Recolonization following past persecution questions the importance of persistent snow cover as a range limiting factor for wolverines.

Globally, climate is changing rapidly, which causes shifts in many species' distributions, stressing the need to understand their response to changing environmental conditions to inform conservation and management. Northern latitudes are expected to experience strongest changes in climate, with milder winters and decreasing snow cover. The wolverine (Gulo gulo) is a circumpolar, threatened carnivore distributed in northern tundra, boreal, and subboreal habitats. Previous studies have suggested that wolverine distribution and reproduction are constrained by a strong association with persistent spring snow cover. We assess this hypothesis by relating spatial distribution of 1589 reproductive events, a fitness-related proxy for female reproduction and survival, to snow cover over two decades. Wolverine distribution has increased and number of reproductive events increased 20 times in areas lacking spring snow cover during our study period, despite low monitoring effort where snow is sparse. Thus, the relationship between reproductive events and persistent spring snow cover weakened during this period. These findings show that wolverine reproductive success and hence distribution are less dependent on spring snow cover than expected. This has important implications for projections of future habitat availability, and thus distribution, of this threatened species. Our study also illustrates how past persecution, or other factors, that have restricted species distribution to remote areas can mask actual effects of environmental parameters, whose importance reveals when populations expand beyond previously restricted ranges. Overwhelming evidence shows that climate change is affecting many species and ecological processes, but forecasting potential consequences on a given species requires longitudinal data to revisit hypotheses and reassess the direction and magnitude of climate effects with new data. This is especially important for conservation-oriented management of species inhabiting dynamic systems where environmental factors and human activities interact, a common scenario for many species in different ecosystems around the globe.

Functional traits driving species role in the structure of terrestrial vertebrate scavenger networks.

Species assemblages often have a non-random nested organization, which in vertebrate scavenger (carrion-consuming) assemblages is thought to be driven by facilitation in competitive environments. However, not all scavenger species play the same role in maintaining assemblage structure, as some species are obligate scavengers (i.e., vultures) and others are facultative, scavenging opportunistically. We used a database with 177 vertebrate scavenger species from 53 assemblages in 22 countries across five continents to identify which functional traits of scavenger species are key to maintaining the scavenging network structure. We used network analyses to relate ten traits hypothesized to affect assemblage structure with the "role" of each species in the scavenging assemblage in which it appeared. We characterized the role of a species in terms of both the proportion of monitored carcasses on which that species scavenged, or scavenging breadth (i.e., the species "normalized degree"), and the role of that species in the nested structure of the assemblage (i.e., the species "paired nested degree"), therefore identifying possible facilitative interactions among species. We found that species with high olfactory acuity, social foragers, and obligate scavengers had the widest scavenging breadth. We also found that social foragers had a large paired nested degree in scavenger assemblages, probably because their presence is easier to detect by other species to signal carcass occurrence. Our study highlights differences in the functional roles of scavenger species and can be used to identify key species for targeted conservation to maintain the ecological function of scavenger assemblages.

Rubbing behavior of European brown bears: factors affecting rub tree selectivity and density.

Scent-mediated communication is considered the principal communication channel in many mammal species. Compared with visual and vocal communication, odors persist for a longer time, enabling individuals to interact without being in the same place at the same time. The brown bear (Ursus arctos), like other mammals, carries out chemical communication, for example, by means of scents deposited on marking (or rub) trees. In this study, we assessed rub tree selectivity of the brown bear in the predominantly deciduous forests of the Cantabrian Mountains (NW Spain). We first compared the characteristics of 101 brown bear rub trees with 263 control trees. We then analyzed the potential factors affecting the density of rub trees along 35 survey routes along footpaths. We hypothesized that: (1) bears would select particular trees, or tree species, with characteristics that make them more conspicuous; and (2) that bears would select trees located in areas with the highest presence of conspecifics, depending on the population density or the position of the trees within the species' range. We used linear models and generalized additive models to test these hypotheses. Our results showed that brown bears generally selected more conspicuous trees with a preference for birches (Betula spp.). This choice may facilitate the marking and/or detection of chemical signals and, therefore, the effectiveness of intraspecific communication. Conversely, the abundance of rub trees along footpaths did not seem to depend on the density of bear observations or their relative position within the population center or its border. Our results suggest that Cantabrian brown bears select trees based on their individual characteristics and their location, with no influence of characteristics of the bear population itself. Our findings can be used to locate target trees that could help in population monitoring.

