Bottom-up rather than top-down mechanisms determine mesocarnivore interactions in Norway.

Interactions among coexisting mesocarnivores can be influenced by different factors such as the presence of large carnivores, land-use, environmental productivity, or human disturbance. Disentangling the relative importance of bottom-up and top-down processes can be challenging, but it is important for biodiversity conservation and wildlife management. The aim of this study was to assess how the interactions among mesocarnivores (red fox Vulpes vulpes, badger Meles meles, and pine marten Martes martes) were affected by large carnivores (Eurasian lynx Lynx lynx and wolf Canis lupus), land cover variables (proportion of agricultural land and primary productivity), and human disturbance, as well as how these top-down and bottom-up mechanisms were influenced by season. We analyzed 3 years (2018-2020) of camera trapping observations from Norway and used structural equation models to assess hypothesized networks of causal relationships. Our results showed that land cover variables were more strongly associated with mesocarnivore detection rates than large carnivores in Norway. This might be caused by a combination of low density of large carnivores in an unproductive ecosystem with strong seasonality. Additionally, detection rates of all mesocarnivores showed positive associations among each other, which were stronger in winter. The prevalence of positive interactions among predators might indicate a tendency to use the same areas and resources combined with weak interference competition. Alternatively, it might indicate some kind of facilitative relationship among species. Human disturbance had contrasting effects for different species, benefiting the larger mesocarnivores (red fox and badger) probably through food subsidization, but negatively affecting apex predators (wolf and lynx) and smaller mesocarnivores (pine marten). In a human-dominated world, this highlights the importance of including anthropogenic influences in the study of species interactions.

Scavenger community structure along an environmental gradient from boreal forest to alpine tundra in Scandinavia.

Scavengers can have strong impacts on food webs, and awareness of their role in ecosystems has increased during the last decades. In our study, we used baited camera traps to quantify the structure of the winter scavenger community in central Scandinavia across a forest-alpine continuum and assess how climatic conditions affected spatial patterns of species occurrences at baits. Canonical correspondence analysis revealed that the main habitat type (forest or alpine tundra) and snow depth was main determinants of the community structure. According to a joint species distribution model within the HMSC framework, species richness tended to be higher in forest than in alpine tundra habitat, but was only weakly associated with temperature and snow depth. However, we observed stronger and more diverse impacts of these covariates on individual species. Occurrence at baits by habitat generalists (red fox, golden eagle, and common raven) typically increased at low temperatures and high snow depth, probably due to increased energetic demands and lower abundance of natural prey in harsh winter conditions. On the contrary, occurrence at baits by forest specialists (e.g., Eurasian jay) tended to decrease in deep snow, which is possibly a consequence of reduced bait detectability and accessibility. In general, the influence of environmental covariates on species richness and occurrence at baits was lower in alpine tundra than in forests, and habitat generalists dominated the scavenger communities in both forest and alpine tundra. Following forecasted climate change, altered environmental conditions are likely to cause range expansion of boreal species and range contraction of typical alpine species such as the arctic fox. Our results suggest that altered snow conditions will possibly be a main driver of changes in species community structure.

Multiple factors influence local perceptions of snow leopards and Himalayan wolves in the central Himalayas, Nepal.

An understanding of local perceptions of carnivores is important for conservation and management planning. In the central Himalayas, Nepal, we interviewed 428 individuals from 85 settlements using a semi-structured questionnaire to quantitatively assess local perceptions and tolerance of snow leopards and wolves. We used generalized linear mixed effect models to assess influential factors, and found that tolerance of snow leopards was much higher than of wolves. Interestingly, having experienced livestock losses had a minor impact on perceptions of the carnivores. Occupation of the respondents had a strong effect on perceptions of snow leopards but not of wolves. Literacy and age had weak impacts on snow leopard perceptions, but the interaction among these terms showed a marked effect, that is, being illiterate had a more marked negative impact among older respondents. Among the various factors affecting perceptions of wolves, numbers of livestock owned and gender were the most important predictors. People with larger livestock herds were more negative towards wolves. In terms of gender, males were more positive to wolves than females, but no such pattern was observed for snow leopards. People's negative perceptions towards wolves were also related to the remoteness of the villages. Factors affecting people's perceptions could not be generalized for the two species, and thus need to be addressed separately. We suggest future conservation projects and programs should prioritize remote settlements.

The impact of leopards (Panthera pardus) on livestock losses and human injuries in a human-use landscape in Maharashtra, India.

