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The study of attitudes toward wild elephants and human–elephant conflict (HEC) is vital to understanding what attitudes are held by local people and how to incorporate them into wild elephant conservation. This study investigated the interlinkages between the HEC experience and local people’s attitudes toward the conservation of wild elephants and which exploratory factors influence these attitudes. We used a Bayesian Belief Network (BBN) framework to highlight the interlinkages between HEC and people’s attitudes. The basic BBN model indicated that HEC was central in determining the attitudes of local people. Although people generally hold positive attitudes toward elephants, people support having elephants in the country but not in their own regions. Conservation willingness was not obvious due to the lack of deliberate assistance from the government to the affected communities. We suggest implementing education programs to promote local awareness of conflict mitigation techniques. Bayesian belief network(BBN); conservation; elephas maximus; human–elephant conflict (HEC); wildlife

Understanding the underlying causes behind human–elephant conflict (HEC)-driven mortality of humans and elephants will help improve both parties’ wellbeing. The objective of this study was to examine the temporal and spatial mortality patterns of humans and elephants and the influence of local attitudes, conflict factors and habitat factors on elephant poaching. We used the Myanmar Forest Department data from 2001 to 2020 for humans and 2011 to 2020 for elephants together with explanatory data on human attitudes, habitat, and conflict factors. Approximately seven persons were killed annually in elephant attacks, with a bias towards men. The annual mortality of elephants during the study period was on average 16 individuals, and most elephants were killed by humans. There was a significant relationship between the number of killed humans and human-killed elephants around HEC villages. Villages with more property damage exhibited a higher rate of human mortality, which also correlated with negative feelings of local people towards elephants. Elephant poaching was higher in villages with less suitable habitat available for elephant use. Human encroachment is an important cause of HEC, leading to human loss and forming the main threat to the survival of wild elephants. We suggest local involvement to ensure good governance in conflict resolution and mitigation strategies and to strengthen law enforcement. Asian elephant (Elephas maximus) Human–elephant conflict (HEC) Attack Mortality Poaching

Quantifying the total number of individuals (abundance) of species is the basis for spatial ecology and biodiversity conservation. Abundance data are mostly collected through professional surveys as part of monitoring programs, often at a national level. These surveys rarely follow exactly the same sampling protocol in different countries, which represents a challenge for producing biogeographical abundance maps based on the transboundary information available covering more than one country. Moreover, not all species are properly covered by a single monitoring scheme, and countries typically collect abundance data for target species through different monitoring schemes. We present a new methodology to model total abundance by merging count data information from surveys with different sampling protocols. The proposed methods are used for data from national breeding bird monitoring programs in Norway and Sweden. Each census collects abundance data following two different sampling protocols in each country, i.e., these protocols provide data from four different sampling processes. The modeling framework assumes a common Gaussian Random Field shared by both the observed and true abundance with either a linear or a relaxed linear association between them. The models account for particularities of each sampling protocol by including terms that affect each observation process, i.e., accounting for differences in observation units and detectability. Bayesian inference is performed using the Integrated Nested Laplace Approximation (INLA) and the Stochastic Partial Differential Equation (SPDE) approach for spatial modeling. We also present the results of a simulation study based on the empirical census data from mid-Scandinavia to assess the performance of the models under model misspecification. Finally, maps of the expected abundance of birds in our study region in mid-Scandinavia are presented with uncertainty estimates. We found that the framework allows for consistent integration of data from surveys with different sampling protocols. Further, the simulation study showed that models with a relaxed linear specification are less sensitive to misspecification, compared to the model that assumes linear association between counts. Relaxed linear specifications of total bird abundance in mid-Scandinavia improved both goodness of fit and the predictive performance of the models. Supplementary materials accompanying this paper appear on-line. Data integration; Joint species distribution models; Bayesian statistics; Latent Gaussian Models; Gaussian Random Fields