La comunicación olfativa se considera el principal canal de comunicación en muchas especies de mamíferos. En comparación con la comunicación visual y la vocal, los olores persisten durante más tiempo, lo que permite a los individuos interactuar sin estar en el mismo lugar al mismo tiempo. El oso pardo (Ursus arctos), al igual que otros mamíferos, emplea la comunicación química, por ejemplo, por medio de olores depositados en árboles a través del marcaje o rascado. En este estudio, evaluamos la selección de árboles de marcaje por el oso pardo en los bosques predominantemente caducifolios de la Cordillera Cantábrica (noroeste de España). En primer lugar, comparamos las características individuales de 101 árboles de marcaje de oso pardo con 263 árboles control. Después, analizamos los factores potenciales que afectan la densidad de árboles de marcaje en 35 trayectos de prospección a lo largo de caminos y pistas forestales. Planteamos las hipótesis que: (1) los osos seleccionan árboles particulares, o especies de árboles, con características que los hacen más conspicuos; y (2) que los osos seleccionan árboles ubicados en áreas con mayor presencia de conespecíficos, dependiendo de la densidad de población osera o de la posición de los árboles dentro del rango de distribución de la especie. Usamos modelos lineales y modelos aditivos generalizados para probar estas hipótesis. Nuestros resultados mostraron que los osos pardos generalmente seleccionaron árboles más conspicuos, con preferencia por los abedules (Betula spp.). Esta elección puede facilitar el marcaje y/o detección de señales químicas y, por tanto, la eficacia de la comunicación intraespecífica. Por el contrario, la abundancia de marcaje a lo largo de los trayectos no parece depender de la densidad de las observaciones de osos o de su posición relativa con respecto al centro o los límites del rango de la población. Nuestros resultados sugieren que los osos pardos cantábricos seleccionan árboles en función de sus características individuales y de su ubicación, sin que influyan en ello las características de la población osera. Nuestros hallazgos pueden servir para localizar árboles específicos que podrían ayudar al monitoreo de la población.

Visual marking in mammals first proved by manipulations of brown bear tree debarking.

The rather limited human ability to understand animal vision and visual signalling has frequently clouded our expectations concerning the visual abilities of other animals. But there are multiple reasons to suspect that visual signalling is more widely employed by animals than previously thought. Because visibility of visual marks depends on the background in which they are seen, species spending most of their time living in dark conditions (e.g., in forests and/or having crepuscular and nocturnal habits) may rely on bright signals to enhance visual display. Here, as a result of experimental manipulations, we present, for the first time ever, evidence supporting the use of a new channel of intraspecific communication by a mammal species, i.e., brown bear Ursus arctos adult males relying on visual marks during mating. Bear reactions to our manipulation suggest that visual signalling could represent a widely overlooked mechanism in mammal communication, which may be more broadly employed than was previously thought.

Estimating and forecasting spatial population dynamics of apex predators using transnational genetic monitoring.

