The effects of local and landscape-scale habitat characteristics and prey availability on corridor use by carnivores: A comparison of two contrasting farmlands

Corridors are thought to reduce the negative biological effects of habitat loss and fragmentation by providing connectivity and suitable habitat for many species, including carnivores. Although corridor structure maintenance is considered to be an essential tool for carnivore conservation in a human-dominated landscape, surprisingly little is known about the effects of different factors at various spatial scales. The main aim of this study was to determine how local and landscape-scale habitat characteristics and prey availability influence the corridor use by carnivores in a Central European agricultural landscape. Moreover we investigated carnivore corridor occurrence in two contrasting landscapes that differ in level of habitat loss and fragmentation to evaluate relative effect of species-specific response to regional context. Results show that the availability of principal prey (small mammals) was the most crucial factor affecting carnivore corridor use. Other important factors influencing corridor use were corridor width (positive), proportion of shrubs (positive) and presence of local, low-traffic, roads (positive). Single species models revealed interspecific and area-specific differences in carnivore preference at both spatial scales. Our findings confirm the general importance of multi-level approach to evaluating species-specific habitat requirements as a crucial tool for determining suitable methods for carnivore efficient conservation in human-dominated agricultural landscape.     

Assessing species traits and landscape relationships of the mammalian carnivore community in a neotropical biological corridor

Mammalian carnivores play an important role in regulating food webs and ecosystems. While many carnivore populations are facing various threats such as habitat loss and fragmentation, poaching, and illegal trade, others have adapted to human-dominated landscapes. Information about Neotropical carnivore communities in particular is limited, especially in disturbed landscapes. We conducted a camera trap survey at 38 sites across the San Juan–La Selva Biological Corridor in Costa Rica to assess occupancy and detection probabilities of the carnivore community. We developed hypotheses within a likelihood-based framework in order to determine the landscape features and species traits (diet and size) that influenced their occupancy. We detected nine of the 13 native carnivores predicted to occur in the corridor. When modeled separately, each species responded to land cover changes differently, suggesting no strong community-wide predictors of occupancy. We then modeled three separate guilds within the carnivore community: omnivorous mesopredators, obligate carnivorous mesopredators, and apex predators. These community guild models revealed a negative relationship between omnivorous mesopredators and increasing forest and tree plantation cover, suggesting omnivores utilize forest fragments and edge habitats in agricultural landscapes. Obligate carnivorous mesopredator models did not reveal any strong habitat relationships, but landscape effects tended to contradict our a priori predictions. Apex predators were positively associated with increasing forest and tree plantation cover, protected areas, and increasing distances to villages. Alarmingly, apex predators and obligate carnivorous mesopredators were generally rare within the biological corridor. A lack of top-down control alone might result in heightened occupancy for all mesopredators, but because the community is dominated by omnivorous species, bottom-up release from human-induced land cover changes and resource provision may better explain their high occupancy.     

Use of agroecosystem matrix habitats by mammalian carnivores (Carnivora): a global‐scale analysis

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Scale‐dependent strategies for coexistence of mesocarnivores in human‐dominated landscapes

Identifying factors influencing the distribution of and interactions within carnivore communities is important for understanding how they are affected by human activities. Species differ in their ability to adapt to humans depending on their degree of specialization in habitat use and feeding habits. This results in asymmetric changes in the ecology of co‐occurring species that can influence their interactions. We investigated whether human infrastructures and free‐ranging domestic dogs (a species typically associated with humans) influenced the co‐occurrence and habitat use of mesocarnivores in a landscape of high human population density in Maharashtra, India. We used 40 camera trap locations during 233 trapping nights and used Bayesian co‐occurrence occupancy models to investigate the habitat use and coexistence of species at different spatial scales. Additionally, we investigated their temporal overlap in space use. Indian foxes altered their habitat use both spatially and temporally in order to avoid free‐ranging domestic dogs and other larger competitors. The use of human infrastructure by jackals and jungle cats was limited by the presence of dogs. Our results illustrate how habitat use of smaller carnivore species changes both spatially and temporally in order to avoid larger competitors. We also show that the presence of species associated with humans mediates the influence of human infrastructures on the habitat use of mesocarnivores. We highlight the importance of acknowledging the potential impact of urbanization not only on single species, but also on the interactions within the community.     