There are many ways in which large carnivores and humans interact in shared spaces. In this study we provide insights into human-leopard relationships in an entirely modified, human-dominated landscape inhabited by dense populations of humans (266 per km2), their livestock (162 per km2) and relatively high densities of large predators (10 per 100 km2). No human deaths were recorded, and livestock losses to leopards numbered only 0.45 per km2per year (averaged over three years) despite the almost complete dependency of leopards on domestic animals as prey. Predation was not the major cause of livestock mortality as diseases and natural causes caused higher losses (80% of self-reported losses). We also found that ineffective night time livestock protection and the presence of domestic dogs increased the probability of a farmer facing leopard attacks on livestock. Resident farmers faced much lower livestock losses to leopard predation in contrast to the migratory shepherds who reported much higher losses, but rarely availed of the government compensation schemes. We recommend that local wildlife managers continue to shift from reactive measures such as leopard captures after livestock attacks to proactive measures such as focusing on effective livestock protection and informing the affected communities about safety measures to be taken where leopards occur in rural landscapes. The natural causes of livestock deaths due do diseases may be better prevented by involving animal husbandry department for timely vaccinations and treatment.

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.

The alternative prey hypothesis revisited: Still valid for willow ptarmigan population dynamics.

The alternative prey hypothesis predicts that the interaction between generalist predators and their main prey is a major driver of population dynamics of alternative prey species. In Fennoscandia, changes in climate and human land use are assumed to alter the dynamics of cyclic small rodents (main prey) and lead to increased densities and range expansion of an important generalist predator, the red fox Vulpes vulpes. In order to better understand the role of these potential changes in community structure on an alternative prey species, willow ptarmigan Lagopus lagopus, we analyzed nine years of population census data from SE Norway to investigate how community interactions affected their population dynamics. The ptarmigan populations showed no declining trend during the study period, and annual variations corresponded with marked periodic small rodent peaks and declines. Population growth and breeding success were highly correlated, and both demographic variables were influenced by an interaction between red fox and small rodents. Red foxes affected ptarmigan negatively only when small rodent abundance was low, which is in accordance with the alternative prey hypothesis. Our results confirm the important role of red fox predation in ptarmigan dynamics, and indicate that if small rodent cycles are disrupted, this may lead to decline in ptarmigan and other alternative prey species due to elevated predation pressure.

Variation in home range size of red foxes Vulpes vulpes along a gradient of productivity and human landscape alteration.

Home range size is a fundamental concept for understanding animal dispersion and ecological needs, and it is one of the most commonly reported ecological attributes of free-ranging mammals. Previous studies indicate that red foxes Vulpes vulpes display great variability in home range size. Yet, there has been little consensus regarding the reasons why home range sizes of red foxes vary so extensively. In this study, we examine possible causes of variation in red fox home range sizes using data from 52 GPS collared red foxes from four study areas representing a gradient of landscape productivity and human landscape alteration in Norway and Sweden. Using 90% Local Convex Hull home range estimates, we examined how red fox home range size varied in relation to latitude, elevation, vegetation zone, proportion of agricultural land and human settlement within a home range, and sex and age. We found considerable variation in red fox home range sizes, ranging between 0.95 km2 to 44 km2 (LoCoH 90%) and 2.4 km2 to 358 km2 (MCP 100%). Elevation, proportion of agricultural land and sex accounted for 50% of the variation in home range size found amongst foxes, with elevation having the strongest effect. Red foxes residing in more productive landscapes (those in more southern vegetation zones), had home ranges approximately four times smaller than the home ranges of foxes in the northern boreal vegetation zone. Our results indicate that home range size was influenced by a productivity gradient at both the landscape (latitude) and the local (elevation) scale. The influence of the proportion of agriculture land on home range size of foxes illustrates how human landscape alteration can affect the space use and distribution of red foxes. Further, the variation in home range size found in this study demonstrates the plasticity of red foxes to respond to changing human landscape alteration as well as changes in landscape productivity, which may be contributing to red fox population increases and northern range expansions.

The range of the mange: Spatiotemporal patterns of sarcoptic mange in red foxes (Vulpes vulpes) as revealed by camera trapping.