The ongoing recovery of terrestrial large carnivores in North America and Europe is accompanied by intense controversy. On the one hand, reestablishment of large carnivores entails a recovery of their most important ecological role, predation. On the other hand, societies are struggling to relearn how to live with apex predators that kill livestock, compete for game species, and occasionally injure or kill people. Those responsible for managing these species and mitigating conflict often lack fundamental information due to a long-standing challenge in ecology: How do we draw robust population-level inferences for elusive animals spread over immense areas? Here we showcase the application of an effective tool for spatially explicit tracking and forecasting of wildlife population dynamics at scales that are relevant to management and conservation. We analyzed the world's largest dataset on carnivores comprising more than 35,000 noninvasively obtained DNA samples from over 6,000 individual brown bears (Ursus arctos), gray wolves (Canis lupus), and wolverines (Gulo gulo). Our analyses took into account that not all individuals are detected and, even if detected, their fates are not always known. We show unequivocal quantitative evidence of large carnivore recovery in northern Europe, juxtaposed with the finding that humans are the single-most important factor driving the dynamics of these apex predators. We present maps and forecasts of the spatiotemporal dynamics of large carnivore populations, transcending national boundaries and management regimes.

Wolf habitat selection when sympatric or allopatric with brown bears in Scandinavia.

Habitat selection of animals depends on factors such as food availability, landscape features, and intra- and interspecific interactions. Individuals can show several behavioral responses to reduce competition for habitat, yet the mechanisms that drive them are poorly understood. This is particularly true for large carnivores, whose fine-scale monitoring is logistically complex and expensive. In Scandinavia, the home-range establishment and kill rates of gray wolves (Canis lupus) are affected by the coexistence with brown bears (Ursus arctos). Here, we applied resource selection functions and a multivariate approach to compare wolf habitat selection within home ranges of wolves that were either sympatric or allopatric with bears. Wolves selected for lower altitudes in winter, particularly in the area where bears and wolves are sympatric, where altitude is generally higher than where they are allopatric. Wolves may follow the winter migration of their staple prey, moose (Alces alces), to lower altitudes. Otherwise, we did not find any effect of bear presence on wolf habitat selection, in contrast with our previous studies. Our new results indicate that the manifestation of a specific driver of habitat selection, namely interspecific competition, can vary at different spatial-temporal scales. This is important to understand the structure of ecological communities and the varying mechanisms underlying interspecific interactions.

Methodological Considerations for Using Fecal Glucocorticoid Metabolite Concentrations as an Indicator of Physiological Stress in the Brown Bear (Ursus arctos).

Reliable methods to measure stress-related glucocorticoid responses in free-ranging animals are important for wildlife management and conservation. Such methods are also paramount for our ability to improve our knowledge of the ecological consequences of physiological processes. The brown bear (Ursus arctos) is a large carnivore of ecological and cultural importance and is important for management. Here, we provide a physiological validation for an enzyme immunoassay (EIA) to quantify glucocorticoid metabolites in brown bear feces. We also provide an evaluation of the effects of sample exposure to ambient temperature on measured fecal glucocorticoid metabolite (fGCM) concentrations. We evaluated three EIA systems: a cortisol assay, an 11-oxoetiocholanolone assay, and an 11ß-hydroxyetiocholanolone assay. Of these, the cortisol assay provided the best discrimination between peak fGCM concentrations detected 1-4 d after injections of synthetic adrenocorticotrophic hormone and preinjection baseline concentrations in four individual brown bears. The time of exposure to ambient temperature had substantial but variable effects on measured fGCM concentrations, including variation both between samples from the same individual and among samples from different bears. We propose that the validated EIA system for measuring fGCM concentrations in the brown bear could be a useful noninvasive method to monitor stress in this species. However, we highlight that this method requires that fecal samples be frozen immediately after defecation, which could be a limitation in many field situations.

High frequency GPS bursts and path-level analysis reveal linear feature tracking by red foxes.

