. Thus, optimal foraging behavior may have contributed to the selection of urban areas. Urban use was lowest during peak months of parturition, suggesting that reproductive strategies may also have influenced temporal patterns of urban use. Although urban areas provided bighorn sheep with forage and water resources, the use of these areas may have substantial costs. For example, females using urban areas had internal parasites that were not found elsewhere in the Peninsular Ranges.     

Multi-trophic consequences of kelp harvest

Coastal kelp forest ecosystems provide important habitats for a diverse assemblage of invertebrates, fish and marine top-predators such as seabirds and sea mammals. Although kelp is harvested industrially on a worldwide scale little is known about the multi-trophic consequences of this habitat removal. We investigated how kelp fisheries, which remove feeding and nursery grounds of coastal fish, influence local food webs and the availability of food to a marine top predator, the great cormorant (Phalacrocorax carbo). We conducted experimental harvesting of the canopy-forming kelp (Laminaria hyperborea) during a 3 year period (2001-2003) in an area at the coast of Central Norway while synoptically monitoring fish occurrence and cormorant foraging parameters. Our results demonstrate that cormorants preferentially foraged within kelp-forested areas and performed significantly more dives when feeding in harvested versus un-harvested areas suggesting lower foraging yield in the former case. In kelp areas that were newly harvested the number of small (<15 cm) gadid fish was 92% lower than in un-harvested areas. This effect was persistent for at least 1 year following harvest. To our knowledge, this is the first time that the ecological consequences of kelp harvesting have been tested at a multi-trophic level. The results presented strongly suggest that kelp harvesting affects fish abundance and diminishes coastal seabird foraging efficiency. Kelp fisheries are currently managed in order to maximize the net harvest of kelp biomass, and the underlying effects on the ecosystems are partly ignored. This study calls for re-assessment of such management practices.     

Habitat selection by the Mediterranean horseshoe bat, Rhinolophus euryale (Chiroptera: Rhinolophidae) in a rural area of southern Italy and implications for conservation

We studied habitat selection by Rhinolophus euryale in a rural area of southern Italy in 1998-2000 by radio-tracking. Two comparisons were carried out, one between habitat occurrence within individual home ranges and within the study area, the other between time spent in each foraging habitat and habitat occurrence within the home range. The first analysis showed that olive groves and conifer plantations were, respectively, the most and the least important habitats. The second analysis highlighted the importance of woodland for R. euryale, while urban sites, open areas and conifer plantations were avoided. We recommend that clearing of continuous, large areas of woodland for tree harvesting should be avoided. Conifers should not be used for reforestation. Urbanisation should be limited in the areas of greatest importance for the species, and linear landscape elements such as tree lines and hedgerows should be maintained.     

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.     

Hunting habitat selection by hen harriers on moorland: Implications for conservation management

We examine habitat use by hunting hen harriers Circus cyaneus at three study sites in Scotland to evaluate whether foraging patterns differ between sexes, sites, and stages of the breeding period. We modelled time spent hunting in focal plots as a function of habitat and nest proximity. Male hunting intensity (time spent hunting per hour of observation and km²) varied between sites and breeding periods, being lower during the nestling than the incubation period. Habitat use patterns were mostly consistent among study sites, which is important for developing species management recommendations applicable over the species’ range. Males avoided improved grassland, and selected areas of mixed heather and rough grass (with an optimum at ca. 50% heather cover). The effect of nest proximity was small. In contrast, females hunted mainly within 300-500 m of the nest, with a small additive effect of vegetation cover, areas of fragmented heather being preferred. Habitat management to benefit foraging harriers will involve creating (or maintaining) mosaics of heather/grassland around nest areas. Additionally, it might be possible to manipulate habitat to reduce conflict in areas where harrier predation on red grouse is important by segregating areas holding highest grouse densities (with high heather cover) from those favoured for harrier foraging (heather-grass mosaics). However, it would be necessary to test whether these manipulations might also influence harrier nest distribution, an effect which could negate any benefits from this strategy.     

Higher breeding densities of the threatened little bustard Tetrax tetrax occur in larger grassland fields: Implications for conservation

Previous studies have found that densities of little bustard Tetrax tetrax breeding males tend to be higher in areas with smaller agricultural fields, presumably due to increased habitat diversity. However, exceptionally high densities have been found in large grassland fields in Portugal, which suggests that the influence of field size varies geographically, and that the role of this factor is not yet fully understood, despite its importance as a key management issue.We studied how field size, together with vegetation structure, influences the presence and density of breeding little bustards in a region of southern Portugal. Fifty-four grassland fields were sampled in 2007 and another 29 in 2008, with sizes ranging from 23 to 172 ha. A total of 183 breeding males were found in 47 of these fields, reaching densities of up to 37 males/100 ha. A higher probability of occurrence of breeding males was found in larger fields with a vegetation height below 40 cm and field size alone explained 46% of the variability in male density. These results suggest that larger continuous areas of suitable habitat attract many males, most likely as a consequence of their lek mating system. We conclude that conservation efforts, in a landscape context of large farm sizes, should: (1) be channelled to farms with large fields; (2) ensure adequate livestock grazing to create suitable habitat and (3) promote management at a landscape level to ensure the most continuous grassland habitat patches possible.     