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This study aimed to understand the movement behaviour and utilization distributions of Kori bustards in space and time in the Serengeti ecosystem. A total of 14 individuals were tracked with the aid of GPS (Geographical positioning system) satellite transmitters, and their sexes were identified using DNA analysis. A species utilization distribution was estimated using the Brownian bridge movement model (hereafter dBBMM) in which the probability of being in an area is conditioned by starting and ending (GPS) relocations. Resource selections were analysed by comparing the GPS relocations with locations randomly placed within each individual’s region of utilization in a spatio-temporal approach. Vegetation information was derived from a Serengeti GIS vegetation map and Data Centre and was reclassified as Open grassland, Dense grassland, Shrubbed grassland, Treed grassland, Shrubland, and Woodland. The Shannon diversity index for vegetation was calculated based on the original vegetation classification. Used versus non-used habitats were contrasted using a generalized linear mixed-effects model with a binomial distribution. The results indicated that males were 21.5% more mobile than females, and movements were 6.3% more diffuse during the nonbreeding period compared to the breeding period (7.59 versus 7.14, respectively). Contrasting models indicated that males preferred more open grasslands during the non-breeding period and also preferred closed and shrubbed grassland during the breeding period. Females preferred more woody vegetation during the non-breeding season compared to the breeding season. The most parsimonious model indicated that females preferred to stay closer to rivers and diverse areas during the non-breeding period whereas males preferred areas that were farther from rivers and homogenous. Homogeneous areas were preferred during the breeding period, and heterogeneous areas were preferred during the non-breeding period. We conclude that the movement behaviours of Kori bustards changes with the season and habitat. Further research is needed to understand the factors driving the seasonal movement of Kori bustards in the Serengeti ecosystem. © 2019 Mmassy et al. This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.

Adaptive management (AM) is a systematic process intended to improve policies and practices and reduce scientific uncertainty by learning from the outcome of management decisions. Although many nations are considering the use of AM for wind energy, its application in practice and in policy has been limited. Recent applications of AM have revealed fundamental differences in the definition of AM, its applications, and the projects or planning processes to which it might be applied. This chapter suggests the need for a common understanding and definition of and framework for AM and its application to wind energy. We discuss a definition of AM and technical guidance created by the United States (US) Department of the Interior’s (DOI’s) Adaptive Management Working Group. The chapter also examines how AM has been applied to wind energy development in several European nations and in the USA. The challenges and opportunities associated with implementation of AM for wind development are addressed, management actions in nations that exhibit attributes of AM are compared, and pathways to appropriate application and potential broader use of AM are explored.

Understanding how anthropogenic activities and management actions influence the delivery of ecosystem services is complicated by the interrelated nature of diverse factors. We present a Bayesian Belief Network to highlight the likely consequences of a set of interventions on four wildlife-related ecosystem (dis)services and for supporting biodiversity and human welfare in the Greater Serengeti-Mara Ecosystem. According to the model, core protected areas are important for biodiversity and safari tourism provision. In adjacent game reserves safari tourism opportunities may be hampered by trophy and bushmeat hunting causing fear in wildlife. Most multipleuse areas strike a good balance between the costs and benefits derived from wildlife. Loliondo, however, requires drastic changes in management to either maximize green value creation or sustainable welfare. Although further globalization is expected to render highest levels of welfare, this will be at the expense of biodiversity and related ecosystem services. An online version of the model is available (https://africanbioservices.shinyapps.io/ servicescape) to interactively explore five future scenarios with alternative management strategies, and visualization of the resultant consequences thereof. Identifying areas of conflicts and potential trade-offs between ecosystem (dis)services are crucial to find pathways to nature-based tourism strategies that simultaneously maintain biodiversity and promote the socioeconomic viability of local communities. Open Access, CC-BY-NC-ND

Societal concerns regarding the negative impacts of wind turbines on species and ecosystems have placed more emphasis on mitigation efforts pre- and post-construction. While the mitigation hierarchy is usually fronted to deal with negative ecological impacts, it is hardly employed accordingly. This calls for the core of the problem to be addressed, namely, the lack of an appropriate framework for mitigation as a concept to properly address ecological impacts caused by wind-power development. In this chapter, mitigation is defined as the intervention(s) implemented to affect the risk of wind turbines impacting species or ecosystems. This concept is placed within a social-ecological context where the consecutive steps of the mitigation hierarchy may be affected by socio-economic, technological or environmental spheres of interest. Decisions relating to mitigation are in principle normative, which necessitates addressing three central ethical questions: (1) In which circumstances should mitigation be implemented? (2) How much mitigation is required? (3) Who is responsible for mitigation? Implementing mitigation requires decision-makers to acknowledge that trouble never comes alone, which requires balancing trade-offs and embracing uncertainty into the decision-making process. Adaptive and participatory management may be the best decision-making framework to do this, as it allows for improved ecological understanding through monitoring and a flexible approach to mitigate locally but manage regionally.