Sarcoptic mange is a widely distributed disease that affects numerous mammalian species. We used camera traps to investigate the apparent prevalence and spatiotemporal dynamics of sarcoptic mange in a red fox population in southeastern Norway. We monitored red foxes for five years using 305 camera traps distributed across an 18000 km2 area. A total of 6581 fox events were examined to visually identify mange compatible lesions. We investigated factors associated with the occurrence of mange by using logistic models within a Bayesian framework, whereas the spatiotemporal dynamics of the disease were analysed with space-time scan statistics. The apparent prevalence of the disease fluctuated over the study period with a mean of 3.15% and credible interval [1.25, 6.37], and our best logistic model explaining the presence of red foxes with mange-compatible lesions included time since the beginning of the study and the interaction between distance to settlement and season as explanatory variables. The scan analyses detected several potential clusters of the disease that varied in persistence and size, and the locations in the cluster with the highest probability were closer to human settlements than the other survey locations. Our results indicate that red foxes in an advanced stage of the disease are most likely found closer to human settlements during periods of low wild prey availability (winter). We discuss different potential causes. Furthermore, the disease appears to follow a pattern of small localized outbreaks rather than sporadic isolated events.

Snow Leopard and Himalayan Wolf: Food Habits and Prey Selection in the Central Himalayas, Nepal.

Top carnivores play an important role in maintaining energy flow and functioning of the ecosystem, and a clear understanding of their diets and foraging strategies is essential for developing effective conservation strategies. In this paper, we compared diets and prey selection of snow leopards and wolves based on analyses of genotyped scats (snow leopards n = 182, wolves n = 57), collected within 26 sampling grid cells (5×5 km) that were distributed across a vast landscape of ca 5000 km2 in the Central Himalayas, Nepal. Within the grid cells, we sampled prey abundances using the double observer method. We found that interspecific differences in diet composition and prey selection reflected their respective habitat preferences, i.e. snow leopards significantly preferred cliff-dwelling wild ungulates (mainly bharal, 57% of identified material in scat samples), whereas wolves preferred typically plain-dwellers (Tibetan gazelle, kiang and argali, 31%). Livestock was consumed less frequently than their proportional availability by both predators (snow leopard = 27%; wolf = 24%), but significant avoidance was only detected among snow leopards. Among livestock species, snow leopards significantly preferred horses and goats, avoided yaks, and used sheep as available. We identified factors influencing diet composition using Generalized Linear Mixed Models. Wolves showed seasonal differences in the occurrence of small mammals/birds, probably due to the winter hibernation of an important prey, marmots. For snow leopard, occurrence of both wild ungulates and livestock in scats depended on sex and latitude. Wild ungulates occurrence increased while livestock decreased from south to north, probably due to a latitudinal gradient in prey availability. Livestock occurred more frequently in scats from male snow leopards (males: 47%, females: 21%), and wild ungulates more frequently in scats from females (males: 48%, females: 70%). The sexual difference agrees with previous telemetry studies on snow leopards and other large carnivores, and may reflect a high-risk high-gain strategy among males.

Adaptable neighbours: movement patterns of GPS-collared leopards in human dominated landscapes in India.

Understanding the nature of the interactions between humans and wildlife is of vital importance for conflict mitigation. We equipped five leopards with GPS-collars in Maharashtra (4) and Himachal Pradesh (1), India, to study movement patterns in human-dominated landscapes outside protected areas. An adult male and an adult female were both translocated 52 km, and exhibited extensive, and directional, post release movements (straight line movements: male = 89 km in 37 days, female = 45 km in 5 months), until they settled in home ranges of 42 km2 (male) and 65 km2 (female). The three other leopards, two adult females and a young male were released close to their capture sites and used small home ranges of 8 km2 (male), 11 km2 and 15 km2 (females). Movement patterns were markedly nocturnal, with hourly step lengths averaging 339±9.5 m (SE) during night and 60±4.1 m during day, and night locations were significantly closer to human settlements than day locations. However, more nocturnal movements were observed among those three living in the areas with high human population densities. These visited houses regularly at nighttime (20% of locations <25 m from houses), but rarely during day (<1%). One leopard living in a sparsely populated area avoided human settlements both day and night. The small home ranges of the leopards indicate that anthropogenic food resources may be plentiful although wild prey is absent. The study provides clear insights into the ability of leopards to live and move in landscapes that are extremely modified by human activity.

Bears are simply voles writ large: social structure determines the mechanisms of intrinsic population regulation in mammals.