With Allee effects,life for the social carnivore is complicated

Anthropogenic modification of the landscape, resultant habitat loss, and decades of persecution have resulted in severe decline and fragmentation of large carnivore populations worldwide. Infectious disease is also identified as a primary threat to many carnivores. In wildlife species, population demography and group persistence are strongly influenced by group or population size. This is referred to as the Allee effect, in which a population or group is at an increased risk of extinction when the number or density of individuals falls below some threshold due to ecological and/or genetic factors. However, in social mammalian species, the relationship between the number of individuals and the risk of extinction is complicated because aggregation may enhance pathogen exposure and transmission. Although theoretical studies of the interaction between infectious disease transmission and Allee effects reveal important implications for carnivore management and population extinction risk, information about the interaction has yet to be synthesized. In this paper, we assess life history strategies of medium to large carnivore species (≥2.4 kg) and their influence on population dynamics, with a special focus on infectious disease. While declining population trends are observed in 73 % of all carnivores (both social and solitary species), infectious disease is identified as a significant cause of population decline in 45 % of social carnivores and 3 % of solitary carnivores. Furthermore, where carnivores suffer a combination of rapid population decline and infectious disease, Allee effects may be more likely to impact social as compared to solitary carnivore populations. These potentially additive interactions may strongly influence disease transmission dynamics and population persistence potential. Understanding the mechanisms that can result in Allee effects in endangered carnivore populations and the manner in which infectious disease interfaces at this nexus may define the outcome of developed conservation strategies.     

Conservation of biodiversity in Scandinavian boreal forests: large carnivores as flagships, umbrellas, indicators, or keystones?

Large carnivores are often used as focal species (indicators, umbrellas, flagships or keystones) in conservation strategies either aimed at conserving carnivores, the rest of the biodiversity that occupies their habitats, or both. We evaluate their suitability for these roles in the context of boreal forest biodiversity conservation in the muti-use landscapes of Scandinavia. The enormous conflicts, especially with livestock, that carnivores cause in these areas makes them very controversial flagships to the extent that it may affect rural people's attitudes to conservation in general. Because of the broad habitat tolerance of large carnivores and their prey, and the difficulties in surveying carnivore numbers, they are very insensitive and impractical indicators of forest biodiversity. This ability of large carnivores to thrive in industrial forests means that the many species that are sensitive to modern forestry will not fall under the umbrella of areas managed for large carnivores. If large carnivores have a keystone function with respect to affecting the density of their ungulate prey it is likely to lead to even further conflicts with hunters who gain economic benefit from harvesting wild ungulates. In other words, none of the classic `ecological' arguments are likely to help justify large carnivore conservation, and large carnivore conservation is unlikely to help conserve the rest of the boreal forest's biodiversity. Based on these arguments we recommend that (1) justification for large carnivore conservation focus on the real philosophical and value orientated reasons rather than ecological justifications, (2) that this conservation should be brought about in practice by dedicated management programs that specifically address the conflicts caused by large carnivores, and (3) that boreal forest biodiversity is best conserved by specific actions designed to establish reserves or change forestry practices.     

Using Landscape and Bioclimatic Features to Predict the Distribution of Lions,Leopards and Spotted Hyaenas in Tanzania's Ruaha Landscape