Grassland nature reserves for breeding wading birds in England and the implications for the ESA agri-environment scheme

The two principal mechanisms available to conserve breeding wading birds on lowland wet grassland in England are nature reserves and agri-environment schemes, particularly the Environmentally Sensitive Area (ESA) scheme. ESAs offer payments to landowners in return for maintaining (‘low tiers’) or enhancing (‘high tiers’) the landscape, biodiversity and historic value of the land. Lowland wet grassland nature reserves managed by the Royal Society for the Protection of Birds (RSPB) have been successful at conserving breeding lapwings and redshank but habitat management usually only results in short-term increases in numbers of breeding snipe. Within ESAs, populations of breeding wading birds fare better on land managed under high tiers than on land managed under low tiers. Land managed under high tiers also supports more pairs of breeding wading birds per £ of ESA grant received than land managed under low tiers. Means of increasing the cost-effectiveness of the ESA scheme in conserving breeding wading birds are discussed in the light of these results.     

Habitat management and patterns of predation of Northern Lapwings on wet grasslands: The influence of linear habitat structures at different spatial scales

In managed landscapes, habitat structure is frequently manipulated through the creation of features such as tracks, hedges, and waterways. If predator and prey activity are concentrated around these features, levels of predation may be elevated in these landscapes. This issue is of particular importance when habitat structures are used to attract species of conservation concern. For example, the installation of linear waterways in wet grasslands is a common form of habitat management to benefit breeding waders and wader nests and foraging chicks tend to be aggregated around wet features. If predator activity is also focused around these features, and if their linearity increases the probability of prey being located, then the conservation benefits of this management technique may be eliminated. We explore predator movement in relation to the structure and complexity of linear wet features within a lowland wet grassland landscape. We examine patterns of nest and chick predation in lapwing (Vanellus vanellus) at the whole-site, between-field and within-field scales. Mammalian predators were responsible for the majority of nest predation. However, we found no evidence that mammalian predators used linear wet features disproportionately within the landscape, or that wet feature distribution influenced the probability of nest or chick predation. At the whole-site scale, nest predation rates were significantly higher in areas with greater predator presence and lowest where the number of breeding neighbours was high. Thus, predation levels were influenced by large-scale patterns of predator presence and lapwing density but not by the use of linear wet features as a habitat management tool. Managing predator impacts is therefore likely to require empirical assessments of local predator distribution and abundance in order to target measures effectively.     

Determining the spatial scale for conservation purposes - an example with grizzly bears

This study suggests procedures for determining the spatial scale for conservation guidelines for animals, giving an illustration with an analysis of grizzly bear habitat selection. Bear densities were sampled by identifying hairs at bait stations in British Columbia. Habitat variables were measured using remote sensing. Spatial scale was changed by varying the window size over which the variables were averaged. First, the spatial pattern of bears was studied, measuring the patchiness in bear densities at a variety of spatial scales, by calculating the correlation in bear densities between adjacent windows. This was repeated for the habitat variables. Finally, the overall interaction between bears and habitats was analysed, measuring the strength of habitat selection at different spatial scales. There are three domains of scale: at 2-4 km, bears and habitats are patchy, at 5-10 km, bears select for habitats, and at 40+ km, habitats are patchy and bears select for habitats. At scales of 40+ km, bears selected for: (i) higher slopes, or (ii) higher slopes, and some combination of more avalanche chutes, fewer roads and trees, higher elevations, and less logged land. Within 15 km areas, bears selected for 6 km areas that are either at higher elevations, or at higher elevations and had fewer trees. The relationship of conservation guidelines at different spatial scales should be determined by measuring and comparing hierarchical to non-hierarchical selection. The scales that bears select for habitats roughly correspond to the scales used in present grizzly bear conservation plans in British Columbia.     

Identifying indirect habitat loss and avoidance of human infrastructure by northern mountain woodland caribou

Understanding the impact of indirect habitat loss resulting from avoidance of human infrastructure is an important conservation priority. We evaluated resource selection for 10 global positioning system collared northern mountain woodland caribou (Rangifer tarandus caribou) in British Columbia, Canada, with seasonal resource selection functions (RSF) developed at the second-order (landscape) and third-order (within home range) scales. To estimate how much habitat was lost due to avoidance, we estimated the zone of influence (ZOI) around multiple developments and modeled realized and potential habitat. Potential habitat was approximated by removing the ZOI from RSF models. By calculating the spatial difference between potential and realized habitat we estimated the amount of indirect habitat loss. Caribou displayed hierarchical avoidance of development, with the greatest avoidance occurring at the second-order. During both seasons caribou avoided high-use roads by 2 km and low-use roads by 1 km. In winter, caribou avoided town by 9 km compared to 3 km in summer. However, in summer caribou avoided mines by 2 km and cabins and camps by 1.5 km, while in winter when human activity was low, avoidance of these features was minor. As a result of avoidance of the cumulative ZOI, approximately 8% and 2% of high quality habitat was lost in the study area in winter and summer, respectively. Our study provides an approach to identify the extent and quality of habitat influenced by indirect avoidance. Conservation efforts should prioritize protecting areas of high quality habitat degraded by avoidance in the vicinity of human development.