There is a need to quantify and better understand how wildlife interact with linear features, as these are integral elements of most landscapes. One potentially important aspect is linear feature tracking (LFT), yet studies rarely succeed in directly revealing or quantifying this behavior. In a proof-of-concept study, we employed short-term intensive GPS monitoring of red foxes (Vulpes vulpes) in a multiple-use landscape in southern Norway. Using periodic bursts of high frequency GPS position fixes, we performed modified path selection analyses to estimate the propensity of foxes to track natural and man-made linear features (roads, forest edges, and streams) once they are encountered. Foxes in our study tracked primarily forest edges and roads. Forty-three percent of bursts that encountered any linear feature resulted in LFT. LFT, although prominent, was manifested as a short-lived behavior, with overall median times to linear feature abandonment around two minutes. Movement speeds were highest along roads, perhaps due to greater ease of travel or higher perceived risk. In the highly heterogeneous habitats that characterize human-dominated landscapes, LFT may be manifested at such a fine spatio-temporal scale that it would remain hidden during telemetry studies employing conventional position fix frequencies. The approach described here may aid others studying spatial behaviors that are manifested over very short durations, yet are biologically significant.

Scavenging in the Anthropocene: Human impact drives vertebrate scavenger species richness at a global scale.

Understanding the distribution of biodiversity across the Earth is one of the most challenging questions in biology. Much research has been directed at explaining the species latitudinal pattern showing that communities are richer in tropical areas; however, despite decades of research, a general consensus has not yet emerged. In addition, global biodiversity patterns are being rapidly altered by human activities. Here, we aim to describe large-scale patterns of species richness and diversity in terrestrial vertebrate scavenger (carrion-consuming) assemblages, which provide key ecosystem functions and services. We used a worldwide dataset comprising 43 sites, where vertebrate scavenger assemblages were identified using 2,485 carcasses monitored between 1991 and 2018. First, we evaluated how scavenger richness (number of species) and diversity (Shannon diversity index) varied among seasons (cold vs. warm, wet vs. dry). Then, we studied the potential effects of human impact and a set of macroecological variables related to climatic conditions on the scavenger assemblages. Vertebrate scavenger richness ranged from species-poor to species rich assemblages (4-30 species). Both scavenger richness and diversity also showed some seasonal variation. However, in general, climatic variables did not drive latitudinal patterns, as scavenger richness and diversity were not affected by temperature or rainfall. Rainfall seasonality slightly increased the number of species in the community, but its effect was weak. Instead, the human impact index included in our study was the main predictor of scavenger richness. Scavenger assemblages in highly human-impacted areas sustained the smallest number of scavenger species, suggesting human activity may be overriding other macroecological processes in shaping scavenger communities. Our results highlight the effect of human impact at a global scale. As species-rich assemblages tend to be more functional, we warn about possible reductions in ecosystem functions and the services provided by scavengers in human-dominated landscapes in the Anthropocene.

Testing the influence of habitat experienced during the natal phase on habitat selection later in life in Scandinavian wolves.

Natal habitat preference induction (NHPI) occurs when characteristics of the natal habitat influence the future habitat selection of an animal. However, the influence of NHPI after the dispersal phase has received remarkably little attention. We tested whether exposure to humans in the natal habitat helps understand why some adult wolves Canis lupus may approach human settlements more than other conspecifics, a question of both ecological and management interest. We quantified habitat selection patterns within home ranges using resource selection functions and GPS data from 21 wolf pairs in Scandinavia. We identified the natal territory of each wolf with genetic parental assignment, and we used human-related characteristics within the natal territory to estimate the degree of anthropogenic influence in the early life of each wolf. When the female of the adult wolf pair was born in an area with a high degree of anthropogenic influence, the wolf pair tended to select areas further away from humans, compared to wolf pairs from natal territories with a low degree of anthropogenic influence. Yet the pattern was statistically weak, we suggest that our methodological approach can be useful in other systems to better understand NHPI and to inform management  about human-wildlife interactions.

Patterns of brown bear damages on apiaries and management recommendations in the Cantabrian Mountains, Spain.