Tanzania''s Ruaha landscape is an international priority area for large carnivores, supporting over 10% of the world''s lions and important populations of leopards and spotted hyaenas. However, lack of ecological data on large carnivore distribution and habitat use hinders the development of effective carnivore conservation strategies in this critical landscape. Therefore, the study aimed to (i) identify the most significant ecogeographical variables influencing the potential distribution of lions, leopards and spotted hyaenas across the Ruaha landscape; (ii) identify zones with highest suitability for harbouring those species; and (iii) use species distribution modelling algorithms (SDMs) to define important areas for conservation of large carnivores. Habitat suitability was calculated based on environmental features from georeferenced presence-only carnivore location data. Potential distribution of large carnivores appeared to be strongly influenced by water availability; highly suitable areas were situated close to rivers and experienced above average annual precipitation. Net primary productivity and tree cover also exerted some influence on habitat suitability. All three species showed relatively narrow niche breadth and low tolerance to changes in habitat characteristics. From 21,050 km² assessed, 8.1% (1,702 km²) emerged as highly suitable for all three large carnivores collectively. Of that area, 95.4% (1,624 km²) was located within 30 km of the Park-village border, raising concerns about human-carnivore conflict. This was of particular concern for spotted hyaenas, as they were located significantly closer to the Park boundary than lions and leopards. This study provides the first map of potential carnivore distribution across the globally important Ruaha landscape, and demonstrates that SDMs can be effective for understanding large carnivore habitat requirements in poorly sampled areas. This approach could have relevance for many other important wildlife areas that only have limited, haphazard presence-only data, but which urgently require strategic conservation planning.     

Seasonal variation in prey preference,diet partitioning and niche breadth in a rich large carnivore guild

Large carnivore community structure is affected by direct and indirect interactions between intra-guild members. Co-existence between different species within a carnivore guild may occur through diet, habitat or temporal partitioning. Since carnivore species are highly dependent on availability and accessibility of prey, diet partitioning is potentially one of the most important mechanisms in allowing carnivores to co-exist. Intra-guild interactions may vary over time as carnivore prey preference and diet overlap can change due to seasonal changes in resource availability. We conducted scat analysis to compare the seasonal changes in prey preference, diet partitioning and niche breadth of four large carnivore species, namely leopard Panthera pardus, spotted hyena Crocuta crocuta, brown hyena Parahyaena brunnea and wild dog Lycaon pictus in central Tuli, Botswana. Large carnivores in central Tuli display a high dietary overlap, with spotted hyena and brown hyena displaying almost complete dietary overlap and the other carnivore species displaying slightly lower but still significant dietary overlap. Dietary niche breadth for both hyena species was high possibly due to their flexible foraging strategies, including scavenging, while leopard and wild dog showed a relatively low niche breadth, suggesting a more specialised diet. High dietary overlap in central Tuli is possibly explained by the high abundance of prey species in the area thereby reducing competition pressure between carnivore species. Our research highlights the need to assess the influence of diet partitioning in structuring large carnivore communities across multiple study sites, by demonstrating that in prey rich environments, the need for diet partitioning by carnivores to avoid competition may be limited.     

Body Size Overlap,Habitat Partitioning and Living Space Requirements of Terrestrial Vertebrate Predators: Implications for the Paleoecology of Large Theropod Dinosaurs

Ecological studies of extant tetrapod predators indicate that morphologically similar species which coexist in the same habitats routinely reduce interspecific competition for food by regular spacing of body size. The biggest predator species in the assemblage often differ more from one another in size than the smallest species. When coexisting carnivore species do not differ greatly in size, they commonly show morphological differences related to prey handling that may reduce dietary overlap. If carnivore species are very similar in both size and morphology, competition is avoided by habitat partitioning. Two tyrannosaurid species from the late Campanian Dinosaur Park Formation of western Canada are similar in both size and morphology, suggesting that they were segregated on the basis of habitat and/or biogeographic province. However; consideration of the living-space requirements of predator species of such large body size suggests that this kind of spatial separation would only have been possible had tyrannosaurids been more like ectotherms than endotherms in their metabolic rates. Distribution of different large theropod species across different, and surprisingly small (for the size of the animals) portions of Mesozoic landscapes may also account for the remarkably high diversity of morphologically similar large theropods in other dinosaur faunas.     