Wind energy development continues to expand rapidly to meet international climate targets. However, these developments impact biodiversity and associated sustainability targets negatively. Environmental impact assessments (EIA) predominately focus on site-specific impacts; failing to account for cumulative impacts at larger spatial scales. Life cycle assessments (LCA) can take such a holistic view, however models quantifying the main impact pathways on biodiversity are still lacking. To address this gap, we present a methodology to quantify habitat loss, disturbance and collision impacts of onshore wind power plants on bird biodiversity globally. By overlaying species ranges with wind-power plant locations, species-area relationships were used to calculate the potentially disappeared fraction of species (PDF) for the three impact pathways. Results revealed habitat-specific bird richness values, at order level, which were greatest in tropical and subtropical regions. Temporal changes in PDF were related to operationalizing wind-power plants. Despite similar PDF curves throughout the time period, disturbance and habitat loss had a greater effect compared to collisions. Annual energy production values strongly affected overall impacts for all impact pathways, which in turn differed between continents. When controlling for continent, bird order rather than country more strongly influenced variation in the pathway-specific PDF per GWh. Our approach represents the first step towards incorporating the impacts of wind power production in strategic environmental planning and the LCA framework. Failing to account for these negates an assessment of the trade-offs between biodiversity and energy production, and therefore the balancing of global sustainability goals. © 2020 The Author(s). Published by Elsevier Inc. This is an open access article under the CC BY license (http://creativecommons.org/licenses/by/4.0/).

Concerns for potential wildlife impacts resulting from land†based and offshore wind energy have created challenges for wind project development. Research is not always adequately supported, results are neither always readily accessible nor are they satisfactorily disseminated, and so decisions are often made based on the best available information, which may be missing key findings. The potential for high impacts to avian and bat species and marine mammals have been used by wind project opponents to stop, downsize, or severely delay project development. The global nature of the wind industry—combined with the understanding that many affected species cross†national boundaries, and in many cases migrate between continents—also points to the need to collaborate on an international level. The International Energy Agency (IEA) Wind Technology Collaborative Programs facilitates coordination on key research issues. IEA Wind Task 34—WREN: Working Together to Resolve Environmental Effects of Wind Energy–is a collaborative forum to share lessons gained from field research and modeling, including management methods, wildlife monitoring methods, best practices, study results, and successful approaches to mitigating impacts and addressing the cumulative effects of wind energy on wildlife. WREN develops products such as white papers, fact sheets, and short science summaries, and is involved in a number of activities including hosting a webinar series and outreach and information dissemination through participation in meetings, workshops, and conferences to increase and expand the knowledge base pertaining to wildlife challenges at wind energy facilities. This information is available on WREN Hub, hosted on the Tethys website. This article is categorized under: Wind Power > Climate and Environment

Social behaviour in solitary carnivores has long been an active area of investigation but for many species remains largely founded in conjecture compared to our understanding of sociality in group-living species. The social organization of the wolverine has, until now, received little attention beyond its portrayal as a typical mustelid social system. In this chapter the authors compile observations of social interactions from multiple wolverine field studies, which are integrated into an ecological framework. An ethological model for the wolverine is proposed that reveals an intricate social organization, which is driven by variable resource availability within extremely large territories and supports social behaviour that underpins offspring development.

© The Ecological Society of America The removal of thousands of structures associated with oil and gas development from the world's oceans is well underway, yet the environmental impacts of this decommissioning practice remain unknown. Similar impacts will be associated with the eventual removal of offshore wind turbines. We conducted a global survey of environmental experts to guide best decommissioning practices in the North Sea, a region with a substantial removal burden. In contrast to current regulations, 94.7% of experts (36 out of 38) agreed that a more flexible case-by-case approach to decommissioning could benefit the North Sea environment. Partial removal options were considered to deliver better environmental outcomes than complete removal for platforms, but both approaches were equally supported for wind turbines. Key considerations identified for decommissioning were biodiversity enhancement, provision of reef habitat, and protection from bottom trawling, all of which are negatively affected by complete removal. We provide recommendations to guide the revision of offshore decommissioning policy, including a temporary suspension of obligatory removal.