Large carnivores are often persecuted due to conflict with human activities, making their conservation in human-modified landscapes very challenging. Conflict-related scenarios are increasing worldwide, due to the expansion of human activities or to the recovery of carnivore populations. In general, brown bears Ursus arctos avoid humans and their settlements, but they may use some areas close to people or human infrastructures. Bear damages in human-modified landscapes may be related to the availability of food resources of human origin, such as beehives. However, the association of damage events with factors that may predispose bears to cause damages has rarely been investigated. We investigated bear damages to apiaries in the Cantabrian Mountains (Spain), an area with relatively high density of bears. We included spatial, temporal and environmental factors and damage prevention measures in our analyses, as factors that may influence the occurrence and intensity of damages. In 2006-2008, we located 61 apiaries, which included 435 beehives damaged in the study area (346 km2). The probability of an apiary being attacked was positively related to both the intensity of the damage suffered the year before and the distance to the nearest damaged apiary, and negatively related to the number of prevention measures employed as well as the intensity of the damage suffered by the nearest damage apiary. The intensity of damage to apiaries was positively related to the size of the apiary and to vegetation cover in the surroundings, and negatively related to the number of human settlements. Minimizing the occurrence of bear damages to apiaries seems feasible by applying and maintaining proper prevention measures, especially before an attack occurs and selecting appropriate locations for beehives (e.g. away from forest areas). This applies to areas currently occupied by bears, and to neighbouring areas where dispersing individuals may expand their range.

Brown bear (Ursus arctos) attacks resulting in human casualties in Scandinavia 1977-2016; management implications and recommendations.

Human persecution and habitat loss have endangered large carnivore populations worldwide, but some are recovering, exacerbating old conflicts. Carnivores can injure and kill people; the most dramatic form of wildlife-human conflict. In Scandinavia, the brown bear (Ursus arctos) population increased from ~500 bears in 1977 to ~3300 in 2008, with an increase in injuries, fatalities, and public fear of bear attacks. We reviewed media coverage and interviewed victims to explore how bear population trends, hunter education, and other factors may have influenced the number of injuries and fatalities in Scandinavia from 1977 to 2016. We found 42 incidents with 42 injuries and 2 fatalities; 42 were adult men, one was an adult woman conducting forestry work, and one was a boy skiing off-piste. Thirty-three adult men were hunting bears, moose, or small game, often with a hunting dog, and 26 had shot at the bear at 8±11 m before injury. Eleven nonhunters were conducting forestry work, inspecting a hunting area, picking berries, tending livestock, hiking, harassing a denned bear, and one person was killed outside his house at night. Eight of the 11 incidents of nonhunters involved female bears with cubs; three of these family groups were in dens and two were on carcasses. The annual number of hunters injured/killed was mostly influenced by the increase in the bear population size. The pattern was similar regarding injuries/fatalities to other outdoor users, but the relation with the bear population size was weaker than for hunters, and the null model was equally supported. Bear physiology at denning may make encounters with bears more risky in the fall, when bears show prehibernation behavior. Awareness and education efforts, especially among hunters, seem important to ensure human safety. Recreationists and forestry workers should avoid dense vegetation or make noise to warn bears of their presence.

Habitat segregation between brown bears and gray wolves in a human-dominated landscape.

Identifying how sympatric species belonging to the same guild coexist is a major question of community ecology and conservation. Habitat segregation between two species might help reduce the effects of interspecific competition and apex predators are of special interest in this context, because their interactions can have consequences for lower trophic levels. However, habitat segregation between sympatric large carnivores has seldom been studied. Based on monitoring of 53 brown bears (Ursus arctos) and seven sympatric adult gray wolves (Canis lupus) equipped with GPS collars in Sweden, we analyzed the degree of interspecific segregation in habitat selection within their home ranges in both late winter and spring, when their diets overlap the most. We used the K-select method, a multivariate approach that relies on the concept of ecological niche, and randomization methods to quantify habitat segregation between bears and wolves. Habitat segregation between bears and wolves was greater than expected by chance. Wolves tended to select for moose occurrence, young forests, and rugged terrain more than bears, which likely reflects the different requirements of an omnivore (bear) and an obligate carnivore (wolf). However, both species generally avoided human-related habitats during daytime. Disentangling the mechanisms that can drive interspecific interactions at different spatial scales is essential for understanding how sympatric large carnivores occur and coexist in human-dominated landscapes, and how coexistence may affect lower trophic levels. The individual variation in habitat selection detected in our study may be a relevant mechanism to overcome intraguild competition and facilitate coexistence.