The literature reveals opposing views regarding the importance of intrinsic population regulation in mammals. Different models have been proposed; adding importance to contrasting life histories, body sizes and social interactions. Here we evaluate current theory based on results from two Scandinavian projects studying two ecologically different mammal species with contrasting body sizes and life history traits: the root vole Microtus oeconomus and the brown bear Ursus arctos. We emphasize four inter-linked behavioral aspects-territoriality, dispersal, social inhibition of breeding, and infanticide-that together form a density-dependent syndrome with potentially regulatory effects on population growth. We show that the two species are similar in all four behaviors and thus the overall regulatory syndrome. Females form matrilineal assemblages, female natal dispersal is negatively density dependent and breeding is suppressed in philopatric young females. In both species, male turnover due to extrinsic mortality agents cause infanticide with negative effects on population growth. The sex-biased and density-dependent dispersal patterns promote the formation of matrilineal clusters which, in turn, leads to reproductive suppression with potentially regulatory effects. Hence, we show that intrinsic population regulation interacting with extrinsic mortality agents may occur irrespective of taxon, life history and body size. Our review stresses the significance of a mechanistic approach to understanding population ecology. We also show that experimental model populations are useful to elucidate natural populations of other species with similar social systems. In particular, such experiments should be combined with methodical innovations that may unravel the effects of cryptic intrinsic mechanisms such as infanticide.

Concurrent effects of age class and food distribution on immigration success and population dynamics in a small mammal.

During the settlement stage of dispersal, the outcome of conflicts between residents and immigrants should depend on the social organization of resident populations as well as on individual traits of immigrants, such as their age class, body mass and/or behaviour. We have previously shown that spatial distribution of food influences the social organization of female bank voles (Myodes glareolus). Here, we aimed to determine the relative impact of food distribution and immigrant age class on the success and demographic consequences of female bank vole immigration. We manipulated the spatial distribution of food within populations having either clumped or dispersed food. After a pre-experimental period, we released either adult immigrants or juvenile immigrants, for which we scored sociability and aggressiveness prior to introduction. We found that immigrant females survived less well and moved more between populations than resident females, which suggest settlement costs. However, settled juvenile immigrants had a higher probability to reproduce than field-born juveniles. Food distribution had little effects on the settlement success of immigrant females. Survival and settlement probabilities of immigrants were influenced by adult female density in opposite ways for adult and juvenile immigrants, suggesting a strong adult-adult competition. Moreover, females of higher body mass at release had a lower probability to survive, and the breeding probability of settled immigrants increased with their aggressiveness and decreased with their sociability. Prior to the introduction of immigrants, resident females were more aggregated in the clumped food treatment than in the dispersed food treatment, but immigration reversed this relationship. In addition, differences in growth trajectories were seen during the breeding season, with populations reaching higher densities when adult immigrants were introduced in a plot with dispersed food, or when juvenile immigrants were introduced in a plot with clumped food. These results indicate the relative importance of intrinsic and extrinsic factors on immigration success and demographic consequences of dispersal and are of relevance to conservation actions, such as reinforcement of small populations.

Big cats in our backyards: persistence of large carnivores in a human dominated landscape in India.

Protected areas are extremely important for the long term viability of biodiversity in a densely populated country like India where land is a scarce resource. However, protected areas cover only 5% of the land area in India and in the case of large carnivores that range widely, human use landscapes will function as important habitats required for gene flow to occur between protected areas. In this study, we used photographic capture recapture analysis to assess the density of large carnivores in a human-dominated agricultural landscape with density >300 people/km(2) in western Maharashtra, India. We found evidence of a wide suite of wild carnivores inhabiting a cropland landscape devoid of wilderness and wild herbivore prey. Furthermore, the large carnivores; leopard (Panthera pardus) and striped hyaena (Hyaena hyaena) occurred at relatively high density of 4.8±1.2 (sd) adults/100 km(2) and 5.03±1.3 (sd) adults/100 km(2) respectively. This situation has never been reported before where 10 large carnivores/100 km(2) are sharing space with dense human populations in a completely modified landscape. Human attacks by leopards were rare despite a potentially volatile situation considering that the leopard has been involved in serious conflict, including human deaths in adjoining areas. The results of our work push the frontiers of our understanding of the adaptability of both, humans and wildlife to each other's presence. The results also highlight the urgent need to shift from a PA centric to a landscape level conservation approach, where issues are more complex, and the potential for conflict is also very high. It also highlights the need for a serious rethink of conservation policy, law and practice where the current management focus is restricted to wildlife inside Protected Areas.