Carnivore Use of Avocado Orchards across an Agricultural-Wildland Gradient

Wide-ranging species cannot persist in reserves alone. Consequently, there is growing interest in the conservation value of agricultural lands that separate or buffer natural areas. The value of agricultural lands for wildlife habitat and connectivity varies as a function of the crop type and landscape context, and quantifying these differences will improve our ability to manage these lands more effectively for animals. In southern California, many species are present in avocado orchards, including mammalian carnivores. We examined occupancy of avocado orchards by mammalian carnivores across agricultural-wildland gradients in southern California with motion-activated cameras. More carnivore species were detected with cameras in orchards than in wildland sites, and for bobcats and gray foxes, orchards were associated with higher occupancy rates. Our results demonstrate that agricultural lands have potential to contribute to conservation by providing habitat or facilitating landscape connectivity.     

Predictors of mammal species richness in KwaZulu-Natal,South Africa

The management of multi-functional landscapes warrants better knowledge of environment-richness associations at varying disturbance levels and habitat gradients. Intensive land-use patterns for agricultural purposes lead to fragmentation of natural habitat resulting in biodiversity loss that can be measured using landscape metrics to assess mammalian richness. Since carnivores and herbivores are likely to show different responses to disturbance, we calculated carnivore, non-carnivore, and total mammal species richness from camera surveys using a first order Jackknife Estimator. Richness was compared along a habitat gradient comprising coastal forest, Acacia thicket, and highland in KwaZulu-Natal, South Africa. We used standardized OLS regression models to identify climatic and disturbance variables, and landscape metrics as predictors of species richness. The estimated total and non-carnivore species richness were highest in coastal forest, while carnivore species richness was highest in highland followed by coastal forest and Acacia thicket. Average monthly maximum temperature was a significant predictor of all richness groups, and precipitation of the wettest month and isothermality determined total and non-carnivore species richness, respectively. These climatic variables possibly limit species distribution because of physiological tolerance of the species. Total mammal richness was determined by mean shape (+) and habitat division (−) while diversity (+) and patch richness (−) explained carnivore species richness. Mean shape index (+) influenced non-carnivore richness. However, habitat division and patch richness negatively influenced total mammal richness. Though habitat patch size and contiguity had a weak positive prediction, these metrics demonstrated the importance of habitat connectivity for maintaining mammal richness. The identification of these climatic and landscape patterns is important to facilitate future landscape management for mammal conservation in forest-mosaics.     

Spatial variation in the density and vulnerability of preferred prey in the landscape shape patterns of Amur tiger habitat use

Theoretical and empirical research suggests that carnivore distributions are largely determined by prey availability. Availability depends not only on prey density but also on prey accessibility which is affected, in part, by the configuration of landscape attributes that make prey vulnerable to predation. Exactly how spatial variation in these processes shape patterns of carnivore habitat use at the home range scale remains poorly understood. We examined the influence of prey density (negative binomial resource selection function) and vulnerability (kill site resource selection function), mapped separately for each of three species of primary prey, on habitat use patterns within the home range for Amur tigers Panthera tigris altaica in Far East Russia over 20 winters. We developed spatially‐explicit mixed linear regression models to assess these patterns and found that models with parameters for specific primary prey were more robust than models with composite parameters for all primary prey species. This emphasizes the importance of evaluating predation dynamics at a species‐specific level. We also found that Amur tigers used habitat within the home range where red deer Cervus elaphus and wild boar Sus scrofa were dense. These two species were clearly preferred by tigers accounting for 72% (201 of the 278) of the tiger kills detected. The effect of red deer density however, was modulated by the vulnerability of red deer in the landscape. Amur tigers tended to establish their home ranges on habitat where red deer were most vulnerable to predation, but would use habitat where red deer were dense in the peripheral regions of their home ranges. This suggests that tigers may utilize two separate strategies for acquiring prey. As the configuration of resource patches within the home range influences carnivore survival and reproduction, our analysis has implications for tiger conservation that extend beyond our improved understanding of tiger‐prey ecology.     