No place like home? A test of the natal habitat-biased dispersal hypothesis in Scandinavian wolves.

Natal dispersal is an important mechanism for the viability of populations. The influence of local conditions or experience gained in the natal habitat could improve fitness if dispersing individuals settle in an area with similar habitat characteristics. This process, defined as 'natal habitat-biased dispersal' (NHBD), has been used to explain distribution patterns in large carnivores, but actual studies evaluating it are rare. We tested whether grey wolf Canis lupus territory establishment was influenced by the habitat characteristics of the natal territory using the long-term monitoring of the Scandinavian wolf population. We paired the locations of natal and established territories, accounted for available habitats along the dispersing route, and compared their habitat characteristics for 271 wolves during 1998-2012. Wolves with the shortest dispersal distances established in natal-like habitat types more than expected by chance, whereas wolves that dispersed longer distances did not show NHBD. The pattern was consistent for male and female wolves, with females showing more NHBD than males. Chances to detect NHBD increased with the size of habitat defined as available. This highlights the importance of considering the biological characteristics of the studied species when defining habitat availability. Our methodological approach can prove useful to inform conservation and management to identify habitats to be selected by reintroduced or naturally expanding populations.

Staying cool or staying safe in a human-dominated landscape: which is more relevant for brown bears?

Pigeon et al. (2016) Staying cool in a changing landscape: the influence of maximum daily ambient temperature on grizzly bear habitat selection. Oecologia 181:1101. doi: 10.1007/s00442-016-3630-5 analyzed the effect of ambient temperature on the habitat selection of grizzly bears (Ursus arctos) in Alberta, Canada. They concluded that temperature played a significant role in bear habitat selection and that it was unlikely that human activity introduced biases to the habitat selection of bears. However, Pigeon et al. did not consider variables related to human activities in their analyses. They also misinterpreted previous research that has accounted for temperature in the habitat selection of brown bears. There is much literature published on the negative effects of human disturbance on wildlife in general and on bears in particular. Downplaying the role of human disturbance could have important negative consequences if, in fact, human disturbance were a more important factor than thermoregulation. Indeed, dismissing the importance of human influence, in the face of contradictory evidence, could tempt managers to disregard an important factor that is difficult and often unpopular to deal with in their conservation plans.

Competition between apex predators? Brown bears decrease wolf kill rate on two continents.

Trophic interactions are a fundamental topic in ecology, but we know little about how competition between apex predators affects predation, the mechanism driving top-down forcing in ecosystems. We used long-term datasets from Scandinavia (Europe) and Yellowstone National Park (North America) to evaluate how grey wolf (Canis lupus) kill rate was affected by a sympatric apex predator, the brown bear (Ursus arctos). We used kill interval (i.e. the number of days between consecutive ungulate kills) as a proxy of kill rate. Although brown bears can monopolize wolf kills, we found no support in either study system for the common assumption that they cause wolves to kill more often. On the contrary, our results showed the opposite effect. In Scandinavia, wolf packs sympatric with brown bears killed less often than allopatric packs during both spring (after bear den emergence) and summer. Similarly, the presence of bears at wolf-killed ungulates was associated with wolves killing less often during summer in Yellowstone. The consistency in results between the two systems suggests that brown bear presence actually reduces wolf kill rate. Our results suggest that the influence of predation on lower trophic levels may depend on the composition of predator communities.

Conflict Misleads Large Carnivore Management and Conservation: Brown Bears and Wolves in Spain.