Impacts of anthropogenic activities on coastal seabirds might be extensive, especially in the breeding season. Identifying important foraging areas and associated habitats is important for a proper management of seabirds. To identify habitat characteristics driving the selection of foraging sites of breeding European shags Phalacrocorax aristotelis, this study used tracking data (GPS- and TDR-loggers), from 282 individual birds comprising 905 foraging trips and 27,303 dives with known locations, to create habitat selection models. To explore possible effects of regional differences in habitat on foraging behavior, the study was performed at two Norwegian colonies, Sklinna in the Norwegian Sea (65°N, 11°E) and Hornoya in the Barents Sea (70°N, 31°E), with distinct differences in seascape structure and habitat availability. Shags at Sklinna foraged further away from the colony than those at Hornoya but diving depth and duration were similar at the two colonies. In both colonies, sea depth was an important predictor of habitats selected by chick-rearing shags during foraging, with birds preferring shallow depths. At Sklinna, shags also selected for flat areas with high probability of kelp forest occurrence. There was no difference in trip length and duration between sexes, but males dived deeper than females in both colonies. This suggests that males and females might utilize different microhabitats within the same foraging area. The study discusses the application of habitat selection modeling to identify important foraging areas for coastal seabirds, and how this may contribute to the management, conservation and assessment of impacts of human activities.

Human–wildlife interactions are of pressing concern for conservation biologists. We show that African elephants are chronically stressed, have smaller group sizes and may have lower calf recruitment outside of a protected area compared to inside. Additionally, this paper discusses some strengths and weaknesses of using faecal hormone analyses in the field., The conservation of the African savanna elephant (Loxodonta africana) is of prime importance for many African countries. Interactions between elephants and humans are known to induce stress and thereby have the potential to affect elephants’ fitness. In Namibia, anthropogenic disturbances are increasing due to increasing human population size and development, particularly near protected areas, such as national parks. In this study, we investigated elephant stress levels in relation to their land use, specifically their protection status, comparing elephants within Etosha National Park in Namibia with elephants residing outside the park. We noninvasively collected dung samples of 91 elephants and determined the concentration of faecal glucocorticoid metabolites (fGCM), an indicator of physiological stress. Elephants outside the park (N = 35) had significantly higher concentrations of fGCM than those inside ENP (N = 56), suggesting that, despite including community-based conservancies, unprotected areas are more stressful for elephants than protected areas, most likely due to increased interactions with humans. We also found that males had lower fGCM concentrations than females, but no significant effect of age, body size or group size was detected. Additionally, herd sizes were significantly smaller and calf recruitment was potentially lower in unprotected areas. These findings underpin the importance of protected areas such as ENP, while encouraging decision-makers to continue reducing and mitigating potential human-induced disturbances.

Concurrent with the development of wind energy, research activity on wind energy generation and wildlife has evolved significantly during the last decade. This chapter presents an overview of remaining key knowledge gaps, consequent future research directions and their significance for management and planning for wind energy generation. The impacts of wind farms on wildlife are generally site-, species- and season-specific and related management strategies and practices may differ considerably between countries. These differences acknowledge the need to consider potential wildlife impacts for each wind farm project. Still, the ecological mechanisms guiding species’ responses and potential vulnerability to wind farms can be expected to be fundamental in nature. A more cohesive understanding of the causes, patterns, mechanisms, and consequences of animal movement decisions will thereby facilitate successful mitigation of impacts. This requires planning approaches that implement the mitigation hierarchy effectively to reduce risks to species of concern. At larger geographical scales, population-level and cumulative impacts of multiple wind farms (and other anthropogenic activity) need to be addressed. This requires longitudinal and multiple-site studies to identify species-specific traits that influence risk of mortality, notably from collision with wind turbines, disturbance or barrier effects. In addition, appropriate pre- and post-construction monitoring techniques must be utilized. Predictive modelling to forecast risk, while tackling spatio-temporal variability, can guide the mitigation of wildlife impacts at wind farms.