Guild composition and habitat use of voles in 2 forest landscapes in south-eastern Norway.

It is widely believed that intensive forestry has influenced small mammal population dynamics, and thereby the entire mammalian community in Fennoscandian boreal forests. The nature of these impacts on the different species is subject to debate. We live-trapped voles between 2006 and 2009 in 2 commercially harvested forests in south-eastern Norway. We investigated the variation in vole abundance among habitat types (e.g. mature forest and clear-cut) and the hypothesis that graminivorous species such as field voles (Microtus agrestis L.) benefit from clear-cuts at the expense of forest dwellers (i.e. the bank vole, Myodes glareolus Schreb.), using fine-scale descriptors of the ground vegetation. We could not find support for the hypothesis that field voles show a preference for clear-cuts, and their overall abundance was low, while bank voles were the dominant species in all habitat types, including clear-cuts in the peak and pre-peak years. We found a positive association between bank vole abundance and bilberry (Vaccinium myrtillus L.) availability rather than a specific habitat type. Low field vole density in clear-cuts might be due to variation in local productivity and ground vegetation as well as to large variation in the species temporal dynamics. The latter is particularly associated with the widespread decline of field voles in Scandinavia. Logging has the potential to negatively affect bank vole population dynamics because of the negative effect on bilberry development.

Translocation as a tool for mitigating conflict with leopards in human-dominated landscapes of India.

We examined the efficacy of a translocation program in which large numbers of leopards (Panthera pardus fusca) were trapped in human-dominated landscapes where livestock attacks were common and human attacks rare and released into adjoining forested areas in an attempt to reduce leopard presence and mitigate conflicts at the capture site. In the year starting in February of 2001, 29 leopards were captured in the human-dominated rural landscape of the Junnar region (4275 km(2) , 185 people/km(2) ), Maharashtra, India, and released an average of 39.5 km away in adjoining forests. Eleven leopards were also relocated to the same forests from other districts. Prior to the large-scale translocation program, an average of four leopard attacks on humans occurred each year between 1993 and 2001. After the translocation program was initiated, the average increased substantially to 17 attacks. Linear and logistic models showed that attack frequency increased in Junnar following nearby releases of leopards and decreased when leopards were removed for releases far away; that attacks became more lethal when the number of leopards introduced from other districts increased; and that attacks were most likely to occur in the regions where the largest number of leopards had been introduced from other areas. These results suggest that leopards did not stay at the release sites and that translocation induced attacks on people. Potential explanations for these results include increased aggression induced by stress of the translocation process, movement through unfamiliar human-dominated landscapes following release, and loss of fear of humans due to familiarity with humans acquired during captivity. Our results show that reactive solutions to attacks on humans by leopards, such as translocation, could in fact increase human-leopard conflict. Measures to reduce human-carnivore conflicts may include more effective compensation procedures to pay livestock owners for the loss of animals to predation by carnivores, providing better methods of protection for livestock, and encouraging greater social acceptance of the presence of carnivores in human-dominated landscapes.

Predicting spacing behavior and mating systems of solitary cervids: a study of hog deer and Indian muntjac.

This study investigates the validity of current theory for predicting ecological and allometric effects on space use, social structure and mating systems of poorly known solitary cervids, based on a comparative analysis of radio-telemetry data on hog deer Axis porcinus (N=32) and Indian muntjac Muntiacus muntjak (N=28). The larger and sexually size-dimorphic hog deer inhabit highly productive alluvial floodplains, where resource distribution is patchy and spatiotemporally unpredictable. As predicted for this species, site fidelity was low and range sizes varied among sex and age groups and among seasons. Hog deer were probably non-territorial, as home range sizes seemed too large to be exclusive when taking into account their high population density. Extensive movements of adult males during the rut implied "roaming" as a mating strategy. The smaller, forest-dwelling and sexually size-monomorphic muntjacs inhabit a more uniform and stable habitat. As predicted, muntjacs exhibited higher site fidelity than hog deer, and no seasonal variations in home range sizes. Adults exhibited relatively large home range overlap, both inter- and intrasexually. Hence, strict territoriality did not occur, but their well-defined home ranges and high site fidelity indicated some form of site-specific dominance. In conclusion, habitat characteristics were appropriate predictors of home range sizes and site fidelity. Body mass appeared to be a suitable predictor of intraspecific patterns in space use but a poor predictor of interspecific patterns, probably due to a confounding effect of habitat productivity.