Multi-criteria spatial identification of carnivore conservation areas under data scarcity and conflict: a jaguar case study in Sierra Nevada de Santa Marta,Colombia

Human–wildlife conflict, habitat loss, and prey hunting are the main threats to carnivore species worldwide. Forest conversion as consequence of deforestation and agricultural expansion increases the proximity between carnivores and humans, thereby escalating conflicts. Knowledge about carnivore species in data-poor countries, such as Colombia, is scarce which has the potential to result in poor landscape planning decisions. For many species, the only existing spatial information resides in expert-driven approaches which result in coarse-resolution ‘extent-of-occurrence’ maps. There is an increasing need for the development of methodologies to identify conservation and management areas at appropriate scales. Multi-criteria approaches will allow the inclusion of diverse species attributes enabling environmental institutions to address complex landscape decisions that result in conservation and management of carnivore habitat. We present a multi-criteria spatial identification tool for conservation and management areas, focused on Jaguars (Panthera onca) in the Sierra Nevada de Santa Marta, in northern Colombia. Our approach identifies areas based on the relationship between three spatial criteria: (1) suitable habitat patches, (2) habitat connectivity, and (3) zones of higher likelihood of human–jaguar conflict. We identified areas with the presence of at least one spatial criteria in 32% of the study area. Only 16.28% of these occur within protected areas (PAs) and the remaining fall on private lands (83.72%), either within (35.68%) or outside (48.04%) buffer zones of PAs. Our results highlight the need for multi-stakeholder collaborative approaches given that most proposed conservation areas fall on private rather than public lands.     

Changes in home range sizes and population densities of carnivore species along the natural to urban habitat gradient

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The priority value of scrubland habitats for carnivore conservation in Mediterranean ecosystems

Carnivores are umbrella species with a key role in ecosystems. In the Mediterranean, they face with several conservation problems, mainly habitat loss or transformation. In this region, scrubland habitats are considered of minor conservation relevance as compared to mature forest formations. Conservation of scrublands in Mediterranean ecosystems is also difficult because to control fires, they are removed over large areas. For carnivores, scrublands may be essential to guarantee shelter and food. Here, we analyzed the importance of scrublands and other habitat variables in determining the richness of medium-sized carnivores in a typical Mediterranean area of central Spain (Monfragüe Natural Park). The Park was divided into plots of 2 × 2 km (n = 30). In each plot, a 2 km-length survey route was walked searching for carnivore scats. We recorded the number of species of carnivores and we related this value to several habitat variables. Habitat variables were summarized by means of Principal Component Analysis (PCA). Habitat models were developed by, using multiple regression models, including PCA factors and the habitat type as predictors. The only variable included in the final model was the first PCA factor. Carnivore richness was positively associated with large tree, shrub, and rock cover areas, which is the typical habitat structure of scrublands. This habitat offers the best available area for shelter and may be considered as a key element for carnivore conservation in the Mediterranean region. To reconciliate carnivore conservation and forestry management, we proposed small-scale clearances as a management alternative to typical large scrubland removals.     

Insect diversity of a Costa Rican dry forest: why keep it,and how?

The dry forest of the 108 km2 Santa Rosa National Park in northwestern Costa Rica contains as many as 13 000 species of insects (including 3140 species of moths and butterflies) sustained by and sustaining about 700 species of plants and 400 species of vertebrates. These insects require explicit conservation attention. They are more than just decorations on the plants; rather, they are the building blocks and glue for much of the habitat. They are the food for much of the carnivore community, and the insect species are not merely interchangeable bits of nutrients. They are major killers of seeds, and thereby influence both the plant species composition of a habitat and prevent the better competitors from taking it over. They are the primary pollinators and are specific enough to be neither interchangeable nor replaceable with other animals; the seeds resulting from their pollination activities are major animal foods in the habitat. They are a diverse, puzzling, complex, intrinsically attractive, and major part of the intellectual display offered by tropical wildlands, the display that will be the eventual foundation for most of the reason why tropical wildlands will be retained as such in the future.
The retention and maintenance of insect species richness in a tropical wildland is strongly rooted in the preservation of plant species richness, in maintaining habitat mosaics (different members of which are used by a given insect in different seasons), in preserving a large diversity of habitats, and in recognizing the threat posed by insect crop associates in nearby agricultural lands.     