On average, 7.8 white-tailed eagles (Haliaeetus albicilla) collide with wind turbines annually at the operating wind-power plant on the island of Smøla off the coast of central Norway. To better understand the impact of this wind-power plant on white-tailed eagles, we investigated how habitat utilization affected displacement effects. We collected data on habitat utilization in non-territorial subadult white-tailed eagles using global positioning system satellite telemetry (2004–2009). From these data, we estimated utilization distributions using the Brownian bridge movement model and analyzed them using Resource Utilization Functions. Home ranges were circa 10–30% smaller for subadults hatched on skerries and islets farther from the wind-power plant, and tended to be circa 40% larger during spring. Shallow sea, skerries, and islets were utilized extensively, as was marsh, heathland, and forest on the main island of Smøla. We noted lower selectivity for arable land and higher levels of differentiation in utilization among individuals for forest and islets. Females also had a nearly 4 times higher between-individual variation, which was likely due to longranging excursions. The within- and between-individual variation among seasons showed an annual pattern, with increasing between-individual variation toward summer. Displacement (indicated by an overall 40% proportional reduction in utilization) was more pronounced in the birds’ second and third calendar year compared with their first calendar year, and during autumn–winter. Reduced displacement during spring coincides with the white-tailed eagle pre-breeding period with increased flight activity. This may, in part, explain increased collision risk during spring. Possible displacement effects in white-tailed eagles may be avoided by siting wind-power plants farther inland or offshore, avoidance, collision risk, displacement, habitat use, habitat utilization, Haliaeetus albicilla, Norway,wind turbines

Evidence is increasing of bird mortality due to large-scale wind-energy development. Soaring raptors, such as the white-tailed eagle (Haliaeetus albicilla), have proven particularly vulnerable to collisions. In this study, we compared white-tailed eagle flight behavior both inside and outside of the Smola wind-power plant on coastal Central Norway. During the eagle breeding period (mid-Mar–end May 2008), we collected data on flight activity (directional flight, social behavior, and soaring) and flight altitude (below, within, and above the rotor-swept zone [RSZ]) at 12 vantage points; 6 within the wind-power plant and 6 outside (control area). We found that white-tailed eagles did not show any clear avoidance flight responses to the wind turbines. Hence, we found no significant differences in the total amount of flight activity within and outside the power-plant area. However, we found less flight activity among adults than among subadults within the power plant compared with the control area. We also found a slightly increased probability of flight activity in the RSZ within the power plant, which obviously may increase the risk of collision with wind turbines. Our findings may help explain the relatively high mortality rate of white-tailed eagles in the wind-power-plant area and the recorded peak in eagle fatalities during the breeding season. © 2013 The Wildlife Society.

Seabird movements outside the breeding season are generally poorly known, but can cover thousands of square km and a multitude of habitats, feeding conditions and potential threats. During the last decades, many seabird species in the North Atlantic have experienced large reductions in population size and breeding success, probably caused by reduced prey abundance caused by climate alterations and overfishing. One of these seabird species is the common guillemot. We used global location sensors (geolocators) to identify inter-breeding movements of 10 individuals breeding at Sklinna, a colony off the coast of Central Norway during July 2009–July 2010. All individuals moved northwards after breeding, and eight of them (80 %) entered the Barents Sea where they probably completed their moult. Three individuals moved southwards before the winter, but in total, half of the individuals stayed in the Barents Sea during winter. The other half wintered off the coast of Central Norway–Lofoten. The fact that all individuals moved northwards to winter was surprising as ringing recoveries suggest they also moves southwards (to the Skagerrak area) to winter. This suggests variation (individual or annual) in wintering movements and calls for a multi-year geolocator study at a number of colonies. Much of the area in the Barents Sea–Lofoten area is classified as vulnerable with respect to specific environmental pressures such as oil pollution and other anthropogenic factors, and the importance of the Barents Sea as a major wintering area for common guillemots from central Norway certainly has implications for the management authorities.

Successful wolverine (Gulo gulo) reproduction, and thereby population viability especially in multiple-use landscapes, is likely to be enhanced by availability of suitable den sites. We investigated which topographic features were selected for reproductive den sites by female wolverines in southern Norway (2000–2006) at three spatial scales (landscape, home-range and site-specific scales). At the sitespecific scale, den sites were associated with steep, rugged terrain with bare rock. At the home-range and landscape scales, den sites were placed in rugged terrain at 1100 m a.s.l. and away from infrastructure (private roads and public roads). These features provide snowdrifts into which wolverines can excavate dry, safe cavities. Den sites were also placed away from infrastructure, indicating that den-site distribution, and possibly successful reproduction, may be partly influenced by human activities. Recurrent use of specific topographic features may provide valuable information for guiding geographically differentiated management and monitoring efforts, and augmenting recovery of endangered wolverine populations.