Large carnivores inhabiting human-dominated landscapes often interact with people and their properties, leading to conflict scenarios that can mislead carnivore management and, ultimately, jeopardize conservation. In northwest Spain, brown bears Ursus arctos are strictly protected, whereas sympatric wolves Canis lupus are subject to lethal control. We explored ecological, economic and societal components of conflict scenarios involving large carnivores and damages to human properties. We analyzed the relation between complaints of depredations by bears and wolves on beehives and livestock, respectively, and bear and wolf abundance, livestock heads, number of culled wolves, amount of paid compensations, and media coverage. We also evaluated the efficiency of wolf culling to reduce depredations on livestock. Bear damages to beehives correlated positively to the number of female bears with cubs of the year. Complaints of wolf predation on livestock were unrelated to livestock numbers; instead, they correlated positively to the number of wild ungulates harvested during the previous season, the number of wolf packs, and to wolves culled during the previous season. Compensations for wolf complaints were fivefold higher than for bears, but media coverage of wolf damages was thirtyfold higher. Media coverage of wolf damages was unrelated to the actual costs of wolf damages, but the amount of news correlated positively to wolf culling. However, wolf culling was followed by an increase in compensated damages. Our results show that culling of the wolf population failed in its goal of reducing damages, and suggest that management decisions are at least partly mediated by press coverage. We suggest that our results provide insight to similar scenarios, where several species of large carnivores share the landscape with humans, and management may be reactive to perceived conflicts.

Physiological evidence for a human-induced landscape of fear in brown bears (Ursus arctos).

Human persecution is a major cause of mortality for large carnivores. Consequently, large carnivores avoid humans, but may use human-dominated landscapes by being nocturnal and elusive. Behavioral studies indicate that certain ecological systems are "landscapes of fear", driven by antipredator behavior. Because behavior and physiology are closely interrelated, physiological assessments may provide insight into the behavioral response of large carnivores to human activity. To elucidate changes in brown bears' (Ursus arctos) behavior associated with human activity, we evaluated stress as changes in heart rate (HR) and heart rate variability (HRV) in 12 GPS-collared, free-ranging bears, 7 males and 5 females, 3-11 years old, using cardiac-monitoring devices. We applied generalized linear regression models with HR and HRV as response variables and chest activity, time of day, season, distance traveled, and distance to human settlements from GPS positions recorded every 30 min as potential explanatory variables. Bears exhibited lower HRV, an indication of stress, when they were close to human settlements and especially during the berry season, when humans were more often in the forest, picking berries and hunting. Our findings provide evidence of a human-induced landscape of fear in this hunted population of brown bears.

Behavioural differences between single scandinavian brown bears (Ursus arctos) and females with dependent young when experimentally approached by humans.

Carnivore-human encounters that result in human injury present a conservation and management challenge and it is therefore important to understand under what conditions such incidents occur. Females with cubs are often involved when humans are injured by brown bears Ursus arctos. In Scandinavia, this is particularly true for unarmed recreational forest users. Our aim was to document behavioural differences between single bears and females with cubs in order to develop recommendations to minimize the risk of injuries to recreational forest users. We documented the reactions of GPS-collared females with cubs and single brown bears to experimental approaches by humans to 50 m from the bear on 42 and 108 occasions, respectively. The majority of females with cubs (95%) and single bears (89%) left when approached. Bears that left were passed at shorter distances and were in more open areas than those that stayed. Both groups had similar flight initiation distances, which were longer for bears that were active at the time of the disturbance. Females with cubs selected more open habitat than single bears, also for the new site they selected following disturbance. Females with cubs, particularly active females with cubs of the year, moved greater distances and spent more time active following the approach. Females with cubs and single bears were seen or heard in 26% and 14% of the approaches, respectively. None of the bears displayed any aggressive behaviour during the approaches. Females with cubs selected more open habitat, perhaps predisposing them to encountering people that are not involved in hunting activities, which might be the primary explanation why females with cubs are most frequently involved when unarmed people are injured by bears in Scandinavia. To mitigate injury risks, one must consider factors that bring bears closer to human activity in the first place.