An ecological reassessment of the southern African carnivore guild: a case study from Member 4, Sterkfontein, South Africa

The southern African late Pliocene to early Pleistocene carnivore guild was much larger than that of the present day. Understanding how this guild may have functioned is important for the reconstruction of carnivore-hominin interactions and to assess the potential for hominin scavenging in southern Africa. In modern ecosystems, the coexistence of larger carnivore species is constrained by several factors, which include high levels of interspecific competition. Here, the composition of the fossil carnivore guild is examined using Sterkfontein Member 4 (Cradle of Humankind, South Africa) as a case study. Sterkfontein Member 4 contains 10 larger carnivore taxa (body mass > 21.5 kg) and may also contain two Australopithecus species. Two possible causes of higher numbers of carnivore species in the South African fossil record are initially considered. First, that there is a bias introduced through comparing assemblages of differing sizes; second, carnivore biodiversity may have been artificially inflated due to previous taxonomic splitting of carnivore species, such as Crocuta. These possibilities are rejected and modern ecological data are used to construct a simple spatial model to determine how many carnivores could have co-existed. Although the resulting model indicates that the carnivore taxa present in Member 4 could have co-occurred, modern ecological studies indicate that it is highly unlikely that they would have co-existed simultaneously. Considering the complex depositional processes that operate in the southern African cave sites, it is proposed that the larger carnivore guild observed in the Sterkfontein Member 4 fossil assemblage is a palimpsest created by time-averaging. In light of this, we suggest that sites which have a large number of carnivore taxa should be examined for time-averaging, while those sites which have relatively few species may be a better reflection of carnivore communities.     

Global patterns of fragmentation and connectivity of mammalian carnivore habitat

Although mammalian carnivores are vulnerable to habitat fragmentation and require landscape connectivity, their global patterns of fragmentation and connectivity have not been examined. We use recently developed high-resolution habitat suitability models to conduct comparative analyses and to identify global hotspots of fragmentation and connectivity for the world's terrestrial carnivores. Species with less fragmentation (i.e. more interior high-quality habitat) had larger geographical ranges, a greater proportion of habitat within their range, greater habitat connectivity and a lower risk of extinction. Species with higher connectivity (i.e. less habitat isolation) also had a greater proportion of high-quality habitat, but had smaller, not larger, ranges, probably reflecting shorter distances between habitat patches for species with restricted distributions; such species were also more threatened, as would be expected given the negative relationship between range size and extinction risk. Fragmentation and connectivity did not differ among Carnivora families, and body mass was associated with connectivity but not fragmentation. On average, only 54.3 per cent of a species' geographical range comprised high-quality habitat, and more troubling, only 5.2 per cent of the range comprised such habitat within protected areas. Identification of global hotspots of fragmentation and connectivity will help guide strategic priorities for carnivore conservation.     

Demographic consequences of habitat loss and crowding in large carnivores: A natural experiment

Habitat loss is one of largest threats to the persistence of large carnivore populations. However, because most large carnivores are long‐lived, cryptic and wide‐ranging, few studies examine the demographic consequences of habitat loss, and the resultant crowding that ensues, on these species. We used long‐term data to examine the demographic responses of an African lion (Panthera leo) population to flooding‐induced habitat loss in a seasonal wetland, the Okavango Delta, during a transition from low to high annual flooding patterns. We found that intraspecific competition reduced cub survival and that this effect was exacerbated by habitat loss and consequent crowding. The proportion of cubs recorded in the population also declined as crowding increased, and both the survival of cubs and proportion of cubs recorded declined as prey abundance decreased. Apparent sub‐adult survival declined with increasing pride size, but this likely reflects emigration rather than mortality. Adult survival remained relatively constant throughout the study period, a population response which is important in buffering populations against short‐term fluctuations in ecological conditions. As many large carnivore populations face future habitat loss, it is important that we understand the demographic consequences of habitat loss in order to better mitigate its effects in the future.