In predator-prey theory, habitat heterogeneity can affect the relationship between kill rates and prey or predator density through its effect on the predator's ability to search for, encounter, kill and consume its prey. Many studies of predator- prey interactions include the effect of spatial heterogeneity, but these are mostly based on species with restricted mobility or conducted in experimental settings. 2. Here, we aim to identify the patterns through which spatial heterogeneity affects predator-prey dynamics and to review the literature on the effect of spatial hetero- geneity on predator-prey interactions in terrestrial mammalian systems, i.e. in freely moving species with high mobility, in non-experimental settings. We also review current methodologies that allow the study of the predation process within a spatial context. 3. When the functional response includes the effect of spatial heterogeneity, it usually takes the form of predator-dependent or ratio-dependent models and has wide applicability. 4. The analysis of the predation process through its different stages may further contribute towards identifying the spatial scale of interest and the specific spatial mechanism affecting predator-prey interactions. 5. Analyzing the predation process based on the functional response theory, but separating the stages of predation and applying a multiscale approach, is likely to

Conservation of carnivores in an increasingly changing environment is greatly helped by understanding the decision-making processes underlying habitat patch choice. Foraging theory may give us insight into spatio-temporal search patterns and consequent foraging decisions that carnivores make in heterogeneous and fluctuating environments. Constraints placed on central-place foragers in particular are likely to influence both foraging decisions and related spatio-temporal movement patterns. We used discrete choice models to investigate the spatio-temporal ranging behaviour of GPS collared female wolverines (Gulo gulo) with dependent cubs in south-central Norway. Activity patterns, home range use and selection for elevation were analyzed in relation to spatial and temporal covariates (daily and seasonal) and related to different foraging behaviours. In spring, wolverines showed restricted movement patterns around rendezvous sites at high elevations by day, whereas during the night animals were active at lower elevations. Over the summer, this daily pattern in intensity of use diminished and their overall selectiveness for elevation decreased as cubs grow more mobile and independent. At the onset of autumn, wolverines showed intensive use of the profitable forest-alpine tundra ecotone. We argue that reproducing wolverines deployed a foraging strategy attuned to altering their movement patterns throughout the summer to address a continuous, but diminishing, trade-off between providing both food and shelter for their offspring. Incorporating spatially and temporally explicit activity patterns and home range use in discrete choice resource selection models thus enhances the understanding of the motives behind wolverine resource utilization in space and time. Such knowledge may provide guidance to managers designing regional-scale zoning, in order to facilitate carnivore recovery and to minimize conflicts with human activities.

We propose a fundamental geographic distribution for the wolverine ( Gulo gulo (L., 1758)) based on the hypothesis that the occurrence of wolverines is constrained by their obligate association with persistent spring snow cover for successful reproductive denning and by an upper limit of thermoneutrality. To investigate this hypothesis, we developed a composite of MODIS classified satellite images representing persistent snow cover from 24 April to 15 May, which encompasses the end of the wolverine’s reproductive denning period. To investigate the wolverine’s spatial relationship with average maximum August temperatures, we used interpolated temperature maps. We then compared and correlated these climatic factors with spatially referenced data on wolverine den sites and telemetry locations from North America and Fennoscandia, and our contemporary understanding of the wolverine’s circumboreal range. All 562 reproductive dens from Fennoscandia and North America occurred at sites with persistent spring snow cover. Ninety-five percent of summer and 86% of winter telemetry locations were concordant with spring snow coverage. Average maximum August temperature was a less effective predictor of wolverine presence, although wolverines preferred summer temperatures lower than those available. Reductions in spring snow cover associated with climatic warming will likely reduce the extent of wolverine habitat, with an associated loss of connectivity.

Summary 1. The re-establishment of large carnivores in Norway has led to increased conflicts and the adoption of regional zoning for these predators. When planning the future distribution of large carnivores, it is important to consider details of their potential habitat tolerances and strength of interspecific differentiation. We studied differentiation in habitat and kill sites within the large-carnivore community of south-eastern Norway. 2. We compared habitat selection of the brown bear Ursus arctos L., Eurasian lynx Lynx lynx L., wolf Canis lupus L. and wolverine Gulo gulo L., based on radio-tracking data. Differences in kill site locations were explored using locations of documented predator-killed sheep Ovis aries L. We modelled each species’ selection for, and differentiation in, habitat and kill sites on a landscape scale using resource selection functions and multinomial logistic regression. Based on projected probability of occurrence maps, we estimated continuous patches of habitat within the study area. 3. Although bears, lynx, wolves and wolverines had overlapping distributions, we found a clear differentiation for all four species in both habitat and kill sites. The presence of bears, wolves and lynx was generally associated with rugged, forested areas at lower elevations, whereas wolverines selected rugged terrain at higher elevations. Some degree of sympatry was possible in over 40% of the study area, although only 1·5% could hold all four large carnivores together. 4. Synthesis and applications . A geographically differentiated management policy has been adopted in Norway, aimed at conserving viable populations of large carnivores while minimizing the potential for conflicts. Sympatry of all four carnivores will be most successful if regional zones are established of adequate size spanning an elevational gradient. High prey densities, low carnivore densities, low dietary overlap and scavenging opportunities have most probably led to reduced competitive exclusion. Although regional sympatry enhances the conservation of an intact guild of large carnivores, it may well increase conflict levels and resistance to carnivore conservation locally.

Within the predator guild, wolverines ( Gulo gulo (L., 1758)) have evolved as generalist predators and scavengers on prey killed by other predators. The recovery of wolves ( Canis lupus L., 1758) in the boreal forests of southern Norway during the late 1990s may have triggered consequent recolonization by wolverines through increased carcass availability. We investigated winter foraging behavior of wolverines in the boreal forest with regard to wolf, lynx ( Lynx lynx (L., 1758)), and red fox ( Vulpes vulpes (L., 1758)) presence. We followed 55 wolverine tracks in the snow from at least nine individuals for a total of 237 km during the winters of 2003–2004. We documented 19 moose ( Alces alces (L., 1758)) and 4 bird carcasses, and no successful hunts. Wolverines did not follow guild species trails directly to carcasses; however, they did change their movement patterns after red fox encounters. While wolverines were more active at higher elevations, the probability of encountering a wolf was higher at lower elevations, suggesting a spatial separation between wolverines and wolves. Although wolverines seem to depend on wolf for carrion during winter, they did not use wolf trails to find carcasses. This may indicate that wolverines reduce risk of intraguild predation by avoiding direct confrontation with wolves.

In Norway domestic sheep Ovis aries range unattended in mountainous areas during the summer season. Wolverines Gulo gulo re-established in the alpine regions of southern Norway during recent decades and are viewed as a substantial predator on lambs. Reducing predation on sheep by wolverines would not only reduce the economic loss to farmers but also promote the acceptance of wolverines in their summer ranges. We hypothesized that male lambs would be more prone to wolverine predation, because of higher locomotor activity, lower behavioural ewe–lamb synchrony and larger ewe–lamb distance. We studied ewe and lamb behavioural patterns, synchrony and ewe–lamb distance on a summer range in Knutshoi, south-central Norway. Although no differences were found in ewe–lamb distance or locomotor activity in gender, female lambs synchronized their behaviour more with their mother than males. Only for female lambs, increased synchrony resulted in closer ewe–lamb distances. Overall losses to wolverines based on long-term data indicate that male lambs are more prone to predation than females later in the season. These sex-specific behavioural differences in lambs affect the spatial and social relationships between ewe and lamb, and are likely to increase with age eventually leading to sexual segregation. Male lambs can therefore be expected to be more prone to wolverine predation towards the end of the season, when lambs become independent from the ewe.

A feeding trial was carried out on two captive wolverines Gulo gulo to evaluate methods to assess wolverine diets through scat content. During the feeding trial, wolverines were offered known quantities of five prey species. All scats were collected and their contents analysed. We evaluated four widely used methods of quantifying dietary composition: dry weight, index of relative contribution, frequency of occurrence, and percentage of occurrence. Based on the outcome of this evaluation, percentage of occurrence was found to be the most appropriate method for wolverine diet studies given the extreme variation in prey items (e.g. prey type and age) and undigested items (e.g. hide and bones) in the wolverine's diet. Dry weight may provide additional information on the amount of biomass consumed, which is biologically more meaningful than just the composition assessment derived from using the percentage of occurrence.

Compared to the other northern large carnivores, wolverines Gulo gulo are thought to be the most sensitive species with regard to habitat changes and human disturbance. Nowadays wolverines in Scandinavia are found in remote high alpine areas, and we investigated whether human development through presence of infrastructure has relegated them to these areas. We analysed wolverine habitat selection and the impact of infrastructure in two study areas in Norway using compositional analysis. We found that alpine tundra with low human development was important for wolverines to locate their home ranges. Human development formed a more important factor for home range location than did habitat, because habitat selectivity was much higher in undeveloped habitats than in developed habitats. Within their home ranges, wolverines used alpine shrubland and forest, irrespective of human development. The sympatric distribution of wolverines with wild and semi-domestic reindeer Rangifer tarandus indicates that wol...

Summary 1. Decreasing rate of migration in several species as a consequence of climate change and anthropic pressure, together with increasing evidence of space-use strategies intermediate between residency and complete migration, are very strong motivations to evaluate migration occurrence and features in animal populations. 2. The main goal of this paper was to perform a relative comparison between methods for identifying and charact erizing migration at the individual and population level on the basis of animal location data. 3. We classified 104 yearly individual trajectories from five populations of three deer species as migratory or non-migratory, by means of three methods: seasonal home range overlap, spatio-temporal separation of seasonal clusters and the Net Squared Displacement (NSD) method. For migr atory cases, we also measured timing and distance of migration and resi- dence time on the summer range. Finally, we comp ared the classification in migration cases across methods and populations. 4. All methods consistently identified migration at the population level, that is, they coherently dis- tinguished between complete or almost complete migr atory populations and partially migratory populations. Ho wever, in the latter case, methods co heren tly classified only about 50% of the sin- gle cases, that is they classified differently at the individual-animal level. We therefore infer that the compariso n of methods may help point to ‘less-stereo typed ’ cases in the residency -to-migration continuum. For ca ses consistently classified by all methods, no signifi cant differences were found in migration distance, or residence time on summer ranges. Timing of migration estimated by NSD was ea rlier than by the other two methods, both for spring and autumn migrations. 5. We suggest three steps to identify improper inferences from migration data and to enhance understanding of intermedia te space-use strategies. We recommend (i) classifying migration behaviours using more than one method, (ii) performing sensitivity analysis on method parame- ters to identify the extent of the differences and (iii) investigating inconsistently classified cases as these may often be ecologically interest ing (i.e. less-stereotyped migratory behaviours). adehabitat, hom e range overlap, movement patterns, Net Squared Displacement, red deer, reindeer, residence behaviour, roe deer, spatial clusters

1. Wolves Canis lupus L. recolonized the boreal forests in the southern part of the Scandinavian peninsula during the late 1990s, but so far there has been little attention to its effect on ecosystem functioning. Wolf predation increases the availability of carcasses of large prey, especially moose Alces alces L., which may lead in turn to a diet switch in facultative scavengers such as the wolverine Gulo gulo L. 2. Using 459 wolverine scats collected during winter-spring 2001-04 for DNA identity and dietary contents, we compared diet inside and outside wolf territories while controlling for potential confounding factors, such as prey density. We tested the hypothesis that wolverine diet shifted towards moose in the presence of wolves, while taking into account possible sexual segregation between the sexes. Occurrence of reindeer, moose and small prey was modelled against explanatory covariates using logistic mixed-effects models. Furthermore, we compared diet composition and breadth among habitats and sexes. 3. Occurrence of reindeer, moose and small prey in the diet varied with prey availability and habitat. As expected, diet contained more moose and less reindeer and small prey in the presence of wolves. Their diet in tundra consisted of 40% reindeer Rangifer tarandus L., 39% moose and 9% rodents. In forest with wolf, their diet shifted to 76% moose, 18% reindeer and 5% rodents; compared to 42% moose, 32% reindeer and 15% rodents in forest without wolf. This diet switch could not be explained by higher moose density in wolf territories. Female diet consisted of more small prey than for males, but there was a tendency for females to use the highly available moose carrion opportunistically and to hunt less on small prey within wolf territories. 4. Our study highlights how wolves increase scavenging opportunities for wolverines, and how sexual differences in diet may also apply to large scavengers. Due to their more restricted home range, female wolverines are forced to rely more on hunting small prey. The relatively high occurrence of wolf kills, however, forms an important food source to wolverines in this area. The recolonization of wolves may therefore have contributed to the consequent recolonization of wolverines into the same area.